THE NAUTILUS
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0314
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Volume 130, Number 3
September 2, 2016
ISSN 0028-1344
A quarterly devoted
to malacology.
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The Bailey- Matthews National
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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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Museum National d’Histoire Naturelle
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THE
CONTENTS
Carole S. Hickman
Thomas J. DeVries
Kazutaka Amano
Anton Oleinik
Robert G. Jenkins
Martin Averv Snvder
Geerat J. Vermeij
Emilio F. Garcia
William G. Lyons
Martin Avery Snyder
Research Notes
Mary K. Wicksten
William G. Ly ons
Martin Avery Snyder
NAUTILUS
Volume 130, Number 3
September 2, 2016
ISSN 0028-1344
New species of deep-sea gastropods from tiie Indo-West Pacific Region
(Gastropoda: Vetigastropoda: Seguenzioidea: Calliotropidae) with a
geologic and biogeographic perspective . 83
Latest Oligocene and Miocene whelks (Gastropoda: Neogastropoda:
Buccinidae) from Peru . 101
A new species of Adrnete (Gastropoda: Cancellariidae: Admetinae)
from the Paleoeene of eastern Hokkaido, northern Japan . 116
Hesperaptyxis, a new genus for some western American Fasciolariidae
(Gastropoda), with the description ol a new species . 122
The western Caribbean complex of Fasciola ria tephrina de Souza, 2002
(Gastropoda: Fasciolariidae), with the description of a new species . 127
A tumbling snail (Gastropoda: Vetigastropoda: Margaritidae) . 132
The authorship of Turrilatirus craticulatus (Gastropoda:
Fasciolariidae: Peristerniinae) . 134
Sponsored in part by the State of
Flonda, Department of State,
Division of Cultural Affairs and the
Florida Council on Arts and Culture
r ^CULTURE
Us BUILDS
^FLORIDA
FLORIDA DEPARTMENT o/STATE
DIVISION of CULTURAL AFFAIRS
THE NAUTILUS 130(3):83-100, 2016
Page 83
New species of deep-sea gastropods from the Indo-West Pacific
Region (Gastropoda: Vetigastropoda: Seguenzioidea: Calliotropidae)
witli a geologic and biogeographic perspective
Carole S. Hickman
Department of Integrative Biolog)' and Museum of Paleontology
University of California
Berkeley, California 94720-3140 USA
[email protected]
ABSTRACT
This paper identifies the biogeographic realm of Wallacea as a
center of deep-sea (>200 m) endemism in the basal gastropod
family Calliotropidae Hickman and McLean, 1990. Six new
medium- to large-shelled species are described from Sulawesi:
Calliotropis hukabukaensis , C. tominiensis, and C. new species
from >1000 m in the Gulf of Tomini and C tabaknensis ,
C. locolocoensis , and C. lamuluensis from the > 988 m in the
Gulf of Bone. Two additional large-shelled species are described:
C. enantioserrata from >1000 m on the eastern Australian slope
in the northern Tasman Sea and C. andamanensis from 500 m in
the Andaman Sea off the west coast of Thailand. Previous
descriptions and records of Calliotropis species in Indonesia are
from the margins of Wallacea — the Makassar Strait and Islands
in the Banda Arc, notably Lai. Tanimbar, and Timor. When
calliotropid biodiversity is connected with the large body of new
knowledge of the contorted seafloor relief of the Indonesian
region, it is clear that entire deep basins underlain by oceanic
crust have never been sampled. The widespread occurrence of
Cenozoic calliotropid gastropods in cheinosynthetically based
ecosystems is consistent with new data from petroleum explora¬
tion in the volcanic arcs of Indonesia documenting mud volca¬
noes and expulsion of geofluids (hydrocarbons, sulfides) that
typically support rich microbial communities in the deep sea.
The long and complex history of the collision of the Eurasian
and Australian plates, arc volcanisrn, subduction, fusion of rem¬
nant pieces of continental crust, and opening of new deep gulfs
affect the movement of shallow and deep currents and the
Indonesian Throughllow from the Pacific into the Indian Ocean.
This physical history generates testable hypotheses for reassessing
the phylogenetic history of ancient gastropod lineages that origi¬
nated in the Mesozoic Era or earlier.
Additional Keywords: Indonesia, Sulawesi, Wallacea, Indonesian
Throughflow, plate collision, subduction, cold seep, ehemosyn-
thesis, intritacalx
INTRODUCTION
The Indo-West Pacific (1WP) region is recognized as the
global center of marine biodiversity (Hoeksema, 2007;
Bellwood et ah, 2012). Efforts to understand patterns of
elevated diversity of the invertebrate biota have centered
on a triangular region (the Coral Triangle) identified in
the literature by more than 15 different descriptive terms
(see Hoeksema, 2007). This same region has long been
famous as a terrestrial biodiversity hotspot encompassing
the thousands of tropical islands of eastern Indonesia.
In the marine realm, modem research has concentrated
on characterizing and explaining tropical shallow marine
diversity. In gastropods, this has included the use of molec¬
ular tools to document patterns of speeiation and dispersal
and to investigate genetic connectivity and species bound¬
aries (Reid et ah, 2006; Williams et ah, 2011; Williams
et ah, 2012).
There is increasing evidence of a more localized deep¬
water hotspot for basal gastropod diversity in the region
of Wallacea, in marine basins underlain by oceanic crust
between the shallow continental shelves of the collid¬
ing Australian, Eurasian, and Philippine tectonic plates
(Hickman, 2009a, 2009b, 2009c, 2012). Anatomical and
molecular investigations of this putative deep hotspot are
impeded by lack of deep-water sampling and access to
fresh material. However, emerging geological, geophysi¬
cal and hydrographic understanding of remnant deep
basins and deep forearc settings provides an unparalleled
opportunity to study closely correlated patterns in marine
geology, geophysics, and biology.
The primary purpose of this paper is to describe new
species of narrowly endemic basal marine calliotropid
gastropods in the Indo-West Pacific Region. A preceding
paper (Hickman, 2012) described a new genus and two
new species in the trochoidean family Gazidae Hickman
and McLean, 1990, and will be followed by an account of
new species in the seguenzioidean family Cataegidae
McLean and Quinn, 1987.
Secondary objectives are to offer some new perspec¬
tives on persistent problems and challenges of calliotropid
systematic^ and to present new perspectives on the tec¬
tonics and complex geologic history of the assembly
of Wallacea and the specific deep tectono-sedimentarv
settings in which the new species were collected.
Page 84
THE NAUTILUS, Vol. 130, No. 3
There are >100 available names for putative living
species of Calliotropis Seguenza, 1903. Some species
are narrowly endemic, but others have unusually broad
ranges in which disjunct populations may be separated
by thousands of kilometers. There is no satisfactory basis
for morphological delimitation of subgenera, and a con¬
servative treatment of new species is adopted here pend¬
ing revisionary systematics integrating shell, anatomical,
and radular characters with molecular sequence data.
Most species currently are known only from shells, but
the amount of unstudied material in museum collections
is substantial. There are excellent species-level accounts
for the Indo-Pacifie (e.g.„ Vilvens, 2004; 2006; 2007) and
the Philippines (Poppe et ah, 2006) that include detailed
geographic and bathymetric range data. An important
compilation of data on the taxonomy and distribution
of 245 South African vetigastropod species (Herbert,
2015) lends weight to a previously subjective and per¬
plexing disjunction of taxa between the South African
and Indonesian regions. New data from Late Cenozoie
fossil vetigastropods in the Philippines (Helwerda et ah,
2014) add to a new baseline for revisionary work. Tl le
following brief account of the history of calliotropid
systematics provides a framework and rationale for the
descriptions of new species.
Nomenclature and Classification of
Living Calliotropids
Calliotropid gastropods are known primarily from explor¬
ing expeditions beginning in the late 19'1' and earlv 20'1'
centuries, notably the Challenger, Valdivia, Investica
tor, Travailleur, Talisman, Siboca, Princesse-Alige,
Lightning, Porcupine, Blake, and Albatross. Species
were described or treated under a variety of genus-group
names, including Trochus (Margarita) (e.g., Watson, 1879;
P. Fischer, 1882), Solariella (e.g., Wood-Mason and
Alcock, 1891; Smith, 1S94; Locard, 1897; Sowerby III,
1903); Basilissa (e.g., Dautzenberg and Fischer, 1897;
Martens and Thiele, 1904), and Solariclliopsis (Schepman,
1908). Names were based exclusively on shell features
until Schepman (1908) observed and figured both shell
and radula (reproduced here as Figures 1-2) noting “how
dangerous it is to classify such species without any knowl¬
edge of the soft parts.’
The nominotypieal genus Calliotropis Seguenza, 1903
is based on a Pliocene-Pleistocene fossil species from
Italy and is the name that has served for conservative
treatments of more than 100 species from later deep-sea
expeditions up until the present. Diagnosis of the family
Calliotropidae Hickman and McLean, 1990 included
features of the radula (Figures 3-4) that became clear
with Scanning Electron Microscopy. They include a
small-cusped “hooded” rachidian and large-cusped
hooded lateral teeth with a system of deep interlocking
basal projections and pockets, distinctive lateromarginal
plates, extremely fine and numerous marginal teeth and
peculiar mitten-shaped outermost marginal (Hickman
and McLean, 1990). An enlarged hindgut has been noted
in many species along with evidence for selective deposit
feeding (Hickman, 1981).
Other unique anatomical traits have been noted in
calliotropids. The first is a large, enrolled, cylindrical
elaboration of the right side of the male epipodium
that has been interpreted as an “intromittent organ”
(Dali, 1889) or “penis” or “copulatory organ” (Waren
and Bouehet, 1989; Kano, 2007), or more conservatively
as sperm delivery alternative to broadcast spawning that
may not involve internal fertilization (Hickman, 1992;
Kano, 2007). The second is the location of the “eyes” in
some species at the bases of eyestalks and their modifi¬
cation to large irregular patches of black pigment (Dali,
1889). Observation of asymmetry and variability of the
right and left pigmented regions has been interpreted
as “a beginning of the loss of vision in an environment
in which eyes are no longer of any use” (Waren and
Bouehet, 1989). The asymmetry of the pigmented regions
has been interpreted as evidence of separate genetic
control of development of the right and left eye (Waren
and Bouehet, 1989).
Twentieth century attempts to bring taxonomic order
to Calliotropis include Marshall (1979) and Quinn (1979,
1991), although species have continued to be treated
under inappropriate generic names (e.g., Abbott, 1974)
or in inappropriate families (e.g., Nordsieck, 1968, 1982;
Rubio and Rolan, 1987). A full review is beyond the
scope of this paper.
Most significantly, the late 20'1' and early 21st century
have seen a series of French deep-sea sampling expedi¬
tions recovering many calliotropids from bathyal depths
in Oceania and the Indo-West Pacific Region. This work
has further resulted in description of many new callio¬
tropid species based exclusively on shell features (Poppe
et ah, 2006) and Vilvens, 2004, 2005, 2007) as well as
new geographic and bathymetric records from the west¬
ern Indian Ocean (Vilvens, 2006) and central Eastern
Atlantic (Vilvens and Swinnen, 2008).
The monumental survey of Vilvens (2007) provides
251 figures of shells and classifies 94 species under
Calliotropis , while declining to describe new subgenera
or assign species to existing available genus-group names
in the absence of anatomical data. Species are divided
into 8 “formal groups” that are pragmatically justified
and helpful for comparing shells of the same size and
ratio of height to width. Descriptions and illustrations
provide considerable additional detail such as numbers
of cords on spire whorls and base, numbers of nodes or
beads on spiral cords, whorl shape, and observations of
ontogeny of these features.
Indonesia is the least well-sampled part of the Indo-
Pacifie Region. This is particularly unfortunate because
it is geologically and hydrographieally the most complex
region on Earth and includes the Indonesian Through-
flow (ITF) and the oceanic gateway connecting the
Pacific and Indian Oceans (Tillinger, 2011). It is for¬
tuitous that the Albatross Philippines Expedition of
1907-1910 included some exploratory deep dredging in
C.S. Hickman, 2016
Page 85
Figures 1-4. Calliotropid shell and radula. 1. Schepman’s original illustration ol the shell of Calliotropis pagodaformis .
2. Schepman’s original drawing of the radula of Calliotropis calcarata. 3. Scanning electron micrograph of the rachidian and lateral
teeth from a typical calliotropid radula, scale bar = 100 pm. 4. Detail of rachidian tooth cusp, shaft, hood and base, scale bar = 20 pm.
Indonesia (Sulawesi and Halmahera), and this paper
documents six new calliotropid species from that expe¬
dition that have remained undescribed for >100 years.
The paper also includes a new calliotropid species from
the slope off eastern Australia and one from the eastern
Andaman Sea.
Nomenclature and Classification of
Fossil Calliotropids
A full review of the treatment of the fossil record of this
group is not directly relevant to the description of new
living species. However, there is a long history of using
separate family-group names for Mesozoic and Cenozoic
lineages that differ little in shell morphology. Hickman
and McLean recognized Eucyelidae Koken, 1897 as an
extinct, primarily Mesozoic, family group with maximum
diversity in shallow seas of the Jurassic and introduced
the new family group name Calliotropidae (as Callio-
tropini) for the Cenozoic radiation into deep water of
taxa previously classified under a variety of inappropriate
trochoidean family-group names. During the last ten
years, paleontological treatments have introduced new
family-group names and reallocated both fossil and living
genera (e.g., Bandel, 2010), or advocated the use Callio¬
tropis sensu stricto for the Cenozoic forms and using
Riselloidea Cossmann, 1909 as a temporal subgenus for
the Mesozoic forms (Ferrari et ah, 1014). This may seem
to be a highly unsettled systematic state. However, recog¬
nition of the seguenzioid affinities of many Mesozoic and
Cenozoic taxa (Waren et ah, 2003; Bouehet and Rocroi,
2005) has revolutionized systematics. Equally revolution¬
ary is the increasing molecular support for a deep diver¬
gence and common ancestor of hypothesized seguenzioid
radiations (Kano, 2007; Kano et ah, 2009; Aktipis and
Giribet, 2012), including those into the deep sea. Callio¬
tropis is effectively a form genus at this stage, hut it is
poised for an interesting revision integrating many kinds
of data.
Page 86
THE NAUTILUS, Vol. 130, No. 3
Successful revision will require new anatomical and
molecular data and extensive taxon sampling. In terms
of shell morphology, it will require consistent definition
of characters in terms of clearly differentiated character
states. Size and shape terms, measurements, ratios,
angles, whorl counts, counts of sculptural elements are
useful for defining ranges of variation within species
represented by many specimens. However, their value
is descriptive rather than diagnostic. Several untapped
sources of morphological hypotheses of relationships are
identified in the species descriptions that follow. They
include the recognition of terminal growth in the form
of a thickened or reflected outer lip, a descending aper¬
ture, and modes of complete or partial closure of an
open umbilicus. A columellar denticle is a terminal
growth feature, if it is not a continuation of a columellar
plication or denticulation of the columella present in
earlier columellar ontogeny. Little attention has been
paid to shell microstructure and to modes of decreasing
shell thickness. Nacre that is incompletely obscured by
an unusually thin veneer of outer shell, and fine
microsculptural features on the outer shell are poten¬
tially important for defining clades within Calliotropidae.
They also are potentially useful for evaluating sister-
group relationships of seguenzioidean families. For
example, closely spaced wavy, irregular, and sometimes
bifurcating axial microstructure on the adult shell is
expressed in different patterns in at least three
seguenzioid families. Micrographs for ealliotropids are
figured as “wavy threads” by Marshall (1979) and for
ehilodontids as “scratch-like marks” by Herbert (2012).
They are expressed in a similar pattern on eataegids
(personal observation). This peculiar microsculpture is
typically accompanied by different patterns of infill or
overprint by intritacalx deposits that have not been char¬
acterized. Chalky or flakey, partially calcified exterior
layers on the shells of some gastropod and bivalve taxa
(D’Attilio and Radwin, 1971) are a form of remote bio¬
mineralization ( sensu Hickman, 2013). Intritacalx takes
on new taxonomic potential in terms of the untested
hypothesis that it is produced by sulfate-reducing
microbes (Vermeij, 2014).
MATERIALS AND METHODS
Locality data are given as they were originally reported
at the time of collection and recorded in their respec¬
tive museum data bases. Indonesian place names for
Albatross specimens use spellings in the U.S. Fish
Commission Oceanographic Data Set and data on the
original specimen labels, followed bv parenthetical cur¬
rent names, translations and spellings. Albatross depths
are reported in fathoms, as they were originally recorded,
followed by parenthetical conversions to meters. Acro¬
nyms for specimen repositories are: AMS: Australian
Museum, Sydney; ANSP: Academy of Natural Sciences
of Drexel University (formerly the Academy of Natural
Sciences of Philadelphia), Philadelphia; USNM: National
Museum of Natural History, Smithsonian Institution,
Washington, DC.
SYSTEMATICS
Subclass Vetigastropoda Salvini-Plawen, 1980
Superfamily Seguenzioidea Verrill, 1884
(Eucycloidea Koken, 1897, unranked)
Family Calliotropidae Hickman and McLean, 1990
Genus Calliotropis Seguenza, 1903
Type Species: Trochus ottoi Philippi, 1844. Pliocene-
Pleistocene, Italy, by original designation.
Calliotropis enantioserrata new species
(Figures 5-11)
Diagnosis: Shell conical, 8 whorls, large for genus
(height >25 mm), height exceeding width (hAv = 1.2)
conical, with straight-sided whorls and broad, smooth sub-
sutural ramp between two sharply serrate spiral cords;
suture impressed in deep channel between adapical spiral
with abapically pointed serrations and abapical spiral, on
succeeding whorl, with adapieally pointed serrations; umbi¬
licus completely closed by reflexed columellar callus;
aperture prosocline at 43° to axis of coiling, not descending,
but outer lip with prominent reflected terminal thickening.
Description: Exterior shell layer thin, ivory-colored,
smooth, lacking periostracum or continuous intritacalx
that covers shells of some large ealliotropid species. Fine
microstriations and intritacalx visible with magnification.
Nacre clearly visible through translucent exterior layer
and refracts green and pinkish iridescence. In addition
to two serrate spiral cords on each spire whorl, a third
faintly denticulate spiral present, covered by suture
throughout growth. Five fine, faintly denticulate spirals
on convex shell base. An interior oblique prismatic layer
(sensu Gainey and Wise, 1975) apparently lacking, and
nacre not extending onto terminally thickened and
reflected apertural lip of adult. Protoconch (Figure 9)
translucent, with one smooth whorl and distinct bound¬
ary with opaque and ornamented early teleoconch.
Flattened and broadly triangular serrations pointing
in opposite directions on either side of sutural channel
(Figure 10). Surfaces of tooth-like serrations finely striate
under magnification (Figure 11), and brown intritacalx
most prominently developed within sutural channel
(Figures 10-11).
Remarks: The new species is superficially similar to
Calliotropis excelsior Vilvens, 2004, which also has
flat-sided spire whorls and a suture located in a deeply
excavated channel bordered on either side by prominent
serrate spirals cords. However, C. excelsior has an open
umbilicus and lacks a terminally thickened apertural lip. It
is restricted in distribution to Fiji and New Caledonia,
C.S. Hickman, 2016
Page 87
Figures 5-11. Calliotropis enantioserrata new species. 5. Semi-apertural, 6. Apical, and 7. Basal views of holotvpe, AMS
C152225, height = 29.6 mm. 8. Inclined abapical view of the spire of paratype, AMS C142224, height = 15.8 mm. 9. Detail
of protoconch, from Figure 8. 10, 11. Details from Figure 9 of sutural channel, bordering rows of serrations, microsculpture
and intritaealx.
Page 88
THE NAUTILUS, Vol. 130, No. 3
disjunct from the new Australian species. Shell propor¬
tions of the holotypes of the two species are identical,
although all specimens of C. excelsior are smaller (height
<25 mm).
The flattened and broadly triangular serrations pointing
in opposite directions on either side of the sutural channel
(Figure 10) are one of the most distinctive features of
C. enantioserrata . The opposing direction of the serra¬
tions on the cords bordering the sutural channel is also
shared with some specimens that have been assigned to
Calliotropis pagodaformis (Sehepman, 1908), a species
first described from southeastern Indonesia with a range
that subsequently has been extended to the Solomon
Islands (Vilvens, 2007). The holotype of C. pagodaformis
(Figure 2) is distinctive in its narrower width and higher
spire as well as the concave spire profile giving it a classi¬
cally pagodaform appearance.
Type Material: Holotype, AMS C152225; height,
29.6 mm, maximum width 24.0 mm, from type locality;
Paratype: AMS C152224, height 15.8 mm, maximum
width 14.7 mm, R/V Toncaroa Station U222, 32°49.3' S,
152°49.T E, off Newcastle, 1040-1075 m, 9 October 1982.
Type Locality: R/V Toncaroa Station U223. 32°58.8' S,
152°41.6' E, off Newcastle, New South Wales, 951-
1150 m, 9 October 1982.
Other Material Examined: AMS C152228; a single
small, broken and worn specimen from R/V Toncaroa
Station U223, the type locality.
Distribution: Known only from two stations off
Newcastle, New South Wales, Australia.
Etymology: Greek enantios (opposing) + Latin serrata
(toothed edge), referring to the saw-like appearance of
the spiral cords on either side of the channel between
whorls and the opposing direction in which the flattened
tooth-like projections point.
Calliotropis andamanensis new species
(Figures 12-17)
Diagnosis: Shell conical, 8 whorls, moderately large
(maximum height = 25 mm), width exceeding height
(h/w = 0.92); spire profile straight-sided with three
spiral cords; subsutural ramp broad and unornamented
Figures 12-17. Calliotropis andamanensis new species. 12. Semi-apertural, 13. Apical, and 14. Basal views of holotype, ANSP
291376, Height = 2.5.0 min. 15. Semi-apertnral, 16. Apical, and 17. Basal views of paratype, ANSP 465552, height = 21.2 mm.
C.S. Hickman, 2016
Page 89
between adapical spiral and closely spaced pair of
sharply nodose spirals connected by closely spaced,
strongly sinuous axial threads; nodes most prominent
on middle spiral and weak but more numerous on
abapical spiral; suture closely following third abapical
spiral on spire whorls; aperture strongly descending
on body whorl; terminal lip flared and reflected, but
not strongly thickened; umbilicus broad and deep, par¬
tially covered by reflected projection of parietal and
eolumellar callus.
Description: Outer shell layer very thin, lacking per-
iostracum and intritacalx. Ivory-colored, with nacre visi¬
ble as a predominantly greenish refractive luster. Base
(Figures 14, 17) weakly convex and predominantly
smooth, with four to six smooth to faintly nodose spirals
bordering umbilicus. Nodes most prominent on spiral
immediately bordering umbilicus. Sinuous axial threads
increasingly visible between pair of abapical spirals
(Figures 12, 15), consisting of uniquely distinctive feature
of the species. Teleoconch whorls with mierostriations
hut lacking intritacalx. Prominent callus deposit divided
into non-nacreous reflexed thickening that is only partially
covered by a smaller tongue of nacre over umbilical
portion (Figures 14, 17).
Remarks: This species is superficially similar to the
species in “Formal group 4” of Vilvens (2007), which
includes relatively large shells in which width exceeds
height. Some of the species in this group have a similarly
robust development of parietal and eolumellar callus that
either partially or fully covers a broad umbilicus. How¬
ever, none of these species have a descending aperture
(Figures 12, 15), and they differ in whorl profile as well
as the number and ornamentation of spirals on the spire
and base.
The distinctive features of C. andamanensis are
consonant with its hydrographic isolation.
Although the Andaman Sea is in the Indian Ocean, it
is geologically a deep Sundaland basin, separated tecton¬
ically from shallow Indian Ocean floor by the Sunda
Trench. At the same time, it is topographically and
hydrographically separated from Indonesia and the deep
basins of Wallacea by the shallow Sunda shelf. There
is no significant connection between Wallacea and the
Andaman sea via the narrow and shallow Malacca
Strait between Sumatra and Malaysia. The Indonesian
Throughflow of Pacific water into the Indian Ocean (the
largest movement of ocean water on Earth) is via the
Makassar Strait between Sulawesi and Kalimantan,
entering the Indian Ocean between Lombok and Bali
and through the Timor Passage to join the west-flowing
South Equatorial Current.
Type Material: Holotype: ANSP 291376, height
25.0 mm, maximum width 27.3 mm; Paratype: ANSP
465552, height 21.2 mm, maximum width 26.8 mm. Both
from type locality.
Type Locality: R/V Anton Brulin International
Indian Ocean Expedition, Cruise 1, Station 17, 07°40' N,
97°08' E, Andaman sea, off Phuket Island, Thailand,
503-512 m, green-brown clay, 1963.
Distribution: Known only from the type locality in the
Andaman Sea off Phuket, west Thailand.
Etymology: Named for the type locality.
Calliotropis bukabukaensis new species
(Figures 18-22)
Diagnosis: Shell thin, conical, high-spired, and mod¬
erately large (height >15 mm), height exceeding width
(height/width = 1.23); spire whorls stepped, with two
shaqoly nodose spiral cords, suture impressed and slightly
channeled above adapical spiral row of nodes; shoulder
slope concave to slightly convex adapical to peripheral
spiral cord with coarse nodes drawn out and connected;
base with four thin, sharp spirals with concave interspaces
ornamented with very fine axial lines; umbilicus broadly
open, defined by finely beaded spiral and with fine axials
on umbilical wall; eolumellar lip thickened and with
prominent denticle extending into aperture.
Description: Nacre clearly visible through thin outer
shell layer, remnants of thin, brownish-gold intritacalx
present, best preserved on shell base and in umbilicus
(Figures 20-22). Protoconch-teleoconch boundary not
preserved on worn apex, but shell apparently with six or
seven whorls. Aperture broken on all specimens, with no
evidence of either thickened or descending apertural lip.
Umbilicus of holotype (Figure 20) partially blocked
by thin, partially ruptured papery covering. This appears
to have been formed by another organism, umbilicus of
paratype (Figure 22) not similarly blocked.
Remarks: The new species is clearly distinguished
from all previously described calliotropids by the very
strongly developed, nodular eolumellar tooth (Figures 18,
21). Vilvens (2007) noted and illustrated (p. 42, figs 156,
158, and 160) a eolumellar tooth in three small-shelled
species with depressed spires. In each of these species
the denticle is considerably less well developed.
This is one of three new species recovered by the
Albatross in the Gulf 'of Tomini. It is easily distinguished
from the large-shelled non-umbilicate species C. tominiensis
and the smaller non-umbilicate Calliotropis new species?
Type Material: Holotype: USNM 239247, height
15.9 mm, maximum width 13.0 mm. Paratypes: USNM
1297170, height 12.5 mm, maximum width 11.3 mm;
USNM 1409286, height 12.5 mm, maximum width
1 1 .4 mm. All from type' locality. Actual measurements
are incomplete due to specimen breakages.
Type Locality: U.S. Fish Commission, IVY Albatross,
Station 5613, 00°42'00" N, 122°44'00" E, Gulf of Tomini,
Page 90
THE NAUTILUS, Vol. 130, No. 3
Figures 18-22. Calliotropis bukabukaensis new species. 18. Apertural, 19. Apical, and 20. Basal views of holotype, USNM
239247, height == 15.9 mm. 21. Detail of the columellar tooth and umbilical microsculpture of holotype, scale bar = 1 mm.
22. Basal view of paratype with unoccluded umbilicus, USNM 1297170, height = 12.5 mm.
Celebes (Sulawesi), Togian Islands S. of Buka Buka
Island, 752 Fathoms (=1375 m), gray mud, 20 November
1909.
Distribution: Known only from the type locality.
Etymology: Named for the type locality.
Calliotropis tominiensis new species
(Figures 23-25)
Diagnosis: Shell conical, 7 whorls, high-spired (spire
angle = 58°, large (height >30 mm), height exceeding
width (height/width = 1.31), spire whorls with one
prominent, nodose spiral cord; suture not impressed,
closely following and almost completely covering second
abapical spiral; base convex, with 4 finely beaded spiral
cords; parietal and columellar callus reflexed, completely
obscuring umbilicus and partially covering innermost
basal spiral cord, aperture prosocline at 45° angle to axis
of coiling; terminal lip thickened but not descending.
Description: Exterior shell layer extremely thin, with
fine, closely spaced co-marginal lamellae and brownish-
gold intritacalx w'ell preserved on body whorl (Figure 23)
and base (Figure 25). Outer layer eroded away on spire
whorls (Figure 24) of holotype, exposing thicker nacre¬
ous layer that refracts predominantly green and pinkish
C.S. Hickman, 2016
Page 91
Figures 23-28. Calliotropis tominiensis new species and C. new species? 23. semi-apertural, 24. Apical, and 25. Basal views of
holotvpe of C. tominiensis , USNM 239217, height = 33.6 mm. 26. Apertural, 27. Apical and 28. Basal views of figured specimen
of C. new species, USNM 239216, height 15.7 mm.
iridescence (Figures 23-24). Inside aperture, nacre
covered by very thin oblique prismatic layer that can
be detected by touch — rougher toward interior and
smoother toward apertural margin. Protoconch deeply
eroded (Figure 24), and protoeonch/teleoconch boundary
not visible.
Remarks: There are very few species of Calliotropis
that are as large and high-spired as C. tominiensis.
Calliotropis conoeides Vilvens, 2007, known only from
the Solomon Islands, is similar in size and proportions
(h/w of the largest paratype = F3). However, the
Solomon species lacks the terminally thickened outer
lip. It also lacks the extensive parietal and columellar
callus that covers the umbilicus of C. tominiensis.
Calliotropis cynee Vilvens, 2007 from the Tanimbar
Islands, in the Banda Arc of southeastern Indonesia, is
less than half the height of the new species and has a
less elevated spire (h/w < 1.24), It also lacks the termi¬
nally thickened outer lip and extensive callus covering
the umbilicus.
Tbe biodiversity of the deep water basins of the
Wallacean region of Indonesia is very poorly sampled, in
contrast to the shallow waters of the region, which have
been considered a crucible of species formation. Lack of
sampling is unfortunate for two reasons. First, because
of the megadiverse deep settings resulting from the
complex and dynamic geological history of collisions, arc
Page 92
THE NAUTILUS, Vol. 130, No. 3
volcanism, subduction, suturing of continental fragments,
post-collision extension to form new deep embayments,
and tire topographic barriers to migration. Secondly,
because the region offers hydrographically and geologi¬
cally unique opportunities to sample very great depths
very close to shore in isolated embayments and adjacent
to numerous small islands.
It is fortunate that the Albatross spent a brief time in
Indonesia at the end of the 1907-1910 Philippine Expe¬
dition, and material dredged from the Gulf of Tomini
and Gulf of Bone in Sulawesi provides a unique window
on calliotropid biodiversity in two unusual tectonic set¬
tings. The significance of these settings is considered in
greater detail in the discussion section.
Holotype: USNM 239217, height 33.6 mm, maximum
width 25.7 mm.
Type Locality: U.S. Fish Commission, R/V Albatross,
Station 5606, 00°16'28" N, 1 21°33/30" E, Gulf of Tomini,
Celebes (Sulawesi), Togian Islands S. of Dodepo Id., S34
Fathoms (=1525 m), green mud, 17 November 1909.
Distribution: Known only from the type locality.
Etymology: Named for the type locality.
Calliotropis new species
(Figures 26-28)
Description and Remarks: An incomplete and worn
shell recovered by the R/V Albatross from >1000 meters
in the Gulf of Tomini has a combination of characters
that differentiate it from the five new Sulawesi species
described herein. It is most similar to, and from the same
station as, C. tominiensis. However, the peripheral spiral
cord is prominently nodose and the shell is covered by an
unusually heavy brown intritaealx with an exterior flaky
layer in which there is imbedded sediment. The covering
has adhered over most of the shell in spite of post¬
mortem corrosion and breakages (Figures 26-28). The
spire angle is greater (67°), and the h/w ratio is less (1.1).
The umbilicus is almost completely closed by parietal
callus (Figure 28), and the columellar lip, which is not
broken, is sharp rather than thickened and reflected as in
the holotype of C. tominiensis. Additional material would
be required to justify proposing a new name, and it is
possible that the specimen is a juvenile of C. tominiensis.
If umbilical closure is a terminal growth feature, this
smaller specimen would have to be considered an adult.
Illustrated Specimen: USNM 239216, height
15.7 mm, maximum width 14.1 mm.
Locality: U.S. Fish Commission, R/V Albatross, Sta¬
tion 5606, 00°16'28" N, 121o33,30" E. Gulf of Tomini,
Celebes (Sulawesi), Togian Islands, S. of Dodepo Id.,
834 Fathoms (=1525 m), green mud, 17 November 1909,
Calliotropis tabakaensis new species
(Figures 29-31)
Diagnosis: Shell conical, 7 whorls, large (height
~ 17 mm), high spired (spire angle = 72°); spire moder¬
ately elevated with width and height almost equal (height/
width = 0.9); spire whorls angularly convex, with two
nodose spiral cords separated by concavity, suture shal¬
lowly impressed, following and barely covering fine,
faintly beaded spire cord; adapical spiral on body whorl
weaker with loss of nodes, peripheral spiral with 26 nodes;
base weakly convex with four thin spirals that increase in
width adapieally, ornamented by slight swellings where
crossed fine, sinuous axial threads; umbilicus open, aper¬
ture prosoeline at 30° angle to axis of coiling; terminal lip
thin and not descending.
Description: Protoconch of approximately one whorl,
worn, boundary with teleoconch not clearly preserved
(Figure 30). Outer shell layer thin, underlying nacre
visible on entire shell surface. Intritaealx minimally
developed and visible only under microscopic examina¬
tion. Shell not encrusted during life or after death. Epi¬
sodes of breakage and repair restricted to minor chipping
of apertural lip.
Remarks: In size and proportions the shell fits in For¬
mal group 4 of Vilvens (2007) and is superficially similar
to Calliotropis debriosa Vilvens, 2004, a species known
from 9 stations in the southwestern Pacific from New
Caledonia to Fiji and at comparable depths. The new
species is distinguished by its extremely thin shell,
stepped appearance of the rounded whorls, open umbi¬
licus, two prominent nodose cords on the spire whorls,
and the nearly equal ratio of height to width.
This is one of three new species from the Gulf of
Bone. It is remarkable that the linear distance separating
the type locality of C. tabakaensis, off the western mar¬
gin of the southeast arm, from that of C. locolocoensis ,
off the eastern margin of the south arm is only 50 km.
However, the shelf on either side of the gulf is exception¬
ally narrow and drops off rapidly to depths of 2000 m in
the center of the basin. The basin itself is subdivided into
fault-bounded sub-basins, and study of the deep-water
marine molluscan fauna merits further sampling with
the geology and hydrography in mind. A feature of the
Gulf that is especially pertinent to the occurrence of
large-shelled calliotropid gastropods is the presence of
hydrocarbon seeps (Camplin and Hall. 2014).
Holotype: USNM 239445, height = 17.3 mm, maxi¬
mum width = 17.4 mm.
Type Locality: U.S. Fish Commission, R/V Albatross,
Station 5655, 03°34'10" S, 120°50'30" E, Gulf of Boni
(Bone), Celebese (Sulawesi), Buapinang, 3 miles off
Cape Tabaka (Tobaku). 608 Fathoms (= 1112 m), green
mud and fine sand, 18 December 1909.
Distribution: Known only from the type locality.
C.S. Hickman, 2016
Page 93
Figures 29-34. Calliotropis tabakaensis new species and C. locolocoensis new species. 29. Semi -ape rtural, 30. Apical and
31. Basal views of holotype of C. tabakaensis, USNM 239445, height = 17.3 mm. 32. Apertural, 33. Apical and 34. Basal views of
holotype ofC. locolocoensis , USNM 245478, height = 18.2 mm.
Etymology: Named for the type locality.
Calliotropis locolocoensis new species
(Figures 32-34)
Diagnosis: Shell conical, 9 whorls, unusually slender
and high-spired (spire angle = 55°), large (height >15
mm), height exceeding width (height/width = 1.23); spire
whorls with one bluntly spinose, keel-like spiral cord; base
shallowly convex and smooth, bordered adapieally bv
unomamented spiral; umbilicus almost completely cov¬
ered by reflexed columellar and parietal callus and bor¬
dered by two faintly nodose spiral cords; outer shell layer
covered bv thin but continuous tan-colored intritacalx,
thicker inner nacreous layer visible through the outer
shell layer where intritacalx has eroded away.
Description: Outer lip of holotype broken, but the
trace of the break indicates that the final lip was not
descending. Throughout ontogeny, it followed and covered
the unornamented spiral defining the break between the
spire and the base of the shell. The protoconch is approx¬
imately one whorl and worn so that the protoconch-
teleoconch boundary is indistinct.
Remarks: In size and general appearance there is a
close, but superficial, resemblance to C. pagodaformis
(Schepman, 1908), originally described under Solar-
ielliopsis Schepman, 1908 for inclusion of 1 1 deep-water
Page 94
THE NAUTILUS, Vol. 130, No. 3
Indonesian species. Schepman based die genus on
S. calcarata recognizing that his type species had a radula
(Figure 2) distinct from that of Solariella Wood, 1842.
Although he had no radular data for the odier ten forms
that he placed in Solarielliopsis , he emphasized (p. 53)
"how dangerous it is to classify such species, without any
knowledge of the soft parts.”
Schepman ’s original figure of the shell of C. pagoda-
fomiis is reproduced above (Figure 1) for comparison
with C. locolocoensis . The spire whorls of the new spe¬
cies have a single beaded spiral at the periphery and lack
the subsutural spiral with adapically directed spines. The
suture is not in a channel, and the shoulder slope is
slightly convex rather than flat. The base lacks distinct
spiral cords, and the columella is slightly prosocline
rather than coincident with the axis of coiling.
There are prominent plate tectonic and hydrographic
barriers separating the superficially similar pagodaform
species in Indonesia. Schepman’s localities from the
Siboga Expedition were in the Outer Banda Arc thrust
belt of eastern Indonesia, whereas the Albatross station
is in deep water very close to shore in the Gulf of Bone
in Sulawesi. The gulf is a complex, post-collision, exten-
sional basin, emptying to the south into the Flores Sea.
The gulf opened during the Neogene to separate the
south and southeast arms of Sulawesi (Sudarmono,
2000; Camplin and Hall, 2014). Four Indonesian locali¬
ties from which C. pagodaformis has been collected
more recently (March, 1990) and reported by Vilvens
(2007, p. 7) are also in the Banda Arc (Tanimbar Islands),
tectonically in the collision zone between the Australian
continental margin and the Banda volcanic Arc above
the Timor trough (Barber et al, 1986). The three new
species described here from the Gulf of Bone may
reflect its isolation and separate geologic history.
Holotype: USNM 245478. Height 18.2 mm, maxi¬
mum width 14.8 mm.
Type Locality: U.S. Fish Commission, R/V Albatross,
Station 5658, 03°32'40" S, 120°31'30" E, Gulf of Bom
(Bone), Celebes (Sulawesi), off Loko Loko (Locoloco),
510 Fms. (=933 m), gray mud, 19 December 1909.
Distribution: Known only from the type locality.
Etymology: Named for the type locality.
Calliotropis lamuluensis new species
(Figures 35-37)
Figures 35-40. Calliotropis lamuluesis new species and C. g lypta (Watson, 1879). 35. Semi-apertural, 36. Apical and 37. Basal
views of holotype of C. lamuluensis , USNM 239424. height = 19.4 mm. 38. Semi-apertural, 39. Apical and 40. Basal views of a
hypotype of C. g lypta, AMS C.l 15685, height = 17.2 mm.
C.S. Hickman, 2016
Page 95
Diagnosis: Shell broadly conical, 6 whorls, low spired
(spire angle = 90°, large (height « 20 mm), width
exceeding height (height/width = 0.84); spire whorls
weakly convex, with two (increasing by intercalation to
three) nodose spiral cords; suture shallowly impressed
beneath and following abapical spiral; body whorl with
three coarsely nodose spirals on shoulder slope (30, 34
and 38 nodes from adapical to abapical spiral on body
whorl), peripheral spiral more finely nodose (65 nodes
on body whorl); base weakly convex with seven sharply
nodose spirals that become increasingly wide and more
distinctively nodose abapically; parietal and columellar cal¬
lus strongly reflected and completely covering umbilicus;
aperture strongly prosocline at 43° angle to axis of coiling,
terminal lip very slightly thickened but not descending.
Description: Apertural lip of holotvpe chipped, four
roughly circular attachments scars (Figure 36) present
on body whorl made by an epizoan no longer present.
Two major and several minor episodes of shell breakage
and repair during growth. Periostracum not present.
Fine, sinuous and prosocline axial “scratch-marks” on
shell interspersed with minimally developed intritacalx.
Protoconch and earliest portion of teleoconch broken,
nacre visible in several places where very’ thin exterior
shell layer is broken or eroded away. Nacre in interior of
aperture covered, but visible through a thin oblique pris¬
matic layer that is a characteristic feature of many large
calliotropid shells. Nacreous layer in this species thicker
than in most large calliotropids, giving shell a more
robust appearance.
Remarks: Calliotropis lamuluensis differs dramati¬
cally in shell thickness, proportions, and many discrete
features from the two new species described from far¬
ther within the gulf. As noted above, this may reflect
isolation tied to the complex tectonic history and opening
of the gulf (Sudarmono, 2000; Camplin and Hall, 2014).
In view of the complex geology and hydrography and the
hydrocarbon seepage reported in the Gulf (discussed
above) these locations would profit from further sam¬
pling of the deep-water marine molluscan fauna.
The new species is closest in size, proportions, and
ornamentation to the species in Formal group 4 of
Vilvens (2007). It has the lowest hAv ratio of any of the
species in the group and differs from these openly
umbilicate shells in having a fully reflexed umbilical clo¬
sure (Figure 37). It also differs from group 4 species
in having a thicker shell. An Australian specimen of
Calliotropis glypta (Watson, 1879), is illustrated here for
comparison (Figures 38-40). Thickness of the basal por¬
tion of the apertural lip is 0.4 mm, in contrast to 0.7 mm
for C. lamuluensis . Shells of both species supported
encrusting epizoans to a greater extent than typical in
other calliotropids.
Holotype: USNM 239424, height = 19.4 mm, maxi¬
mum width = 23.1 mm.
Type Locality: U.S. Fish Commission, R/V Albatross,
Station 5650, 4°53'45" S, 121°29'00" E, Gulf of Boni
(Bone), Celebese (Sulawesi), Buapinang, off Lamulu Point,
540 Fathoms (=988 m), green mud, 17 December 1909.
Distribution: Known only from the type locality.
Etymology: Named for the tvpe locality.
DISCUSSION
The new species described above shift the center of diver¬
sity of large-shelled (height > 15 mm) calliotropids into
the tectonically and hydrographically complex Wallacean
region of Indonesia, with its long history of plate colli¬
sions, subduction, and volcanic arc volcanism. They also
shift peak diversity of large-shelled calliotropids to depths
exceeding 1000 m. This is remarkable because the
deep-water fauna of Indonesia is still undersampled. A
previous paper (Hickman, 2012) provides additional
background on the unique geologic, biogeographic, and
geologic aspects of Wallacean deep diversity’.
A simplified map of deep Wallacea (Figure 41) shows
the localities of the species described above and provides
a frame of reference for the following discussion of pat¬
terns of narrow endemism, disjunction, and relictual
accumulation of taxa.
Tl le calliotropid species are spatially distributed in
geologic settings that include tectonically isolated deep¬
water basins underlain by oceanic crust. In one interpre¬
tation (Lee and McCabe, 1986) the Sulu, Celebese,
and Banda seas are characterized as remnants of a
large Cretaceous-Eocene seaway. Tectonic elements of
Wallacea include major forearc subduction complexes,
fusions of colliding microplates, shifting plate boundaries,
and formation of narrow gulfs and extensional basins
opening by rollback at subduction hinges. The Molucca
Sea is unique — the only modern example of double sub¬
duction at the collision of two facing volcanic arcs, elimi¬
nating an entire oceanic plate and trapped Indian Ocean
lithosphere (McCaffrey et al., 1980; Hall, 2006; Hall and
Smyth, 2008). This is the first recognition of the relictual
deep-water fauna in a setting that has heretofore been of
intense interest only to geologists.
The corresponding sedimentary and geochemical set¬
tings associated with the complex geologic history of
marine Wallacea are consistent with the development of
unusual deep marine communities. Convergent plate
margins frequently are sites of expulsion of squeezed per¬
colating fluids containing sulfides and hydrocarbons
(oil, gas, and methane hydrates). These fluids feed micro¬
bial productivity and development of chemosymbiotically
based food chains. Although such settings are typically not
good candidates for the formation of large petroleum res¬
ervoirs, ongoing investigations of petroleum potential in
Indonesia have revealed the presence of discrete hydro¬
carbon seeps, mud volcanoes and explosive methane
release (Barber et al., 2000; Sudarmono, 2000; Charlton,
Page 96
THE NAUTILUS, Vol. 130, No. 3
Figure 41. Simplified map of the deep Wallacean region of the Indonesian Archipelago with localities tor the new calliotropid
species (solid circles) and previously described calliotropids from the Siboga Expedition (open circles) and French KARUBAR
Expedition (open squares). The locality of the previously described new gazid genus and species is shown by a solid star. Isolated
occurrences in deep-deep-sea basins, gulfs and straits are depicted in the context of primary zones of collision and subduction
(solid lines with triangular tick marks) and associated volcanic arcs at plate boundaries where subduction has occurred or
is occurring.
2004; Jablonski et ah, 2007; Camplin and Hall, 2014) as
well as sediment systems with diffusive methane emission
and a deep biosphere of chemosymbiotic microbial com¬
munities (Schippers et ah, 2010; Siegert et ah, 201 1).
Relationship of Shallow and Deep
Biogeographic Patterns
The patterns and explanations of deep-water biodiversity
and connectivity in Indonesia are highly unlikely to cor¬
respond with those that have begun to emerge for shal¬
low water taxa. In shallow-water taxa there is a striking
disconnect between surface currents and routes of gene
flow (e.g., Benzie, 1998, 1999; Benzie and Williams,
1995, 1997; DeBoer et al, 2008), perhaps related to
underestimation of the importance of seasonal cycles in
the upper-ocean mixed layer in which monsoonal forcing
generates reversals in flow directions (Lindstrom et ah,
1987; McCreary et ah, 2001). Current directions and
flow speeds are notably different in deep water. Sills
between deep basins impede flow, and the vetigastropod
taxa under consideration here lack the feeding plank¬
tonic larvae that enhance the dispersal potential of many
shallow-water taxa. Deep-water diversity is likewise
unaffected by the diversity increase in reef-associated
habitats and the Oligo-Miocene pulse of shallow-water
diversification of reef-associated taxa in the Indo-West
Pacific (Williams and Duda, 2008).
In fact, it is argued here that modern marine biogeo¬
graphic patterns in the Indonesian region cannot be
understood apart from geologic history. Reconstruction
of the entire Cenozoic history of plate boundaries (Hall,
2001, 2002), especially the dramatic effects of episodic
volcanic arc processes on marine topography (Hall and
Smyth, 2008), argue against simplistic inferences of marine
connectivity and historical biogeography based solely on
present configurations.
Explaining disjunct Deep-water Taxa in Wallacea and
Southern Africa
Tbe disjunct distribution of calliotropid species in deep
Pacific waters of Wallacean Indonesia and the far west¬
ern Indian Ocean has been recognized for more than
100 Years. Schepman (1908) reported Calliotropis
metallica (Wood-Mason and Alcock, 1891) from four sta¬
tions in Indonesia. The species is better known from
Madagascar and southern Africa, and was described
C.S. Hickman, 2016
Page 97
originally from the Gulf of Mannar in the Indian Ocean.
Calliotropis buccina Yilvens, 2006, described from
Reunion Island in the southwestern Indian Ocean is
now reported from the Kai archipelago in southeastern
Indonesia as well as the Solomon Islands, adding a third
disjunct element (Vilvens, 2007). This peculiar pattern is
not restricted to calliotropid gastropods, although it does
appear to occur in lineages with a long, pre-Cenozoic
geologic history. It is consistent with the hypothesis that
these are relictual occurrences of taxa and lineages with
ranges that have been fragmented and reduced. Two
additional examples are given here.
Abyssochrysid Gastropods
The first is in the distribution of living species of the
caenogastropod family Abyssoehrysidae Tomlin, 1927.
In a revision of the genus, Houbrick (1979) provided a
detailed analysis of the shell morphology, anatomy, and
radulae of the two living species: one described by
Tomlin (1927) from 1645-1828 m off the Cape of Good
Hope and the other described by Schepman (1909) from
the Sulu Sea in Indonesia. Based on further examination
of hydrographic data and search of the fossil record,
Houbrick (1979) proposed that these two living abysso-
ehrisids are deep-water relicts of the Paleozoic- Mesozoic
superfamily Loxonematoidea (Koken, 1889). Although
Houbrick (p. 1) considered his proposal to be “radical”,
description of new material from deep-sea expeditions
(Bouchet, 1991) recognized five species, including a
new species from the Makassar Strait in Indonesia and a
distribution that is localized in the deep-water settings
around Sulawesi.
The Indonesian material examined by Houbrick (1979)
included specimens from the Albatross expedition, and
it is interesting to note that one of the specimens appears
to be from the same locality in the Gulf of Bone as
the tvpe of the Calliotropis species described above as
C. lamuluensis. Houbrick did not give the station number
or coordinates, but the reported depth is the same as well
as the location off Lamulu Point.
Description of a putative abyssochrysid from the mid¬
dle Eocene of the northwestern Olympic Peninsula in
western Washington (Goedert and Kaler, 1996; Kiel,
2008) adds a new paleoenvironmental dimension to the
post-Mesozoic disjunct geographic distribution. The
Eocene species is known from >50 specimens in a local¬
ized methane-derived authigenic carbonate deposit in
association with chemosymbiotyic solemyid, lucinid,
thyasirid, and vesieomyid bivalves as well as abundant
serpulid and vestimentiferan worm tubes.
Living Coelacanths
The same pattern of disjunction is not confined to marine
invertebrates. It has received considerable attention with
discovery of a second population of the “living fossil” coe-
lacanth genus Latimeria off Manado, Sulawesi, Indonesia
(Erdmann et al., 1998). The Sulawesi population is
separated by a distance of >10,000 km from the African
population in the Comros Islands (see Springer, 1999;
Holder et al., 1999). Although the two populations must
share a common ancestor, there is no available evidence
to test alternative historical biogeographic hypotheses.
CONCLUSIONS
Knowledge of the internal phylogenetic structure of
Calliotropidae is highly unsatisfactory, although the ana¬
tomical, and molecular data are consistent with place¬
ment of Calliotropidae in Seguenzioidea. For most of
the living species and available genus-group names there
are no anatomical or molecular data and no consistent
terminology for shell features. There are no published
accounts of the behavior of live animals and little infor¬
mation on habitats aside from records of some large-
shelled deposit-feeding species with enlarged hindguts
in hydrocarbon seep biotopes where primary productiv¬
ity is greatly enhanced by microbial chemosynthesis.
Problems are compounded by the long fossil record and
history of paleontological treatments using separate
names and classification.
On the positive side, resurgence of interest in deep-
sea exploration and use of new sampling methods are
adding considerably to the amount of new material avail¬
able for study. In addition, there seems to be increased
interest in describing older material that has been
languishing in museum collections. New techniques
enable illustration of microsculpture and features that
have not been noted previously. This paper draws special
attention to the potential of new characters for diagnos¬
ing taxa.
The discovery of elevated deep-water diversity of
calliotropid gastropods in the Indo-West Pacific, especially
in the deep basins in the Wallacean region of Indonesia,
should not be surprising in view of new data on regional
tectonic history: collision of the Australian, Eurasian, and
Philippine Sea plates; development of volcanic island arcs;
arc-arc and arc-continent collisions; subduetion; deforma¬
tion, faulting, rotations and translations; openings and
closings of distinct deep-water basins, gulfs and straits; and
the movement and accretion of microplates. Evolutionary
persistence of Indian Ocean faunal elements in Wallacea is
not a difficult fate to imagine for taxa arriving on Indian
Ocean lithosphere trapped and ultimately subducted in
the collision zone.
New understanding of seafloor topography and
hydrography of the Indo-West Pacific is in turn related
to the shallow and deep-water current systems that
affect biogeographic patterns. History and physical ocean¬
ography of Indonesian Throughflow and the gateway
connection between the Pacific and Indian Oceans is
potentially a strong key to understanding some of the
puzzling biogeographic connections between southern
Africa and Indonesia.
THE NAUTILUS, Vol. 130, No. 3
Page 98
ACKNOWLEDGMENTS
I thank the many colleagues who have provided assistance
in my efforts to document deep-water vetigastropod taxa
with uncertain assignments and phylogenetic affinities
and to document specimens that have remained unde-
scrihed for many years in museum collections. I am
especially grateful to Jerry Harasewyeh (USNM), Paul
Callomon (ANSP), and Phil Colman, Ian Loch, Alison
Miller, Mandy Reid, and Janet Waterhouse (AMS) for
assistance with numbers and collection records. Scanning
electron micrographs of radulae were recorded originally
on film at the Biological Electron Microscope Facility at
the University of Hawaii, Manoa, as part of a longterm
project documenting molluscan biomineralized struc¬
tures. I thank Dave Strauss for the digital images of
shells and for his expert assistance with preparation of the
final figures. The map was drafted by Marla Coppolino.
Dili H erbet and Claude Vilvens provided valuable reviews
and suggestions. This is contribution number 2074 from
the University of California Museum of Paleontology.
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THE NAUTILUS 130(3):101-1 15, 2016
Page 101
Latest Oligocene and Miocene whelks (Gastropoda: Neogastropoda:
Bnccinidae) from Peru
Thomas J. DeVries 1
Burke Museum of Natural History and Culture
University of Washington
Seattle, WA 98195 USA
ABSTRACT
Misifulgur new genus includes fusiform buccinid gastropods
with shells that are notably thin medially between the shoulder
and base, have moderately long siphonal canals, spiral cords
that are well developed anteriorly but often fade posteriorly,
an inner lip without plications, a smooth inner surface of the
outer lip, and no axial sculpture other than growth lines. The
oldest Misifulgur , M. dockenji new species, comes from lower
Oligocene deposits in Mississippi. While no later records
of Misifulgur are known from Gulf Coast states orCentral America,
Misifulgur whelks (i.e., M. sula (Olsson, 1931), M. cruziana
(Olsson, 1932), M. macneili new species) are the most com¬
monly preserved neogastropods from the latest Oligocene
through middle Miocene on sandy substrates at shelf depths
in the Peruvian Faunal Province. The most recent Misifulgur,
M. montemarensis new species, lived on protected sandy
substrates along the coast of southern Peru during the late
M ioeene. The late Miocene extinction of Misifulgur was
contemporaneous with the appearance on the inner shelf of
possible competitors, especially muricids of the genus Chorus
Gray, 1847.
Additional Keywords: Miocene, Mississippi, taxonomy, paleon¬
tology, biogeography
INTRODUCTION
The East Pisco Basin was a locus of marine deposition in
south-central Peru from the late Eocene until the early
Pliocene (DeVries, 1998). Forearc sedimentary strata,
which became emergent during the late Pliocene and
Quaternary, record shoreface, shallow subtidal, and shelf
paleoenvironments (DeVries and Schrader, 1997). Mol-
lusks are abundantly preserved in nearshore deposits
and, to a lesser extent, in the medium-grained, biotur-
bated, and winnowed sandstones associated with inner-
shelf water depths. Among neogastropods, the most
prominent on the latest Oligocene and Miocene Peru¬
vian shelf were heretofore unrecognized whelks of the
family Bnccinidae.
1 Mailing address: Box 13061, Burton, WA 98013 USA.
The first evidence of late Oligocene-Miocene Peru¬
vian whelks consisted of three incomplete specimens
from the Talara and Progreso forearc basins of north¬
ern Peru. These gastropods were mischaracterized by
Olsson its two species of muricids, Acanthina (Chorus) sula
Olsson, 1931 and Acanthiza ( Chorus ) sula cruziana
Olsson, 1932. New fossil whelks from the East Pisco
Basin and the more southerly Sacaco Basin, including
specimens with intact protoconchs and siphonal canals,
show that Peruvian whelks were widely distributed and
common in shelf environments throughout the Miocene
until they became extinct at about 6 Ma.
The Peruvian whelks are similar to some Paleogene
buccinids from tire southeastern United States included
by MacNeil and Dockery (1984) in the genus Pseudofulgur
MacNeil, 1984, a taxon comprised of fusiform buccinids
from the lower Oligocene Byram Formation of Mississippi.
MacNeils genus has been re-evaluated after examining the
Peruvian fossils, type specimens of Pseudofulgur, and a
new specimen provided by D. Dockery' (Office of Geology,
Department of Environmental Quality, Mississippi).
GEOLOGY
Onshore outcrops of the Talara Basin in northern Peru
extend from the Illescas Mountains (6° S) nearly to
the Ecuadorian border (3°45' S). The Progreso Basin
passes from northernmost Peru into southwestern Ecuador
(Figure 1). Several thousand meters of marginal marine
and deeper water sediments accumulated in these forearc
basins throughout the Cenozoic (Zuniga and Cruzado,
1979; Higley, 2004a, 2004b; Martinez et ah, 2005). Oligo¬
cene and Miocene deposits are especially well developed
north of Cabo Blanco (Palacios, 1994). Whelks collected
by Olsson (1931) from shallow-water shales at Caleta
Mero (3°50' S) were attributed to the basal upper Oligo¬
cene or lower Miocene Heath Formation (Olsson, 1931;
Engelhardt and Wood, 1993; Higley, 2004a), an assign¬
ment consistent with both current geological mapping
(Palacios, 1994) and the concurrent presence of Turritella
woodsi Lisson, 1925 (= Turritella conquistadorana Hanna
and Israelsky, 1925), which ranges in southern Peru from
Page 102
THE NAUTILUS, Vol. 130, No. 3
the late Eocene to latest Oligocene (DeVries, 2007b). A
single whelk specimen from Quebrada de la Cruz, a lew
kilometers east of Caleta Cruz (3°38' S), was attributed
to basal sandstones of the upper Oligocene to lower
M iocene lower Zorritos Formation, which, depending on
the precise locality, is stratigraphically correlative with
or conformably overlying Heath shales (Olsson, 1932).
Olsson’s (1932) stratigraphic attribution is consistent with
current mapping (Palacios, 1994) and age assignments
(Higley, 2004b). ‘
The East Pisco Basin (Figure 1) is filled with many
hundreds, rather than many thousands of meters of
Cenozoic marine sedimentary rock (Montoya et ah, 1993;
Leon et ah, 2008). Four major depositional sequences are
recognized, each associated with an eponymous forma¬
tion: the Paracas (upper Eocene), Otuma (uppermost
Eocene), Chilcatay (uppermost Oligocene to lower
middle Miocene), and Pisco (middle Miocene to Pliocene)
(DeVries, 1998). Fossil whelks in southern Pern are found
in the Chilcatay and Pisco depositional sequences, rang¬
ing from the base of the former (about 25 Ma) to the
upper half of the latter (about 6 Ma). The most south¬
erly occurrence of Miocene Peruvian whelks is in the
Sacaco Basin (Figure 1), situated about 50 km south of
o°s yBasin
LIMA
1 5°30’S
B8360
B8373
74°44’W
Ccrros
Colorados
PERU
14 22 S
B8376
’
B8378
Pacific
12 S
Ocean
East 1
Pisco
Basin
Sacaco
Basin
400
SCALE (km)
1 8°S
72 W
78 w
Figure 1. Peruvian Cenozoic forearc basins and type localities of Peruvian species of Misifulgur. Progreso and Talara basins alter
Higley (2004). Upper inset shows type localities of M. sula (Olsson, 1931) (Caleto Mero) and M. cruziana (Olsson, 1932) (Quebrada
de la Cruz). Middle inset shows type locality' of M. macneili new species (B8376) and a nearby locality (B8378). Lower inset shows
type locality of M. montemarensis new species (B8360) and a nearby locality (B8373). Inset images are from Google Earth.
A stratigraphic column for the Cenozoic section in the East Pisco Basin is shown at upper right.
T.J. DeVries, 2016
Page 103
the East Pisco Basin, where a few hundred meters of
inner shelf and nearshore sediments were deposited dur¬
ing the middle to late Miocene and Pliocene (Muizon
and DeVries, 1985).
MATERIALS AND METHODS
Specimens from southern Peru were collected by the
author. Comparative material was provided by the Acad¬
emy of Natural Sciences of Philadelphia (Pennsylvania;
ANSP), the Office of Geology, Mississippi Department
of Environmental Quality (Jackson, Mississippi; MGS),
the Paleontological Research Institution (Ithaca, New
York; PRI), and the United States National Museum
of Natural History (Washington, DC; USNM).
Locality-sample descriptions, listed in the appendix,
are designated with a code referencing the author’s field
notes (DV xxxx-xx) and a code referencing collec¬
tions of the University of Washington’s Burke Museum
of Natural History and Culture (Bx x x x). Lengths (L)
and widths (W) are measured in millimeters. Dimen¬
sions of broken specimens are enclosed by parentheses.
Most figured specimens were coated with ammonium
chloride prior to photography. Types and other figured
specimens from southern Peru are deposited at the
Burke Museum (UWBM) in Seattle and the Laboratorio
de Paleontologfa de Vertebrados, Museo de Historia
Natural, Universidad Naeional Mayor de San Marcos
(MUSM INV) in Lima, Peru.
SYSTEMATICS
Superfamily Buccinoidea Rafinesque, 1815
Family Buccinidae Rafinesque, 1815
Genus Misifulgur new genus
Type Species: Misifulgur dockeryi new species.
Oligocene, Mississippi.
Diagnosis: Shell fusiform; shell veiy thin midway
between shoulder and base. Protoconch mammillate,
paucispiral, smooth, slightly deviated. Axial sculpture of
radial orthocline growth lines. Spiral sculpture of pri¬
mary cords at base, usually becoming weak or obsolete
adapieally near shoulder. Columellar and parietal cal¬
luses absent. Siphonal fasciole obsolete. Inner lip with¬
out plications. Inside of outer lip smooth.
Description: Shell medium to large sized, fusiform,
very' thin medially between shoulder and base. Protoconch
mammillate, two to three whorls, smooth, slightly deviated.
Spire moderately elevated; sutures variably impressed.
Sutural ramp planar, slopes and widths highly variable,
marked at shoulder by spiral cord of varying strength.
Axial sculpture consisting of radial orthocline growth lines
without sutural sinus. Spiral sculpture on smaller whorls
with 12 to 20 subequally spaced primary spiral cords;
primary interspaces half as wide to slightly wider than
primary cords, U-shaped or flat-bottomed, rarely with a
medial secondary spiral cord. Primary spiral cords on later
whorls becoming weaker or absent near shoulder and on
sutural ramp. Spire whorls with sutural ramp and three to
seven primary spiral cords anterior to shoulder. Aperture
elongate. Siphonal canal about 20% of shell length,
straight to moderately recurved abaxially. Columellar and
parietal callus absent; inner lip without plications.
Siphonal fasciole obsolete. Inside of outer lip smooth.
Distribution: Early Oligocene, southeastern United
States. Late Oligocene to early Miocene, southwestern
Ecuador, northern and southern Peru. Middle to late
Miocene, southern Peru.
Etymology: Misi,’ from the native North American
Ojibwa word for ‘great.’
Remarks: Misifulgur new genus includes fusiform
Mississippi specimens assigned to Pseudofulgur by
MacNeil and Dockery (1984) that have flattened spiral
cords that fade in prominence posteriorly, a veiy thin
shell medially between shoulder and base, and a smooth
inside of the outer lip (Figures 2-5). Excluded are shells
that have a lirate and and/or thickened outer lip, a pau¬
cispiral mammillate protoconch with closely spaced axial
ribs on the last protoconch whorl, and shells with
whorls that are not thinned medially, i.e., the holotype
of P. vicksburgensis (Conrad, 1848) (Figures 6-7), the
holotype of P. lirata Dockery, 1984, and a new specimen
from the Byram Formation of Mississippi referred
to P. lirata by Dockery (written communication, 2007)
(Figures 8, 9), all of which are probably tonnoideans
assignable to Ranellidae Gray, 1854 (sensu Beu and
Cernohorksy (1986) and Ben (2010)).
Specimens of Misifulgur from Mississippi and western
South America resemble the ranellids Linatella caudata
(Gmelin, 1791), a geographically wide-ranging Tethyan
cymatiine from the Miocene to the present (Beu, 2010),
and Monoplex weigmanni (Anton, 1838), a cymatiine
with a late Cenozoic Atlantic-Pacific trans-isthmus distri¬
bution (Beu, 2010). Shells of both cymatiines, however,
are not thinner medially, have a crenulate or lirate inside
to the outer lip, exhibit nodulosity at the shoulder that
reflects a muted axial sculpture, and on many larger spec¬
imens, a callused columella with transverse lirations — all
characters absent on specimens of Misifulgur of any size.
Modem buccinids and fasciolariids from western South
America with a moderately elongate siphonal canal (e.g.,
Aeneator fontainei (d’Orbigny, 1839); Pleuroploca spp.;
Solenosteira fusiformis (Blainville, 1832); see Alamo and
Valdivieso (1997) for geographic distribution data) have
either axial sculpture, a finely lirate inner margin of the
outer lip or lirate structure beneath a smooth inside sur¬
face of the outer lip, or both. Species of the muricid
genus, Peonza Oliveira, 1994, resembles the most strongly
keeled examples of Misifulgur cruziana, but the nodulo¬
sity of the spiral cord on the shoulder, the absence of
anterior inflation of the last whorl, and a corresponding
THE NAUTILUS, Vol. 130, No. 3
Page 104
presence of a very narrow and elongate siphonal canal
easily distinguish the Eocene-to-Miocene Chilean and
Argentinian taxon from Misifulgur.
The origin of early Oligocene Misifulgur and its system¬
atic position with respect to other buecinids remains
obscure. The well-constrained morphological variation of
American Misifulgur from the early Oligocene to late
Miocene makes it unlikely that European Eocene and
Miocene ta\a cited by MacNeil and Dockery (1984: 140)
have a close affinity. Pliocene North Pacific buecinids
with posteriorly fading spiral cords cited by those authors,
e.g., Chrysodomus .stantoni Arnold, 1908, may be Californian
biogeographic outliers of the northwest Pacific genus,
Clinopegma Grant and Chile, 1931, which has a stubby
recurved siphonal canal with a well developed columellar
callus and siphonal fasciole (Figure 10). Northern hemi¬
sphere buecinids with longer recurved siphonal canals,
i.e., many species of Neptunea Linne, 1758, have a sutural
platform and a strong spiral cord or keel on the shoulder,
as do some large specimens of M. cruziana. The pattern of
spiral cords for all Neptunea species, however, does not
vary from the spire whorls to the last whorl, whereas on
most large specimens of Misifulgur spiral cords anterior
to the shoulder on spire whorls fade to obsolescence on
the last whorl.
Pseudofulgur and, by extension, Mississippi specimens
herein assigned to Misifulgur were compared by Dockery
(MacNeil and Dockery, 1984: 140) with the late Miocene
Halia americana Olsson, 1922, from Panama and Costa Rica
(Olsson, 1922; I bias, 1942: 310, unfigured, as "Halia ef.
americana ’; Woodring, 1964: 290, as Ampulla americana
(Olsson, 1922), not Halia Risso, 1826; Petuch, 2003: 49).
With a deep turrid-like sinus close to the suture (Figures 11,
12), Olsson’s Costa Rican specimen cannot be placed in
the volutid genus. Ampulla Boding, 1798, nor in the
buceinid genera, Pseudofulgur and Misifulgur.
Mansfield (1935) doubted Olsson s (1922) attribution
of the “americana” species to the volutid genus, Halia,
but saw similarities between Olsson’s species and the late
Miocene Floridian Neptunea (?) alaquaensis Mansfield,
1935, including the presence of a sutural sinus (Figures 13,
14). Mansfield’s species does have spiral sculpture similar
to that of Pseudofulgur and Misifulgur, but its sutural
sinus sets it apart from those genera.
Two northern Peruvian buceinid species assigned herein
to Misifulgur, M. sula (Olsson, 1931) and M. cruziana
(Olsson, 1932), were placed by their author in the muricid
taxon, Acanthina (Chorus), also spelled Acanthiza
(Chorus). Olsson’s gastropods are neither Acanthina
Fischer von Waldheim, 1807 nor Chains Gray, 1847, both
ocenebrine murieids with an anterior external groove and
associated enrolled labral tooth (DeVries, 1997, 2003).
Specimens of M. sula and M. cruziana have neither.
Acanthina ( Chorus ) meroensis Olsson, 1931, not assigned
herein to Misifulgur, was also incorrectly assigned to
Acanthina ; specimens do have an anterior external groove
(indeed, two such grooves) and at least one labral tooth,
but the square-bottomed groove and spatulate tooth differ
from the V-shaped groove and enrolled tooth o {Acanthina
and Chorus. Acanthiza (Chorus) solida (Nelson, 1870) and
Acanthiza (Chorus) valuta Olsson, 1932, both Peruvian
taxa, are neither Misifulgur, Acanthina, nor Chorus.
Olssons “voluta” species has been referred to the
pseudolivid genus, Testallium Vermeij and DeVries,
1997 (Vermeij, 1998).
Misifulgur dockeryi new species
(Figures 2-5)
Pseudofulgur vickshurgensis (Conrad, 1848). — MacNeil
and Dockery, 1984: 141, pi. 32, fig. 5; pi. 52, fig. 13; not
pk 32, fig. 11.
Diagnosis: Shell length to 45 mm. Sutural ramp weakly
planar, sloping anteriorly 45 degrees. Spiral sculpture of
about 20 broad primary spiral cords between base and
shoulder. Secondary spiral threads present on both sides
of shoulder inflection.
Description: Shell length to 45 mm, fusiform, moder¬
ately constricted, inflection two-thirds distance from last-
whorl suture to anterior end; very thin medially between
shoulder and base. Protoconch unknown. Teleoconch
with five to six whorls. Spire moderately elevated; sutures
strongly impressed. Sutural ramp weakly planar, sloping
anteriorly about 45 degrees from suture, with a weak
primary spiral cord at shoulder inflection. Axial sculpture
of radial orthocline growih lines, twisted adaperturally at
suture. Spiral sculpture of about 20 broad flat-topped
primary spiral cords between base and shoulder,
subequallv spaced, becoming obsolete adapically towards
shoulder, replaced in part by closely spaced secondary
spiral cords and threads on both sides of shoulder. Pri¬
mary interspaces half as wide to as wide as primary spiral
cords, flat-bottomed, rarely with an intercalated second¬
ary' spiral cord or tertiary thread. Spire whorls with sutural
ramp, shoulder and four or five prominent primary spiral
cords. Aperture elongate, ovate. Siphonal canal 20 per¬
cent of shell length, nearly parallel to shell axis, broadly
open. Columellar and parietal callus absent. Siphonal
fasciole obsolete. Inner lip without plications. Inside of
outer lip smooth.
Type Material: MGS 1227, holotype, locality 115,
L 22.9, W (15.0). Byram Formation, lower Oligocene.
Type Locality: MGS locality 115, east side Highway
61 across from the Anderson Tully Lumber Company at
the boundary of Sections 12 and 35, T.16 N., R.3 E.
(MacNeil and Dockery, 1984: 405).
Other Material Examined: USNM 136798, USNM
locality 259 (top of bluff, Vicksburg, Warren County,
Mississippi), H 42.2, W 21.9. Byram Formation (Marl
Member), lower Oligocene.
Distribution: Lower Oligocene, Byram Formation,
Mississippi.
T.J. DeVries, 2016
Page 105
Figures 2-14. Gastropods assigned to or compared with Pseudofulgur MacNeil, 1984 by MacNeil and Dockery (1984).
2-5. Misifulgur dockeryi new genus new species. Mississippi. Early Oligocene. 2. USNM 136798. USGS locality 259. Apertural
view. Length 42.2 mm. 3. USNM 136798. Abapertural view. 4. MGS 1227. MGS locality 115. Holotype, apertural view.
Length 22.9 mm. 5. MGS 1227. Abapertural view. 6-9. Pseudofulgur vicksburgensis (Conrad, 1848). Mississippi. Early
Oligocene. 6. ANSP 13475. Holotype. Apertural view. Length of exposed shell is 21.4 mm. 7. ANSP 1.3475. Abapertural view.
8. MGS 1905. Apertural view. Length 22.4 mm. 9. MGS 1905. Abapertural view. 10. Clinopegma magnum (Dali, 1895). UWBM
98677. Bering Sea. Recent. Apertural view. Length 71.2 mm. 11, 12. Halia americana Olsson, 1922. PHI 20921. Holotype. Near
Mt. Hope, Panama. Late Miocene. 11. Apertural view. Length 65.4 mm. 12. Lateral close-up showing sutural sinus, including portion
lapping onto penultimate whorl. 13, 14. Neptunea (?) [sic] alaquaensis Mansfield, 1935. USNM 373139. Holotype. Station 12046,
Vaughan Creek, upper locality, Walton County, Florida. Late Miocene. 13. Apertural view showing sutural sinus. Length 35 mm.
14. Abapertural view showing sutural sinus.
Etymology: Named in honor of David T. Dockery, III
(Mississippi Office of Geology), in recognition of his
work on fossil Cenozoic mollusks of the southeastern
United States.
Remarks: Specimens of Misifulgur dockeryi new spe¬
cies differ from those of Pseudofulgur vicksburgensis in
having a thin shell medially between shoulder and base
and a smooth inner surface of the outer lip. Specimens of
THE NAUTILUS, Vol. 130, No. 3
Page 106
the latter species have a strongly lirate inner surface of
the outer lip and no mid-whorl thinning. Specimens
of M. dockeryi lack axial sculpture, whereas those of
P. vicksburgensis have growth lines thickened to the
point of becoming low, flat, axial ribs (Figures 7, 9).
Populations of Misifulgur dockeryi , as well as
Pseudoftdgur spp., inhabited sandy and muddy substrates
on an aggrading continental shelf (Coleman, 19S3).
Misifulgur sit I a (Olsson, 1931)
(Figures 15-18)
Acanthina (Chorus) sula Olsson, 1931: 107, pi. 18, figs 1. 2.
Diagnosis: Shell fusiform, strongly constricted towards
base. Sutural ramp planar to slightly concave, steeply
sloping anteriorly from suture. Spire whorls with three
or four well-developed spiral cords anterior to shoulder.
Description: Estimated shell length to 50 mm; fusi¬
form, strongly constricted, inflection two-thirds distance
from last-whorl suture to anterior end. Shell verv thin
medially between shoulder and base. Protoconch
unknown. Teleoconch with at least three whorls. Spire
moderately elongate, sutures moderately impressed.
Sutural ramp planar, sloping anteriorly 45 degrees from
suture, with low flattened primary spiral cord at shoul¬
der. Axial sculpture of radial orthoeline growth lines,
bent adaperturally at suture. Spiral sculpture of about
17 primary spiral cords between base and shoulder,
posteriormost three cords flattened and weak to obso¬
lete. Sutural ramp smooth. Interspaces flat-bottomed,
wider than spiral cords at mid-whorl, lacking intercalated
secondary spiral cords. Spire whorls with smooth sutural
ramp, prominent shoulder, and three or four strong pri¬
mary spiral cords anterior to shoulder. Aperture elon¬
gate, ovate. Siphonal canal estimated at 20 percent of
shell length, narrow, straight or recurved abaxially.
Columellar and parietal callus absent. Siphonal fasciole
obsolete or weakly developed. Inner lip without plications.
Inside of outer lip smooth.
T>pe Material: PRI 2104, holotype, L (43.3), W 29.0;
PHI 2105, paratype, L (27.4), W 18.0.
Type Locality: Caleto Mero, northern Peru (Figure 1.
upper inset), early Miocene.
Distribution: Uppermost Oligocene to lower Mio¬
cene, Heath and lower Zorritos formations, northern
Peru. Lower Miocene, Posorja, Ecuador (Olsson, 1931).
Remarks: Specimens of Misifulgur sula are more
sharply constricted anteriorly than those of the Gulf
coast M. dockeryi and the latest Oligocene to early Mio¬
cene Peruvian M. macneili new species, have a wider
sutural platform and more prominent corded shoulder,
and have spire whorls with three or four well-developed
spiral cords anterior to tin1 shoulder, rather than the four
or five of M. dockeryi and six or seven of M. macneili.
The sutural platform is more steeply sloped anteriorly
than most specimens of M. cruziana , creating a more
fusiform and less turreted profile.
The holotype (PRI 2104) of Misifulgur sida has a
siphonal canal that is moderately recurved, unlike the
paratype (PRI 2105) and specimens assigned to other
species of Misifulgur , and growth lines on the last whorl
of the holotype trace a small sutural sinus that is absent
for other species of Misifulgur. New material from
northern Peru might indicate the need for a taxonomic
revision of M. sula.
Misifulgur macneili new species
(Figures 19, 20)
Diagnosis: Shell fusiform, moderately constricted towards
base. Sutural ramp narrow, planar, steeply sloping anteri¬
orly. Primary spiral cords well developed between base
and shoulder. Spire whorls with six or seven spiral cords.
Description: Estimated shell length to 70 mm, fusi¬
form, moderately constricted, inflection two-thirds dis¬
tance from last-whorl suture to anterior end. Shell thin
medially between shoulder and base. Protoconch unknown.
Teleoconch with at least five whorls. Spire moderately elon¬
gate, sutures moderately impressed. Sutural ramp narrow,
sloping anteriorly 60 degrees from suture, with low flat¬
tened spiral cord at poorly defined shoulder. Axial sculp¬
ture of radial orthoeline growth lines, bent adaperturally
at suture. Spiral sculpture of about 18 primary spiral cords
between base and shoulder, with anterior cords stronger
and separated by equally wide interspaces; posterior cords
weaker, flattened, and with interspaces half as wide as
cords. Some anterior interspaces with an intercalated sec¬
ondary spiral cord. Spire whorls with smooth, steeply
sloping sutural ramp, rounded and poorly defined shoul¬
der, and seven equally strong spiral cords anterior to shoul¬
der. Aperture oval. Siphonal canal estimated at 20 percent
of shell length, weakly recurved abaxially. Columellar
callus absent; parietal callus not visible. Siphonal fasciole
obsolete. Inside of outer lip smooth.
Type Material: UWBM 98670, holotype, B8376 (type
locality), L (44.8), W 27.6; UWBM 104229, paratype,
B8376, L (30.8); UWBM 104230, paratype, B8378,
L (47.2), W 32.0.
Type Locality: B8376, southwest of Cerros Colorados,
East Pisco Basin (Figure 1, middle inset), south-central
Peru.
Other Material Examined: UWBM 98671, B8358,
L (47.6), W (42.4); MUSM INV 200, B8358, L (49.2),
W (39.9); B8378, two fragments.
Distribution: Chattian to Burdigalian, Chilcatay depo-
sitional sequence. East Pisco Basin, south-central Pern.
Etymology: Named in memory of F. Stearns MacNeil,
posthumously a co-author with David T. Dockery of
a 1984 study of Oligocene mollusks from Mississippi
and author of the genus, Pseudofulgur.
T.J. DeVries, 2016
Page 107
Figures 15-30. Misifulgtir species. 15-18. Misifulgur sula (Olsson, 1931). Caleto Mero, Talara Basin, northern Peru . Early Miocene.
15. PRI 2104. I lolotype. Apertural view. Partially decorticated, partially an internal mold. Length 43.3 mm. 16. PHI 2104. Abapertural
view. 17. PRI 2105. Paratype. Apertural Mew. Length 27.4 mm. 18. PRI 2105. Abapertural Mew. Partially decorticated. 19, 20. Misifulgur
rmcrieili new species. East Pisco Basin, south-central Peru. Early Miocene. 19. UWBM 104230. B8378. Paratype. Abapertural Mew.
Siphonal canal and portion oflast whorl missing. Length 47.2 mm. 20. UWBM 98670. B8376. I lolotype. Abapertural Mew. Portion of
siphonal canal missing. Partially an internal mold. Length 44.8 mm. 21-30. Misifulgur cruziana (Olsson, 1932). 21. PRI 2307.
Holotype. Quebrada de la Cruz, Progreso Basin, northern Peru. Early Miocene. Apertural view. Anterior is missing. Length 28.2 mm.
22. UWBM 98656. B8316, East Pisco Basin, south-central Peru. Middle Miocene. Outer lip: fragment of smooth inside of posterior
portion (upper hall) and mold of spirally corded exterior of anterior portion (lower hall). Leading edge of outer lip at right. Length of
fragment + mold is 28.7 mm. 23. UWBM 98652. B8316. Abapertural view. Anterior end missing, largest visible whorl partially
decomposed/dissolved. Length 23.7 mm. 24. UWBM 98656. Abapertural view. Length 55.2 mm. 25. UWBM 98652. Close-up of
mammillate deviated protoconch. 26. UWBM 98656. Lateral Mew. Partially an internal mold at center left of image. 27. UWBM 98666.
B8371, East Pisco Basin, south-central Peru. Middle Miocene. Apertural view. Portion of siphonal canal missing, partially abraded by
wind-blown sand. Length 64.5 mm. 28. MUSM INV 194. B8313, East Pisco Basin, south-central Peru. Middle Miocene. Apertural view
of partly crushed specimen. Length 48.2 mm. 29. UWBM 98666. Lateral view. Partially an internal mold at left on spire. 30. UWBM
98654. B83I6. Abapertural Mew showing mammillate deviated protoconch. Anterior end missing. Anterionnost whorls decorticated or
with exposed internal mold. Length 19.3 mm.
Page 108
THE NAUTILUS, Vol. 130, No. 3
Remarks: Specimens of Misifulgur macneili new spe¬
cies are less constricted towards the base and have a
narrower sutural ramp than those of M. sulci, which oth¬
erwise is the species of Misifulgur that M. macneili most
resembles. The anterior and mid-whorl spiral cords on
both M. macneili and M. sula are less flattened, narrower,
and with wider interspaces than on specimens of
M. cruziana and M. montemarensis (see below).
At most localities, specimens of Misifulgur macneili
and M. cruziana occur together. The latter species is
highl y variable in terms of spire height, width and slope
of the sutural platform, and the onset of spiral cord
obsolescence on later whorls. Misifulgur macneili is not
thought to be a variant of M. cruziana for the following
reasons. First, specimens of M. macneili have six or
seven spiral cords anterior to the shoulder on spire
whorls; all variants of M. cruziana have three or four.
Second, the posterior spiral cords on specimens of
M. macneili flatten hut do not fade to obsolescence on
later whorls. On all variants of M. cruziana , some poste¬
rior spiral cords do disappear. Lastly, the shoulder on
specimens of M. macneili is barely angled, whereas
shoulders on the least tabulate variants of M. cruziana
have a distinct angulation at the shoulder that coincides
with a strong spiral cord.
Misifulgur macneili occurs at two closely situated sites
near the base of the Chilcatav deposition!] sequence in
coarse-grained sandstone and fine gravel (Figure 1, middle
inset). Specimens are part of a diverse mollusean assem¬
blage that includes gastropods overgrown with bryozoans
or small colonies of barnacles. The Mis ifulgu r- b ea ri ng
horizon is correlative with nearby oyster banks that lap
against outcrops of crystalline paleo-basement. A latest
Oligocene age is inferred from the stratigraphic position
of the Misifulgur specimens close to the transgressive
base of the Chileatay depositional sequence (DeVries,
1998). The youngest specimens were collected from
within the mid-section of the type section of the Chileatay
Formation. Their age is inferred to he about 18 Ma, based
on diatoms in overlying tuffaeeous and diatomaceous
sandstone (DeVries, 1998).
Misifulgur cruziana (Olsson, 1932)
(Figures 21-37)
Acanthiza ( Chorus ) sula cruziana Olsson, 1932: 185,
pi. 20, fig. 7.
(?) Chorus cruzianus (Olsson). — Marks, 1951: 23, 30,
also unpaginated figs 5, 7.
(?) Fasciola ria ? [sic] sp. Marks, 1951: 28, also unpag¬
inated fig. 7.
Diagnosis: Shell length to 120 mm. Sutural ramp
planar, sloping anteriorly five to 45 degrees from suture.
Primary spiral cord on shoulder strong or keeled on
large specimens.
Description: Estimated shell length to 120 mm,
fusiform in small specimens, moderately to strongly
constricted with inflection about 60 percent of distance
from last-whorl suture to anterior end; later whorls pos¬
teriorly inflated, deeply constricted, and strongly tabu¬
late in some larger specimens. Shell very thin medially
between shoulder and base. Protoconch mammillate,
smooth, at least two whorls, slightly deviated. Teleoconeh
with five whorls. Spire moderately elongate, sutures mod¬
erately impressed. Sutural ramp planar, sloping anteriorly
45 degrees from suture on small and medium-sized spec¬
imens; on large specimens, sutural ramp planar to con¬
cave, sloping anteriorly five to 45 degrees from suture.
Shoulder angulate, defined by rounded primary spiral
cord, keeled on some large specimens. Axial sculpture of
orthocline radial growth lines, often bent adaperturally at
suture. Earliest spire whorls with smooth sutural ramp
and three or four primary spiral cords anterior to shoul¬
der. Last whorl on small specimens with about 17 broad
primary spiral cords between base and shoulder, with
posteriormost three diminished, flattened, and more
widely spaced. Primary interspaces U-shaped to flat-
bottomed, becoming as wide as primary cords adapically
towards shoulder, rarely with an intercalated secondary
spiral cord. Either side of shoulder sometimes with closely
spaced subdued secondary spiral cords; sutural ramp with
primary spiral cords absent or with one spiral cord situ¬
ated medially or close to suture. Larger specimens with
broad smooth concavity bounded by shoulder and pri¬
mary spiral cord near midpoint of whorl. Anterior half of
last whorl on large specimens with about 12 spiral cords,
spaced more closely towards base. Aperture ovate and
elongate in small specimens, quadrate in large specimens.
Siphonal canal about 20 percent of shell length, recurved
abaxially, broadly open. Columellar or parietal callus
absent. Inner lip without plications. Siphonal fasciole
obsolete. Inside of outer lip smooth.
Type Material: PRI 2307, holotype, L (28.2), W 26.1.
Type Locality: Quebrada de La Cruz, northern Peru
(Figure 1, upper inset), early Miocene.
Other Material Examined: MUSM INV 190, B8316,
middle Miocene, L (29.7), W (28.8); MUSM INV 191,
B8316, L (29.2), W 17.7; MUSM INV 192, B8316,
L (28.9), W 28.4; MUSM INV 193, B8316, L 62.0,
W (47.1); MUSM INV 194, B8313, middle Miocene,
L (48.2), W 34.1; MUSM INV 195, B8305, earlv Miocene,
L (28.2), W (26.1); MUSM INV 196, B8371, middle
Miocene, L (54.4), W 40.0; MUSM INV 197, B8371,
L (43,5), W 31.3; MUSM INV 198, B8371, L (50.1),
W (53.4); MUSM INV 199, B8358, early Miocene,
L (39,5); UWBM 98652, B8316, L (23.7), W 19.0; UWBM
98653. B8316, L (29.4), W 28.0; UWBM 98654, B8316,
L (19.3), W 16.6; UWBM 98655, B8316, L 30.9,
W (19.9); UWBM 98656, B8316, L (53.0), W 33.6; UWBM
98657, B8316, I, (57.3), W 47,3; UWBM 98658, B8364,
middle Miocene, L (47.0), W 34.7; UWBM 98659,
B8362, middle Miocene, L (48.0), W (31.2); UWBM
98660, B8362, L (28.2), W (25,5); UWBM 98661,
B8372, middle Miocene, L (37.8), W (46,5); UWBM
98662, B8305, L (65,3), W (52.5); UWBM 98663,
T.J. DeVries, 2016
Page 109
Figures 31-37. Misifulgur species. 31-33. Misifulgur cruziana (Olsson, 1932). Large specimens from East Pisco Basin, south-central
Peru. Late Oligocene to early Miocene. 31. UWBM 104232, B8361. Abapertural Mew. Partially decorticated. Length 83.6 mm.
32. UWBM 104231, B8361. Abapertural view. Internal mold. Length 63.1 mm. 33. UWBM 104233, B8358. Abapertural Mew.
Rubber cast of external mold in concretion. Length 61.0 mm. 34-37. Misifulgur cruziana (Olsson. 1932). Molds from the Progreso
Basin of southwestern Ecuador (Marks, 1951). 34. PBI 68827, PBI locality 5098C. Progreso Formation, middle to lower Miocene.
Apertural Mew. Internal mold, most of siphonal canal missing. Image taken without ammonium chloride. Length 107 mm. 35. PRI 68827.
Abapertural view. 36. PRI 68826, PRI locality 4740C. Upper Tosagua Formation, lower Miocene. Apertural view. Internal mold,
anterior end missing. Image taken without ammonium chloride. Length 79 mm. 37. PRI 68826. Abapertural view.
BS305, L (34.6), W (26.4); UWBM 98664, B8376, early
Miocene, L (25.0), W (23.0); UWBM 98665, B8376,
L (24.7), W 18.0; UWBM 98666, B8371, L (64.5), W 46,3;
UWBM 98667, B8371, L (48.0); UWBM 98668, B8371,
L (69.4), W (55.6); UWBM 98669, B8358, L (48.1),
W (44.4); UWBM 98676, B8366, late Miocene, L (61.4),
W (42.8); UWBM 104231, B8361, early Miocene, L (63.1),
W (62.7); UWBM 104232, B8361, L (83.6), W (62,5);
UWBM 104233, B8358, L (61.0), W (57.0); UWBM
104234, B8363, late middle Miocene, L (78.9), W 65.3;
PRI 68826, L (79), W 64; PRI 68827, L (107), W 78. Also
B8359, B8365, B8369, B8370, B8333, B8374, B8375,
B8377, B8343, B8379.
Distribution: Aquitanian, Chil cat-ay depositional
sequence, south-central Peru; lower Zonitos/Heath forma¬
tions, northern Peru. Dos Bocas Member, Tosagua For¬
mation, Progreso Basin, southwestern Ecuador. Burdigalian
to early Tortonian, Chilcatay and Pisco depositional
sequences, south-central Peru. Tosagua and Progreso
formations, Progreso Basin, southwestern Ecuador.
Remarks: The incomplete holotype of Misifulgur
cruziana from northern Peru is identical with more com¬
plete medium-sized specimens from the East Pisco Basin.
Specimens of M. cruziana from south-central Pern
include examples with paucispiral, deviate, mammillate
protoconchs (Figures 23, 25, 30), a complete siphonal
canal (Figure 26), and highly variable specimens much
larger than the northern Peruvian holotype (Figures 31-
33), in which the last whorls may be broader, more
inflated posteriorly, and with keeled shoulders bounded
anteriorly and posteriorly by broad, shallow, smooth con¬
cavities (Figures 29, 33).
THE NAUTILUS, Vol. 130, No. 3
Page I 10
Misifulgur cruziana was the most common medium-
sized to large carnivorous gastropod on the Peruvian
continental shelf from the early to early late Miocene,
living far enough offshore for sardine scales to he
embedded in the sandy matrix, i.e., at water depths of
30 meters or greater (DeVries and Pearey, 1982). The
youngest specimen was found near the base of upper
Miocene outcrops at Cerro Blanco, in the Rio lea valley
near Oeueaje (UWBM 98676), where medium-grained
sandstone lies nonconformably upon a rugged basement
of crystalline basement rock (locality B8366). An ash bed
(DV 494-5Snee) 68 meters above the Misifulgur-bearing
bed yielded an 4(lAr-39Ar date of 6.85 ± 0.07 Ma
(L. Snce, United States Geological Survey, written com¬
munication, 1987), consistent with a late late Miocene
age indicated by diatoms from nearly the same horizon
(H. Schrader, University of Bergen, Norway, written
communication, 1988) and other 40Ar-39Ar ash dates from
correlative same strata (Brand et al., 2011).
A buccinid mold (PBI 68826; figs 36, 37) attributed by
Marks (1951) to Misifulgur cruziana (cited as Chorus
cruziana) was collected in southwestern Ecuador, 1.75 kilo¬
meters west of Carrizal (estimated locality: 02°06'S,
80°27'30"W, based on comparison of data obtained from
MapCarta and Google Earth). A second mold (PBI
68827; figs 34, 35), identified as Fasciolaria? sp. [sic] by
Marks (1951) but similar to Misifulgur molds from Peru
and Ecuador, was collected at IPG locality 11895 (P. Plocic,
Paleontological Research Institution, written communica¬
tion, 26 May 2016; additional locality data are unavail¬
able). According to Marks (1951), the Carrizal specimen
was collected from cross-bedded sandstones of inner
shelf origin from the upper Subibaja Formation, assigned
an earlv Miocene age based on correlations with mol-
lusks in Peru and foraminifera identified by R. M.
Stainforth (1948). The Carrizal locality is more recently
mapped into the lower Miocene Dos Bocas Member of
the Tosagua Formation (Longo and Baldock, 1982). The
“ Fasciolaria ” mold (PRI 68827), as well as additional
specimens of "Chorus cruziana' cited by Marks (1951),
were collected from the marine to brackish -water, mid¬
dle to upper Miocene Progreso Formation (Marks, 1951);
its age is supported by more recent authors (Longo
and Baldock, 1982). Both Ecuadorian molds have a
shape that falls within the broad range shown by
Peruvian specimens.
Figures 38-44. Misifulgur montemarensis new species. Late Miocene. 38. UWBM 98672, B8360. Holotype. Sacaco Basin,
southern Pern. Apertural view. Shell material partially dissolved, internal mold partially visible. Length 67.9 mm. 39. UWB.Vl
98672. Abapertural view. Most of spire missing. 40. UWBM 98674. B8373. Paratype. Sacaco Basin, southern Peru. Apertural view;
note mammillate protoconch. Shell material partially dissolved. Length 26.4 mm. 41. UWBM 98674. Abapertural view. Shell material
partially dissolved. 42. UWBM 98673, B8373. Paratype. Apertural view. Portion of siphonal canal and spire missing, partially an
internal mold. 43. UWBM 9867.5, B8359. East Pisco Basin, south-central Peru. Abapertural view. Portions ol last whorl missing.
Length 77.2 mm. 44. UWBM 98673. Abapertural view.
T. J. DeVries, 2016
Page 1 1 1
Misifulgur montemarensis new species
(Figures 38-44)
Diagnosis: Shell length to 1 10 mm, fusiform, weaklv
constricted towards base. Sutural ramp weakly planar,
sloping anteriorly 50 to 70 degrees from suture. Spiral
sculpture of about 16 very broad primary spiral cords
between base and shoulder, diminished in strength
near shoulder.
Description: Shell to 1 10 mm long, fusiform, weakly
constricted, inflection about 60 percent distance from
last-whorl suture to anterior end; shell very thin medi¬
ally. Protoconch mammillate, smooth, at least two
whorls, slightly deviated. Teleoeonch of four to five
whorls. Spire moderately elevated, sutures weaklv
impressed. Sutural ramp weakly planar, sloping anteri¬
orly 50 to 70 degrees from suture. Shoulder with small
inflection and weak primary spiral cord. Axial sculpture
of fine, closely spaced, orthoeline, radial growth lines.
Spiral sculpture on last whorl of about 16 broad, low,
primary spiral cords between base and shoulder,
diminishing in strength near shoulder. Primary interspaces
broadly U-shaped, narrower than primary spiral cords,
without intercalated spiral elements. Spire whorls with
sutural ramp and five primary spiral cords, one on sutural
ramp and three to four from shoulder anteriorward. Aper¬
ture elongate, ovate. Siphonal canal 20 percent of shell
length, narrow, open, nearly straight. Columellar and pari¬
etal callus absent or nearly so; inner lip without plications.
Siphonal fasciole obsolete. Inside of outer lip smooth.
Type Material: (all late Miocene): UWBM 98672,
B 83*60 (type locality), holotype, L (67.9), W 45.4;
UWBM 98673, B8373, paratype, L (34.6), W 23.2;
UWBM 98674, B8373, paratype, L 26.4, W (13.9).
Type Locality': B8360, eastern flank of Montemar
hills, between Lomas and Saeaco, southern Peru. Late
Miocene, about 6-8 Ma (Figure 1, lower inset).
Other Material Examined (all late Miocene):
MUSM INV 201, B8368, L 102.8, W (49.0); MUSM
INV 202, B8367, L (72.9), W (39.7); UWBM 98675,
B8359, L (77.2), W (42.2).
Distribution: Late Miocene, southern Peru.
Etymology: Named for the Montemar paleo-island
near Lomas, Peru, in the lee of which Misifulgur- bearing
upper Miocene tuffaceous sands were deposited.
Remarks: Specimens of Misifulgur montemarensis
have planar and steeply dipping sutural ramps and
poorly defined shoulders, even on the largest specimens.
Primary spiral cords on specimens of M. montemarensis
are broader and lower than those on specimens of the
round-shouldered early Miocene M. sula and have
narrower interspaces.
The types of Misifulgur montemarensis were found in
shoreface sandstone on the lee side of a granitic paleo-
island - Montemar. A late Miocene age is inferred from
8-Ma tnK- l0Ar-dated ash beds lower in the section (Muizon
and DeVries, 1985). Other specimens (MUSM INV 201,
MUSM INV 202) were found near Nazea on the lee side of
a paleo-peninsula (Cerro Huaricangana; see DeVries,
1988) in littoral sandstones 20 meters above an ash bed
(DV 528-1 Snee) with a late Miocene 4<lAr-'wAr date of
7.51 ± 0.05 Ma (L. Snee, written communications, 1987).
DISCUSSION
Misifulgur docken/i is the oldest species of its genus by
ten million years, so it might be that the Gulf of Mexico,
home to other Paleogene buecinoids (MacNeil and
Dockery, 1984), was home to the first Misifulgur. Bv the
latest Oligocene or earliest Miocene, when no physical
barrier had yet separated the Pacific and Atlantic Oceans
(Collins et ah, 1996), M. sula and M. cruziana had
become established in southwestern Ecuador, north¬
western Peru (Olsson, 1931, 1932), and for the latter
species, south-central Peru. A similar pattern of distribu¬
tion characterized the Caribbean genus, Muracypraea
Woodring, 1957, for which early Miocene northern and
southern Peruvian species constituted southern hemi¬
spheric outliers (DeVries et ah, 2006).
A case against dispersal and in favor of convergence of
the Miocene western South American species with an
unrelated Gulf Coast Oligocene species could be made
based on the absence of Central American records for
Misifulgur. The buccinids herein referred to Misifulgur ,
however, are rare in northern Peru (three specimens)
and southwestern Ecuador (two specimens), indicat¬
ing that their absence in Miocene deposits of Central
America may reflect a Miocene scarcity of individuals or
a modern scarcity of preserved paleo-habitat. The issue
of convergence versus dispersal will not be resolved
until Misifulgur is found in Central America or a plausi¬
ble Oligocene ancestral buceinid is found in western
South America.
Historical Biogeocraphical Context for Buccinids
on the Peruvian Shelf
The modern Peruvian Faunal Province (Dali, 1909;
Tarazona et ah, 2003), which includes the present-day
coastline of the East Pisco and Sacaco basins but not the
Talara nor Progreso basins, is inhabited at intertidal and
shallow subtidal depths by medium-sized carnivorous
murieid gastropods of the genera Acanthina, Concholepas
Lamarck, 1801, Stramonita Schumacher, 1817, and
Xanthochorus Fischer, 1884 (Alamo and Valdivieso, 1997;
Guzman et ah, 1998). At inner shelf depths, Stramonita
and Xanthochorus occur together with the carnivorous /
scavenging buceinid, Solenosteira fusifonnis and, off south¬
ern Pern, the buceinid, Aeneator fontainei , the former
with a distribution northward into the Panamic Faunal
Province and the latter with southern ocean affinities
(Laudien et ah, 2007; Araya, 2013; S.V. Mogollon, written
communication, 2008). On the Chilean shelf, gastropod
THE NAUTILUS, Vol. 130, No. 3
Page I 12
predators include members of the murieid genus Trophon
Montfort, 1810, the murieid Chorus giganteus (Lesson,
1831) (McLean and Andrade, 1982; Gajardo et ah, 2002),
and A. fontainei.
During the early to late Pliocene, soft-bottom substrates
of the Peruvian Faunal Province at comparable shallow
depths were occupied by the same genera of medium¬
sized muricids as would be present during the Quaternary,
although usually die genera were represented by
species now extinct (DeVries, 1995; 2000; 2003; 2005a).
Some murieid genera ( Concholepas , Chorus , Trophon ,
Xanthochoms ) had more expansive distributions, ranging
from Chile to Cabo Blanco, northern Peru (Herm, 1969;
DeVries, 1986; 1995; 1997; 2005b; 2007a).
From the latest Oligocene to the early late Miocene,
Misifulgur , especially M. cruziana, is inferred from its
abundance to have been the predominant medium-sized
to large carnivorous neogastropod on the Peruvian shelf.
Muricids were rare and small (DeVries, 2005b). A variety
of neogastropods inhabited Miocene shelf substrates of
Ecuador’s Progreso Basin, according to Marks (1951),
including Misifulgur, Conus spp., Terebra spp., and turrids,
but notably not muricids. During the late Miocene,
die youngest species of Misifulgur, M. montemarensis,
appeared at inner shelf depths along the southern Peruvian
coast, and dien, at about 6 Ma, became extinct, an event
approximately coinciding with die northward expanding
range of a similarly sized, shelf-dwelling, carnivorous neogas¬
tropod, the Chilean murieid, Chorus frassinetti DeVries,
1997, as well as the appearance of the smaller Peruvian
trophinines, Xanthochorus stephanicus DeVries, 2005
and X. ochuroma DeVries, 2005 (DeVries, 2005a).
CONCLUSIONS
Buccinids in Misifulgur new genus probably evolved along
die Gulf Coast of die United States during the Oligocene.
Misifulgur did not experience the same Neogene diversifi¬
cation in the Atlantic realm as other buceinid genera, but
before the latest Oligocene evidently passed into the trop¬
ical eastern Pacific Ocean, where populations found a
niche on soft substrates of the continental shelf from south¬
western Ecuador to southern Peru. Misifulgur cmziana
became the most abundant and perhaps largest carnivorous
gastropod inhabiting die southern Peruvian sandy inner
shelf during the earlv and middle Miocene. By the late
Micx'ene, only M. montemarensis new species remained,
living close to the shoreline on protected sandy substrates.
The demise of M. monteniarensis and thereby the extinc¬
tion of Misifulgur coincided widi the late Miocene to early
Pliocene advent and diversification of die morphologically
similar, shelf-dwelling, carnivorous murieid genus, Chorus ,
as well as the trophinine genus, Xanthochoms.
ACKNOWLEDGMENTS
I would like to thank David Dockery (Office of Geology,
Mississippi Department of Environmental Quality) for
the loan of Oligocene specimens from the Gulf Coast
and curators of the Paleontological Research Institution
(PRI), the Academy of Natural Science of Philadelphia,
and the United States National Museum of Natural
History for the loan of material. Staff at PRI (Greg Dietl,
Leslie Sldbinski, Page Plocic) were very accommodating
during a visit to Ithaca. Valentin Mogollon (Universidad
National Federico Villarreal, Lima, Peru) offered invalu¬
able information on the modern Peruvian molluscan
fauna. The manuscript was improved by constructive
suggestions from Anton Oleinik (Florida Atlantic Uni¬
versity, Boca Raton, Florida) and Sven Nielsen (Uni¬
versidad Austral, Valdivia, Chile).
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APPENDIX
Locality numbers of the Burke Museum of Natural His¬
tory and Culture (Bxxxx), and corresponding DeVries
locality-sample numbers (DV xxxx-xx). dep. seq. = depo-
sitional sequence.
B8305, DV 377-3
B8313, DV 420-1
B8316, DV 482-8
B8333, DV 1320-1
B8343, DV 3004-1
B8358, DV 395-12
B8359, DV 421a- 1
SW wall of Quebrada Gramonal,
near juncture with Rio lea; deltaic
foresets and bottom-sets of coarse¬
grained sandstone. 14°45'45" S,
75°30'27" W. Chilcatay dep. seq.,
upper lower Miocene.
S end of Filudo Depression; gray
sandstone beds. 13°53'12" S,
76° 07' 15" W. Pisco dep. seq.,
middle Miocene.
As for B8316, middle horizon of
fossils.
Pampa Colorado, along road to
Playa Penon, indurated red sand¬
stones. 14°25'59" S, 75° 49' 19" W.
Pisco dep. setp, lower upper
Miocene.
NW corner of Cerro Submarino,
above unconformity. 14°34,36" S,
75°39'55" W. Lower Pisco dep.
seq., middle Miocene.
Two km W of Cerro La Virgen,
north side of Comotrana-Carhuas
road. 14°10'04" S, 76°07'05" W.
Chilcatay dep. seq., lower Miocene.
N wall of Quebrada Huaricangana,
across from small canyons on S
wall. 14°57'08" S, 75°18,06" W.
Pisco dep. seq., upper Miocene.
T. J. DeVries, 2016
Page 1 15
B8360, DV 430-1
B8361, DV 441-14
B8362, DV 451-2
B8363, DV 482-7
B8364, DV 482-10
B8365, DV 484-6
B8366, DV 494-1
B8367, DV 528-3
B8368, DV 536-1
B8369, DV 573-3
Montemar, three km S of Lomas
intersection; shell-rich bed near
Panamerican Highway. 1 5°3 1 '08"
S, 74°48'43" W. Pisco dep. seep,
upper Miocene.
Lower part of measured type sec¬
tion of Chilcatay Formation, 90.8
meters above measured base,
"skull horizon." 14°1 1'31" S,
76°06,56" W. Lower Chilcatay
dep. seq., lower Miocene.
W side of Filudo depression;
anadarid-bearing coarse-grained
sandstone bed at 21 meters in
measured section. 13°57'19" S,
76°07'19" W. Lower Pisco dep.
seq., middle Miocene.
Bowl between Cerro La Brujita
and Cerro La Bruja on W side
Rio lea valley; lowest continuous
shell level, with large gastropods.
14°30'41" S, 75°40'00" W. Pisco
dep. seq., upper middle Miocene.
As for B8316, two meters above
middle shell bed.
Yesera de Amara, ledges and sand¬
stone horizons on lower NE slopes
of buttes. 14° 35' 38" S, 75°40,10" W.
Lower Chilcatay dep. seq., upper¬
most Oligocene to lower Miocene.
Cerro Blanco on W side Rio lea.
Mollusks 15 meters above base of
measured section, five meters
below diseonformity. 14°25,22" S,
75°41'12" W. Pisco dep. seq.,
upper Miocene.
S wall of Quebrada Huaricangana,
third gulch from W; 40 meters
above base of section near E ridge.
Shell bed with diverse fauna.
14°57'47" S, 75°16'58" W. Pisco
dep. seq., upper Miocene.
S wall of Quebrada Huaricangana,
25 meters above base of exposed
section; cross-bedded sandstone
with scattered in situ mollusks.
14°57'1 1" S, 75° 1 7'08" W. Pisco
dep. seq., upper Miocene.
Faulted slopes about 1-2 km E of
Fundo Santa Rosa, massive sand¬
stone with fish scales. 14°47,16" S,
75°30'22" W. Pisco dep. seq.,
middle Miocene.
B8370, DV 576-6
B8371, DV 579-1
B8372, DV 580-1
B8373, DV 1348-1
B8374, DV 1403-1
B8375, DV 1611-3
B8376, DV 1648-1
B8377, DV 2005-1
B8378, DV 2241-1
B8379. DV 4076-1
DV 494-5Snee
DV 528-1 Snee
Slopes E of Quebrada Gramonal.
14°45'38"S, 75°30,21"W. Pisco
dep. seq., lower upper Miocene.
South of Quebrada Gramonal,
one km E of double-knobbed hill
above Rio lea in faulted zone of
basement blocks and Turritella-
sandstone; bed just above marine
mammal level. 14°4630" S,
75°30'06" W. Pisco dep. seq.,
middle Miocene.
1.5 to two km ENE of B8371;
white bed with mollusks.
14°46'55" S, 75°29'28" W. Pisco
dep. seq., middle Miocene.
E side of Montemar, near
Panamerican Highway. 15° 3 I'M" S,
74°48'57" W. GPS. Pisco dep.
seq., upper Miocene.
Slope E of Cerro Sombrero,
5 meters above angular unconfor¬
mity. 14° 05' 49" S, 76°11'08" W.
Basal Chilcatay dep. seq., upper¬
most Oligocene.
Campsite at Ullujaya West, bowl¬
shaped depression'. 14°34'52" S,
75°38'40" W. Lower Chilcatay
dep. seq., uppermost Oligocene.
Valley SW of Cerro Colorado;
southwest-facing flank. 14°22'25"
S, 75°53/52" W. Chilcatay dep.
seq., lower Miocene.
SE side of Laguna Seea, near top
of hill, above tnffaceous white
horizon. 1 4°33'28" S, 75°32'46"
W. Pisco dep. seq., lower upper
Miocene.
SE of Cerros Colorados, edge of
tilted surface of alluvial boulders
with W source. 14°22/46" S,
75°53'50" W. Lower Chilcatay
dep. seq., uppermost Oligocene
to early Miocene.
SE Cerros Colorado, above angu¬
lar unconformity. 14°22'55" ‘ S,
75°53'52" W. Lower Chilcatay
dep. seq., uppermost Oligocene.
As for B8366, but 82 m above base
of measured section, ash bed,
6.85 ± 0.07 Ma. No Burke local¬
ity number.
As for B8367, but 20 m above base
of measured section, ash bed,
7.51 ± 0.05 Ma. No Burke local¬
ity number.
THE NAUTILUS 130(3): 1 16-121 , 2016
Page 1 1 6
A new species of Achnete (Gastropoda: Cancellariidae: Admetinae)
from the Paleocene of eastern Hokkaido, northern Japan
Kazutaka Ainano
Department of Geoscience
Joetsu University of Education
1 Yamayashiki
|oetsu 943-8512, JAPAN
Anton Oleinik
Department of Geosciences
Florida Atlantic University
777 Glades Road
Boca Raton. FL 33431 USA
Robert G. Jenkins
School of Natural System
College of Science and Engineering
Kanazawa University
Kanazawa City, Ishikawa 920-1 192, JAPAN
ABSTRACT
A new cancellariid species, Achnete katsuhiraensis new species is
described from the Paleocene Katsuhira Formation in Urahoro
Town, eastern Hokkaido, Japan. This is the oldest record of
cancellariid gastropods in Japan and also the oldest record of
the genus Achnete and the subfamily Admetinae worldwide.
Although the known fossil record of the genus is very incom¬
plete, occurrence of Paleocene Admetinae in deep sea deposits
of the Katsuhira Formation in northern Japan provides an
interesting insight into the timing of origin of the present day
cold water mollusean fauna in the northern Hemisphere.
INTRODUCTION
Cancellariids consist of a diverse group of mostly warm-
water mollusks. Biodiversity estimates for Cancellariidae
include 1200 fossil and 300 recent species (Hemmen,
2007), and 124 genera and 1864 recent and fossil species,
according to Petit and Harasewych (2005). The oldest
geological records of the family date from the Late Cre¬
taceous (Cenomanian) of Texas and the family reached
its maximum diversity in the Eocene and Miocene
(Stephensen, 1952; Taylor and Morris, 1988; Hemmen,
2007). No Cretaceous cancellariids have been recorded
from Japan (see Hayami and Kase, 1977; Kase, 2001).
The oldest species from Japan are Cancellaria ? sp. indet.
a, b, c, d from the upper Eocene Kvuragi and Kijima
Formations and the lower Oligoeene Yamaga Formation
in northern Kyushu (Nagao, 1928). However, columellar
folding, one of the distinct characteristics of cancellariids,
has not been recognized on those Eocene species from
Kyushu. These Paleogene species from Kyushu were
reexamined by Oyama et al. (1960), and C.? sp. indet. a,
b from the upper Oligoeene Yamaga Formation were
reassigned to Trigonostoma ? ( Scalptia ?) sp. No Paleo¬
gene cancellariids have been recorded in Sakhalin, while
several cancellariid species have been described and
illustrated from the Paleogene deposits of western
Kamchatka (Gladenkov et ah, 1991).
Specimens of a small cancellariid have been recently
collected from the Paleocene (Danian-Selandian)
Katsuhira Formation (see Amano and Jenkins, 2014) in
eastern Hokkaido. In this paper, we describe these as a
new species and discuss its biogeographic significance.
MATERIALS AND METHODS
Two specimens of cancellariid gastropods were collected
from dark gray mudstone of the upper part of Katsuhira
Formation at the cliff along Urahoro River near Katsuhira,
Urahoro Town, eastern Hokkaido (Figure 1). The age of
the formation was assigned to the Paleocene (Danian-
Selandian), based on the planktonic foraminifera and
calcareous nannofossils (Kiminami et ah, 1978; Kaiho,
1984; see also Amano and Jenkins, 2014).
Cancellariid specimens from this locality are associated
with protobranch bivalves such as Acilci , Leionucula,
Malletia, and deep-sea arcid Bentharca steffeni Amano,
Jenkins and Nishida, 2015. Although paleoenvironments
of the Katsuhira Formation have not been studied
in detail, these bivalves indicate deposition in deep
water (Amano and Jenkins, 2014; Amano and Oleinik,
2014; Amano et ah, 2015). One institutional acronym
used is: JUE, Joetsu University of Education, Joetsu,
Niigata Prefecture.
SYSTEMATIC PALEONTOLOGY
Class Gastropoda Cuvier, 1797
Order Neogastropoda Wenz, 1938
Superfamily Cancellarioidea Forbes and Hanley, 1851
Family Cancellariidae Forbes and Hanley, 1851
Subfamily Admetinae Troschel, 1865
Remarks: Admetinae is characterized by thin shell
and columella with arched and weak columellar folds
(Wilson, 1994). Recent molecular phylogeny work
reveals that this subfamily, as it is now understood, is
polyphyletic (Modica et al., 2011). The genus Admetula
K. Amano et al., 2016
Page 117
published by the Geospatial Information Authority of Japan).
Cossmann, 1899 is considered as a separate from
Admetinae clade. This classification was used by
Harzhauser and Landau (2012) when they revised the
Neogene cancellariids of the Paratethys. They also
treated Bonellitia |ousseaume, 1887 as the AcLmetula
clade, despite of lack of molecular data.
Genus Admete Kroyer in MoIIer, 1842
Type Species: Admete crispa Moller, 1842 (= Tritonium
viridulurn Fabricius, 1780) by monotypy.
Remarks: The genus Admete is characterized by a
rather thin shell with large last whorl, deep sutural
groove, narrow shoulder and by having a straight
collumella with two weak folds and narrow umbilicus
or slit (Harasewych and Petit, 1986: 86; this study).
Neadmete Habe, 1961 can be separated from the genus
Admete by having a higher spire and rather straight
collumella with three folds. Z eadmete Finlay, 1926 dif¬
fers from Admete by a fine cancellate sculpture on entire
surface. Genera Admetula and Bonellitia have a thick
crenulated outer lip, three strong columellar folds and
lack umbilicus or slit. The American Cretaceous genus
Admetopsis Meek, 1873 including Admete ? gregaria
Meek, 1873, A.? simfusifomiis and A.? rhomboides Meek,
1873 can be clearly separated from Admete by having
rounded whorls without shoulder, a significantly higher
spire, an anteriorly thickened calcareous callus, by
weaker or absent of collumellar folds, and lack of an
umbilicus or slit. It is presently not clear if Admetopsis
belongs to Admetinae or not. Another American Creta¬
ceous genus Paladmete Gardner, 1916 can be easily dis¬
tinguished from Admete by having a smooth collumella
without folds. Stephenson (1941) established family
Paladmetidae based on this genus.
Admete katsuhiraensis new species
(Japanese vernacular name: Katsuhira-koromogai)
(Figures 2, 3)
Diagnosis: Shell small with cancellate sculpture
consisting of 15 to 21 spiral cords and 19 to 23 axial ribs.
Page 118
THE NAUTILUS, Vol. 130, No. 3
Figures 2, 3. Admete katsuhiraensis new species. 2. Paratype, |UE no. 15940; 2a, adapertnral view; 2b, apical view; 2c, apertural
view. 3. Holotype, JUE no. 15939; 3a, apical view, 3b. adapertural view; 3c, apertural view.
Four whorls with rather deep sutural groove; spire low,
1/8 of shell height. Columella nearly straight with two
weak folds; inner lip broadly covered by thin calcareous
callus; siphonal canal short with very' weak fasciole and
very' narrow umbilical slit.
Description: Shell small, attaining 1 1.4 mm in height,
thin, fusiform, with four whorls. Sutural groove rather
deep; very narrow shoulder present in holotype. Last
whorl large, occupying approximately 7/8 of the height
of shell; spire very low; protoconch poorly preserved,
half of smooth volution remaining. Surface of last whorl
sculptured by 19 to 23 rounded axial ribs that become
obsolete toward base, separated by equal or narrower
interspaces, more distinct near suture. Spiral sculpture
of last whorl consisting of 15 to 21 flat cords sometimes
with one weak cord in between. Sculpture of penulti¬
mate whorl consisting of 20 fine axial ribs and five spiral
cords in holotype. Outer lip thin and not crenulated;
columella nearly straight with two weak folds; inner lip
broadly covered by thin calcareous callus; siphonal canal
short with very weak fasciole and very narrow umbilical
slit recognized in paratype specimen.
Type Material: Holotype, JUE no. 15939 (Shell
height, 8.0 mm; Diameter, 6.0 mm); Paratype, [UE
no. 15940 (Shell height, 1 1.4 mm; Diameter, 9.9 mm)
Type Locality: The cliff along U rahoro River at 750 m
downstream from the mouth of Katsuhirazawa River,
U rahoro Town, eastern Hokkaido. Danian to Selandian
Katsuhira Formation.
Remarks: Admete viridula (Fabricius, 1780) is the only
species of Admete recorded from the upper Miocene to
Pleistocene deposits in Japan as A. couthouyi (Jay, 1839)
(Amano, 1983; Baba, 1990). According to Sneli and
Stockland (1986), the latter name is a junior synonym of
A. viridula. This species differs from A. katsuhiraensis
new species by its larger size (more than 20 mm in
height), a rounded shoulder, very faint columellar plaits
and more strongly curved columella.
Admete katsuhiraensis new species is similar to
Admete profundicola (Okutani, 1964) which lives at
1500 m depth off Sagami Bay, central Honshu. Both
species share small shell size (A. profundicola-, shell
height, 7.8 mm; diameter, 5.0 mm), angulated shoulder.
K. Amano et a]., 2016
Page 1 19
very narrow umbilicus, two columellar folds and similar
number of spiral cords (15 in the last whorl of
A. profundicola). The new species is different from
A. profundicola in having a lower spire and more numer¬
ous axial ribs (17-19 in the last whorl of A. profundicola).
The new species resembles A. californica Dali, 1908
(found around 1103 meters in the Gulf of California) by
having a similar number of spiral cords and axial ribs
(both 20 in the last whorl of A. californica). However,
the new species can be separated from A. californica by
having a larger shell (16 mm in height of A. californica),
lower spire and very narrow umbilical split
The new species differs from the Miocene
A. kamt.schatica Sinelnikova in Gladenkov and Sinelnikova,
1990 from the Ilyinskaya Formation, western Kamchatka
by its smaller size, more compressed shell, with less
whorls, a more inflated and larger (7/8 of the total shell
height) last whorl, and a deep sutural groove.
Cancellaria globulosa Holzapfel, 1888, from the
Cretaceous of western Germany, has a similar outline of
shell and two columellar folds. However, it differs from
the new species by its thick shell with multiple crenula-
tions inside the outer lip, lack of a deep suture, and lack
of an umbilical slit.
Etymology: The new species is named for the location
where the tvpe material was collected.
Distribution: Known only from the type locality,
Paleocene Katsuhira Formation, Danian to Selandian,
Urahoro Town, eastern Hokkaido.
DISCUSSION
The oldest fossil specimens identified as Admete ? were
recorded from the Cretaceous in North America. How¬
ever, as noted above, these species belong to a distinct
genus, Admetopsis. Admete ( Bonellitia ) funigera described
by Staadt in Cossmann, 1913 (202-203, pi. 7, figs 212-9),
from the Paleocene of France, possesses strong collumella
folds, and lack a sutural groove and umbilical slit. These
morphologic features suggest that A. funigera should be
classified as Bonellitia as was originally proposed by the
author. Kollmann and Peel (1983: 93, fig. 209) described
and figured an Admete from the Paleocene (Selandian)
Sonja Member of the Agatdal Formation in central
Nugssuaq of western Greenland. Schnetler and Petit
(2010: 22) reexamined these gastropods and allocated that
species to Eocantharus Clark, 1938 (family Buccinidae).
One more species of Admete was recorded from the
Paleocene (early Selandian) of Greensand at Sundkrogen,
Copenhagen, Denmark. The following species from these
deposits were described as Cancellaria by von Koenen
(1885: 8-12): C. latesulcata, C. conoidea, C. tricarinata,
and C. curta-, Ravn (1939) placed them in “ Admete ?.”
However, the former three species were later allocated
respectively to Admetula, Kroisbachia and Brocchinia by
Schnetler (2001). Although that author eliminated the
question mark from A.? curta , it is difficult to decide
whether to allocate it to Admete because of its very small
shell size (3.7 mm; Ravn, 1939: 86, pi. 3, figs 20a, b) and
relatively large protoconch, indicative of a juvenile indi¬
vidual. The genera Kroisbachia, Brocchinia , Admetula ,
and Unitas are known from the lower Paleocene
(Luzanovka beds) of Ukraine (Makarenko, 1976).
Bonellitia ( Admetula ) paucivaricata (Gabb, 1864) was
illustrated from the Paleocene (Danian) Getkilninskaya
Formation by Gladenkov et al. (1997: pi. 33, fig. 33).
Admete omata Ilyina, 1955 was described from the
Paleogene of the Ustiurt region (vicinity of the Aral Sea)
(Ilyina, 1955: 78, pi. 30, fig. 16). However, that species
has strong axial ribs and lack sutural groove which is not
characteristic of the genus Admete. Although the family
Cancellariidae is present in the Paleocene deposits of
Europe, Greenland and Kamchatka, no proven records
of the subfamily Admetinae are known to date from
Paleogene deposits anywhere in the world. Thus, Admete
katsuhiraensis new species from the Danian to Selandian
Katsuhira Formation is not only the oldest record of
cancellariid in Japan, but also constitutes the oldest record
of the genus Admete and the subfamily Admetinae.
The genus Admete Kroyer in Moller, 1842, today is
restricted to cold waters and considered to be an Arcto-
Boreal taxon (Kantor and Sysoev, 2006; Thorson, 1944;
Macpherson, 1971; Golikov and Scarlato, 1977; Golikov,
1995; Golikov and Sirenko, 1998, 2004; Gulbin, 2004).
Occurrences of the genus Admete in the Southern Hemi¬
sphere and subantarctie waters (Powell, 1951, 1958;
Knudsen, 1964) have yet to be investigated. Some of these
have been already reassigned to different genera, the tax¬
onomic status of other species have to be adjusted after
the study of soft parts morphology. Arctic and Antarctic
mollusks are well known for the convergence in shell
morphology, which makes their identification based on
the slit'll morphology alone uncertain at best.
Paleocene marine isotopic records do not indicate a
significant departure in sea-surface temperature values
from the Late Cretaceous and do not indicate the exis¬
tence of a significant thermocline (Bralower et al., 2002;
Dutton et al., 2005), but there are some indications of
warm global temperatures (Adatte et al., 2002) during
the early Paleocene. These paleoceanographic condi¬
tions, coupled with continuous extended shallow shelf
around the rim of the North Pacific, facilitated the dis¬
persal of molluscan faunas, which resulted in general
similarity between the northeastern and northwestern
Pacific Paleocene molluscan faunas. The majority of
faunas appears to be of warm to warm-temperate affini¬
ties (Oleinik, 2001), lacking cold-water taxa found in the
North Pacific today. Fauna of the Katsuhira Formation,
which was previously virtually unknown, shows a depar¬
ture from this general pattern in having a verv high degree
of endemism and the appearance of new genera, such as
the bueeinid Urahorosphaera (Amano and Oleinik, 2014),
new species such as the aporrhaid, Ka rigid i opt era inouei
(Amano and Jenkins, 2014), or the first appearance of the
modern day cold water genus Admete described herein.
Since there are no true Paleocene Admete species or even
Page 120
THE NAUTILUS, Vol. 130, No. 3
subfamily Admetinae records anywhere in the northern
hemisphere, the occurrence of Admete katsuhiraensis
new species has interesting implications on the history of
the formation of cold water molluscan faunas known today
in the Arctic and boreal regions. As noted above, the lithol¬
ogy and faunal assemblage of the Katsuhira Formation,
from which Admete katsuhiraensis new species was found,
are indicative of relatively deep water deposits. That not
only makes this locality unique among the North Pacific
shallow-marine Paleocene deposits, but also may suggest a
deep water origin of some modem day Arcto- Boreal taxa
in the early part of the Paleogene in the North Pacific.
ACKNOWLEDGM ENTS
This study was partly supported by a Grant-in-aid for
Scientific Research from the Japan Society for Promotion
of Science (C, 26400500, 2014—2016) to KA and RGJ.
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THE NAUTILUS 1 30(3): 122-126, 2016
Page 122
Hesperaptyxis , a new genus for some western American
Fasciolariidae (Gastropoda), with the description of a new species
Martin Avery Snyder
Academy of Natural Sciences of Drexel University
Philadelphia, PA 19103 USA
and
Museum National d’Histoire Naturelle
Paris, FRANCE
[email protected]
Geerat J. Venneij
University7 of California at Davis
One Shield Avenue
Davis, CA 95616 USA
[email protected]
ABSTRACT
Hesperaptyxis is proposed as a new genus for Turbinella
cinerea Reeve, 1847 and five other species from southern
California and northwestern Mexico, including the new species
H. negusi. It differs from Aptyxis Troschel, 1868 (Late Miocene
to Recent, Mediterranean and temperate eastern Atlantic) by
having a slightly upturned siphon and completely adherent
inner lip. It differs from the Late Miocene western Atlantic
genus Pseudaptyxis Petuch, 1988 by much more prominent
axial and spiral sculpture and by lacking adapical and abapical
sinuses on the outer lip. Hesperaptyxis is so far known fossil
only from the Pliocene and Pleistocene.
Additional Keywords: Mollusca, Miocene, Pliocene, Pleisto¬
cene, recent, Baja California
INTRODUCTION
The classification of members of Fusinus Rafinesque,
1815 and related genera has long presented problems.
Until recently, most species have been placed in the
overly broad genus Fusinus. It has become clear, how¬
ever, that although Fusinus and related genera form
a elade, which Couto and colleagues (2016) call the
Fusinus coins elade, there is substantial diversity in this
group. Most attention has been focused on large-shelled
species, leaving small-shelled taxa underrepresented in
most recent studies. Russo (2015) treated the taxon
Aptyxis Troschel, 1868, a member of the F. coins elade
(Couto et c il. , 2016), assigning to it Murex syracusanus
Linnaeus, 1758 (the type species, from the Mediterra¬
nean Sea and adjacent warm-temperate eastern Atlantic)
and the Californian Fusus luteopictus Dali, 1877. Russo
(2015) failed to note that Abbott (1954: 244) had previ¬
ously assigned Fusus luteopictus to Aptyxis, and Keen
(1958: 617-618) had assigned two additional eastern
Pacific species, Turbinella cinerea Reeve, 1847 and
Fusinus felipensis Lowe, 1935, to Aptyxis. Keen (1971)
left the somewhat larger eastern Pacific Fusinus
fredbakeri Lowe, 1935, in Fusinus sensu lato. No ade¬
quate description of Aptyxis has appeared to date, render¬
ing generic assignments arbitrary at best. Our purpose
in this article is to characterize these small species,
to propose the new genus Hesperaptyxis for the small
eastern Pacific species, to describe the new species
//. negusi from Baja California, and to discuss the
biogeographic conundrum presented by small northem-
hemisphere fusinine fasciolariids. An institutional acro¬
nyms is: ANSP, Academy of Natural Sciences of Drexel
University, Philadelphia, PA.
SYSTEMATICS
Family Fasciolariidae Gray, 1853
Subfamily Fusininae Wrigley, 1927
Genus Hesperaptyxis new genus
Type Species: Turbinella cinerea Reeve, 1847, desig¬
nated herein (Recent, northwestern Mexico; Pliocene, Isla
Carmen [Gulf of California, Baja California Sur State],
Mexico; Pleistocene, Isla Coronados [Gulf of California
Baja California Sur state], Mexico) ( Figures 1 , 2, 22)
Diagnosis: Small fusinine fasciolariids with angulated
shoulder cord, convex outer lip, distinct abapical and
adapical sinuses on the outer lip, and siphon with slightly
upturned tip and inner lip adherent throughout its length.
Description: Shell small, maximum length about
70 mm, fusiform. Protoconch paucispirai. Teleoconch
with up to 8 whorls. Suture accentuated bv steep sub-
sutural slope on all whorls. Teleoconch sculpture con¬
sisting of strong spiral cords and axial ribs. Shoulder
cord angulated. Central cord at adapical end of base, less
prominent. Base strongly constricted. Spiral cords pres¬
ent on entire siphon. Tip of siphon slightly upturned
(Figure 22). Aperture elongate-ovate. Outer lip crenu-
lated at edge, smooth or weakly lirate within. Inner lip
M.A. Snyder and G. J. Vermeij, 2016
Page 123
Figures 1-22. 1, 2. Hesperaptyxis cinereus (Reeve, 1847). ANSP 466447, off La Paz, Gulf ol California, Baja California Sur,
Mexico, 1992, 28.3 mm. 3, 4. Hesperaptyxis ambustus (Gould, 1853). ANSP 466448, off Guaymas, Gulf of California, Sonora,
Mexico, 1955, 54.9 mm. 5, 6. Hesperaptyxis felipensis (Lowe, 1935). ANSP 466449, on underside of rocks as low tide, Puertecitos,
Gulf of California, Baja California Norte, Mexico, 1985, 13.9 nun. 7, 8. Hesperaptyxis fredbakeri (Lowe, 1935). 36.0 mm, live
from Cholla Bay, Puerto Penasco, Gulf of California, Sonora, Mexico, ANSP 466450. 9-15. Hesperaptyxis lateopictus (Dali, 1877).
9—11. USNM 32350, Monterey, California., 17.5 mm. 12-15. On rocks by Scuba at 6-9 m, Santa Cruz Island off Santa Barbara,
California, 21.0 mm, 25.1 mm, 16-19. Hesperaptyxis negusi new species. 16-17. Holotype, ANSP 466446, live on rocks at 12-24 m,
San Bonito Island [west of Cedros Island], Pacific Baja California, Mexico, 28.6 nun. 18-19. Paratype, ANSP 466445, from type
locality, 19.1 mm. 20. Aptyxis syracusanus (Linnaeus, 1758). ANSP 466452, off southern Sicily, 1955 m, 46.6 nun. 21. Pseudaptyxis
santamariae Petuch, 1988. ANSP 52871, late Miocene (Turtonian) St. Marys Formation, Maryland, 26 mm. 22. Hesperaptyxis
cinereus (Reeve, 1847). ANSP 466447, 28.3 mm, off La Paz, Gulf of California, Baja California Sur, Mexico, 1992, anterior part of
shell, 28.3 mm.
THE NAUTILUS, Vol. 130, No. 3
Page 124
smooth, columellar margin resorbed in posterior third,
eallused and slightly produced in anterior third, with
transition between (Figure 22). Single entrance fold at
base of siphonal canal, shallow abapical sinus present on
outer lip, aperture slightly folded at terminus of suture.
Included Species: Turbinella cinerea Reeve, 1847
(Figures 1, 2); Fusus ambustus Gould, 1853 (Figures 3, 4);
Fusions felipensis Lowe, 1935 (Figures 5, 6); Fusinus
fredbakeri Lowe, 1935 (Figures 7, 8); Fusus luteopictus
Dali, 1877 (Figures 9—1 5); Hesperaptyxis negusi new spe¬
cies (Figures 16-19). (The upturned canal tip was previ¬
ously noted for Fusinus fredbakeri by Hertz et al. (1999:
80, table 2): [canal] “tip bent to left' .)
Etymology: Combination of Hesperia (Greek, land to
the west) and Aptijxis.
Comparisons: In shell characters, Hesperaptyxis is
extremely similar to Aptijxis Troschel, 1868, type species
Murex syracusanus Linnaeus, 1758, from the Mediterra¬
nean Sea and adjacent warm -temperate eastern Atlantic.
Although there have been occasional placements of addi¬
tional Mediterranean species in Aptijxis, we are basing
our comparison on the type species, Murex syracusanus
(Figure 20). The genus Aptijxis differs from Hesperaptyxis
by having a straight siphon without upturned tip and by
having the abapical end of the inner lip, at the distal end
of the siphonal canal, forming a detached edge to the left
margin of the canal. This occurs in roughly 50% of mature
specimens (n=12).
Another similar genus is Pseudaptyxis Petuch, 1988,
based on P. santamariae Petuch, 1988 from the Late
Miocene (Tortonian) St. Mary’s Formation of Maryland.
Our examination of the holotype of P. santamariae
(ANSP 52871) (Figure 21) shows that this small species
(length about 25 mm) differs from both Aptyxis and
Hesperaptyxis by much finer spiral and axial sculpture
and by having a straight outer lip without abapical and
adapical sinuses. The outer lip is smooth within, and a
parietal tooth or ridge is absent. Pseudaptyxis resembles
Hesperaptyxis in having the tip of the siphon slightly
upturned. In this respect, Hesperaptyxis and Pseudaptyxis
differ from the eastern Atlantic Aptyxis. In addition, the
shoulder is rounded in Pseudaptyxis, whereas it is more or
less angulate in Aptyxis and Hesperaptyxis.
Hesperaptyxis negusi new species
(Figures 16-19)
Description: Shell small, slender, to about 29 mm,
with short siphonal process. Protoconch broken or miss¬
ing in all specimens examined. Teleoconch of 8 convex
whorls at maturity, constricted at sutures. Axial sculpture
of about 12 ribs on early and penultimate whorl, 18 on
body whorl, morphing on last whorl to weak axial ridges
toward lip. Ribs stop short of suture posteriorly and
anteriorly. Spiral cords on entire teleoconch, to tip of
neck, about 5 on early whorls, 6 on penultimate whorl
with intercalated weaker cords; spiral sculpture weaker
on body whorl. Anterior sutural ramp steep with strong
cord at base, 3-4 close-set cords on ramp. Rounded
knobs where spiral cords cross axial ribs, evanescent on
second half of body whorl. Siphonal process short,
straight, with slightly upturned tip. Aperture elongate-
ovate, glazed white within with raised cords terminating
short of lip; small entrance fold at base of siphonal canal.
Parietal callus resorbed, blending smoothly into exterior
shell surface. Columellar margin of siphonal canal raised.
Operculum typical of genus, thin, light caramel color.
Shell background color pale to light brown. Spiral cords
dark brown between axial ribs, cream color at knobs
where crossing axial ribs. Remnants of one protoconch
suggest a dark caramel color.
Type Material: Holotype (Figures 16-17): ANSP
466446, length 28.6 mm, live on rocks at 12-24 m, 1989;
Paratvpe (Figures 18-19), length 19.1 mm, live on rocks
at 12-24 m, ANSP 466445; Both from type locality.
Type Locality: San Benito Island [west of Cedros
Island], Baja California State, Mexico.
Etymology: Named for Rick Negus, who recognized
the taxon as an unnamed species and donated the holo¬
type and paratvpe.
Distribution: From Santa Cruz Island south to San
Benito Island.
Remarks: Hesperaptyxis negusi is most closely related
to II. luteopictus, being of similar size and shape, and
sharing the same or similar habitats. It is easily distin¬
guished from II luteopictus by its more slender and
elongated profile and its less prominent sculpture.
II luteopictus is purplish brown with white spiral
bands where prominent centrally placed cords cross
raised portions of axial ribs, forming sharply pointed
knobs. II. negusi has a smoother more rounded sur¬
face sculpture.
BIOGEOGRAPHY
Despite their great similarity in shell characters, species
of the eastern Atlantic Aptyxis and eastern Pacific
Hesperaptyxis are separated by a large geographic gap.
Aptyxis has been part of the southern European and
eastern Atlantic fauna since at least the Late Miocene
and Pliocene (see Landau et ah, 2013). Hesperaptyxis is
known only from fossils from the Pliocene and Pleisto¬
cene of the Gulf of California, with records of //. cinereus
from Isla Carmen (Pliocene; Durham, 1950) and I si a
Coronados (Pleistocene; Emerson and Hertlein, 1964).
No similar species are known from the rich Neogene
faunas of the Caribbean region. Petuch (1988) described
Pseudaptyxis for the single species P. santamariae from
the Late Miocene of Maryland. Here again, no similar
M.A. Snyder and G.J. Vermeij, 2016
Page 125
taxon is known that might connect this genus phylogenet-
ically with either Aptyxis or Hesperaptyxis . These biogeo¬
graphic separation corroborates a taxonomic separation
among these three very similar genera.
The restriction of Hesperaptyxis to the coasts of south¬
ern California and northwestern Mexico has parallels
in other molluscan genera. The closest parallel is with
the ocenebrine muricid genus Mexacanthina Marko and
Vermeij, 1999, whose three species are common inter¬
tidal gastropods on rocky shores of the Pacific side
of Baja California and the Gulf of California. Like
Hesperaptyxis, Mexacanthina has no obvious relatives in
the Neogene or Recent fauna either in western North
America or elsewhere. It converges in form on the phy-
logenetically distinct eastern Atlantic genus Spinucella
Vermeij, 1993 (Early Miocene to Late Pleistocene).
Another geographically restricted genus in the living
fauna is Macron H. and A. Adams, 1853, with a modern
distribution coincident with that of Hesperaptyxis and
Mexacanthina. In this case, however, there are fossil spe¬
cies in the Early Miocene of Venezuela (Cibson-Smith
et ah, 1997) and the Miocene of Chile (Nielsen and
Frassinetti, 2003). Other examples are Megastraea
McLean, 1970 (Turbinidae), Liocerithium Tryon, 1887
(Cerithiidae), Myrakeena Harry, 1985 (Ostreidae) and
Austrotrophon Dali, 1902 (Muricidae: Ocenebrinae).
Hypotheses for how these restricted distributions came
to be have not been offered, but it is clear from these
examples and from Hesperaptyxis that the region from
southern California to the subtropical coasts of the Gulf
of California is geographically unique.
ACKNOWLEDGMENTS
The authors thank Rick Negus for providing the original
material which stimulated an examination of the small-
shelled west American Fasciolariidae. We also thank Paul
Callomon, collections manager at ANSP, for photograph¬
ing the specimens and assembling the plates, as well as
for helpful discussions and observations.
LITERATURE CITED
Abbott, R.T. 1954. American seashells: a guide to the shells of
the Atlantic, Pacific and Gulf shores of the United States,
Canada, Central America and the islands of the Caribbean.
D. van Nostrand Company, New York, xiv, 541 pp.
Adams, H. and A. Adams. 1853-1858. The genera of Recent
Mollusca; arranged according to their organization. John
Van Voorst, London. 2 vols. 1853: 1: 1-256, pis. 1-32.
Couto, D.R., P. Bouchet. Y. I. Kantor, L.R.L. Simone and
G. Giribet. 2016. A multilocus molecular phytogeny of
Fasciolariidae (Neogastropoda: Buccinoidea). Molecular
Phylogenetics and Evolution, 99: 309-322, figs 1-5, 2 tallies.
Dali, W. H. 1877. On the California species of Fums. [Published
as a preprint of Proceedings of the California Academy of
Sciences 7: 1-5; vol. 7 of Proceedings never published].
Dali, W.H. 1902. Illustrations and descriptions of new,
unfigured, or imperfectly known shells, chiefly American,
in the U.S. National Museum. Proceedings of the United
States National Museum 24(1264): 499-566.
Durham, E.W. 1940. Seripps cruise to the Gulf of California:
Part II. Megascopic Paleontology and Marine Stratigra¬
phy. Geological Society of America, Memoir 43: 1-194.
Emerson, W.K. and L.C. Hertlein. 1964. Invertebrate
megafossils of the Belvedere Expedition to the Gulf of
California. Transactions of the San Diego Society of Natural
History 13: 333-368 .
Cibson-Smith, J., W. Cibson-Smith, and G.J. Vermeij. 1997.
Pacific Mexican affinities of new species of the gastro¬
pod genus Macron (Pseudolividae) and Neorapana
(Muricidae) from the Cantaure Formation (Early Miocene)
of Venezuela. Veliger 40: 358-363.
Gould, A. A. 1853. Descriptions of shells from the Gulf of
California and the Pacific coasts of Mexico and California.
Boston Journal ol Natural History 6: 374-408.
Gray, J.E. 1853. On the division of ctenobranehous gastero-
podous Mollusca into larger groups and families. Proceed¬
ings of the Zoological Society of London 21: 32-44.
Harry, H.W. 1985. Synopsis of the supraspecific classification
of living oysters (Bivalvia: Gyphaeidae and Ostreidae).
The Veliger 28: 121-158.
Hertz, CM., B.W. Meyers, J. Gemmell and D.L. Geiger. 1999.
A discussion of three Fusinus species in the northern Golfo
de California at San Felipe. The Festivus 31(7): 75-84.
Keen, A M. 1958. Sea shells of tropical west America: marine
Mollusks from Lower California to Colombia. Stanford
University Press, Stanford, xii, 624 pp.
Landau, B.M., Harzhauser, M., Islamogl, Y. and C.M. da Silva.
2013. Systematic^ and palaeobiology of the gastropods of
the Middle Miocene (Serrafallian) Karaman Basin Turkey.
Cainozoic Research 11-13: 1-584.
Linnaeus, C. 1758. Systema Naturae per Regna Tria Naturae:
secundum classes, ordines, genera, species, cum charac-
teristilms, differentiis, synonymis, locis. . . Edition decima,
reformata, Tom 1. Animalia. Laurentii Salvii, Holmiae
[Stockholm], iv, 823 pp.
Lowe, H.N. 1935. New marine Mollusca from West Mexico,
together with a list of shells collected at Punta Punasco,
Sonora, Mexico. Transactions of the San Diego Society
of Natural History 8(6): 15-34.
Marko, P. B. and G.| Vermeij. 1999. Molecular phylogenics
and the evolution of labi al spines among eastern Pacific
ocenebrine gastropods. Molecular Phylogenics and Evolu¬
tion 13: 275-288.
McLean, J.H. 1970. New eastern subgenera of Turbo Linnaeus
1758 and Astraea Boding, 1798. The Veliger 13: 71-72.
Nielsen, S.N. and D. Frassinetti. 2003. New and little known
species of Pseudolividae (Gastropoda) from the Tertiary
of Chile. Nautilus 1 17: 91-96.
Petuch, E.J. 1988. Neogene History of Tropical American Mol¬
lusks: Biogeography & Evolutionary Patterns of Tropical
Western Atlantic Mollusca. The Coastal Education and
Research Foundation [CERF], Charlottesville, ii + 217 pp.
Rafinesque, C.S. 1815. Analyse de la nature ou tableau de
l’universe et des corps organises. Palermo. 224 pp.
Reeve, L.[A.]. 1847. Monograph of the genus Turhinella.
Conchologia Iconica 4: [unpaginated text], pis. 1-13.
Russo, P. 2015. On the systematic position of Murex syracusanus
Linnaeus, 1758 (Gastropoda, Fasciolariidae) with reval¬
uation of the genus Aptyxis. Bollettino Malacologico
51: 79-86.
THE NAUTILUS, Vol. 130, No. 3
Page 126
Troschel, F. H. 1866-1893. Das Gebiss der Schnecken zur
Begriindung einer natiirlichen Classification. (Vol. 2),
Nicolaische Verlagsbuchhandlung, Berlin. i-L\, 409 pp.
[Published in parts, part 2, 49-96, pis. 5-8, 1868],
Tryon, G.W. Jr. 1887. Manual of Conehology, structural and
systematic, with illustrations of the species. Volume 9.
G.W. Tryon, Academy of Natural Sciences, Philadelphia,
488 pp.
Vermeij, G.| 1993. Spinucella, a new genus of Miocene to
Pleistocene murieid Gastropods from the eastern Atlantic.
Contributions to Tertiary and Quarternary Geology
30(1-2): 19-27.
Wrigley, A.G. 1927. Notes on English Eocene Mollusca, with
descriptions of new species. II. The Fusinidae. Proceed¬
ings of the Malacological Society of London 17(5, 6):
216-249.
THE NAUTILUS 130(3): 127-13 1 , 2016
Page 127
The western Caribbean complex of Fasciolaria tephrina de Souza,
2002 (Gastropoda: Fasciolariidae), with the description
of a new species
Emilio F. Garcia
115 Oak Crest Drive
Lafayette, LA 70503 USA
Efg21 [email protected]
William G. Lyons
4227 Porpoise Drive SE
St. Petersburg, EL 33705 USA
and
Academy of Natural Sciences of Drexel University
Philadelphia, PA 19103 USA
[email protected]
Martin Avery Snyder
Academy of Natural Sciences of Drexel University
Philadelphia PA 19103, USA
and
Museum National d'Histoire Naturelle
Paris, FRANCE
[email protected]
ABSTRACT
Fasciolaria delicatissima new species, a capacious, thin-shelled,
deep-water faseiolariid f rom a restricted area of the southwestern
Caribbean Sea, is compared with the similar F. tephrina de
Souza, 2002 and F. cf. tephrina, a presumed color form of the
latter; all three taxa have paucispiral protoconchs.
Additional Keywords: Southwestern Caribbean Sea, Honduras,
Nicaragua, Recent
INTRODUCTION
The molluscan fauna of the western Caribbean was mostly
unknown when one of the authors (EFG) began visiting
Roatan Island, Bay Islands, Honduras in 1978. Many
vessels of the commercially important Honduran shrimp
and spiny lobster fisheries land catches at Roatan, and
bycatches from those vessels has revealed a remarkable
molluscan fauna. Most species caught in shrimp trawls
are different than those caught in lobster traps, but both
types of collecting gear have produced many indigenous
species previously unknown to science. The fauna is so
different that Petuch (1988: 62) identified it as one of
several “relict pockets” in the Caribbean region. Although
Petuch considered the pockets center to be around the
Bay Islands (Utila, Roatan, and Guanaja) just off the
Honduran coast, in reality the area comprises the entire
wide continental shelf and upper slope that borders the
Caribbean coasts of Honduras and Nicaragua (Figure 17).
The range of the classic species Valuta polypleura Crosse,
1876 roughly defines the boundary of this “pocket.”
After a lapse of 15 years, trips to Roatan were resumed
in 2012 by EFG. Some previous contacts had lately been
working aboard vessels fish-trapping on the Honduran
slope seaward of the 100-fathom (183 m) isobath
(Figure 17). This activity had yielded species not seen
during some 60 previous trips, over a period of 20 years.
Of particular interest was the recently described Fusilaria
garciai Snyder, 2013, a faseiolariid with a paucispiral
protoconch that grows to 225 mm and seems to inhabit
at least the length of the northern upper continental
slope from north of Cabo Camaron, Honduras to the
Nicaraguan border. Also of great interest was a presumed
color form of the typically white F. tephrina de Souza,
2002. Reported as the “splotched” form of F. tephrina
by Snyder (2013: 128), the form is referred to here as
F. cf. tephrina. Both the white and the colored form have
been collected near the northeastern corner of the con¬
tinental slope (Figure 17), but only the colored form has
been collected with Fusilaria garciai.
The type locality for Fasciolaria tephrina is north of
Quita Sueno Bank, Colombia (off northeastern Nicaragua),
14°40' N, 81°25' W, depth 480 m (de Souza, 2002;
Figure 17). However, de Souza also recorded a specimen
in the Kevan and Linda Sunderland collection that was
taken “off Roatan Ish, Honduras, 420 m, in fish trap.”
The fact that the Sunderland specimen was collected in a
fish trap seems telling. Fishermen who have collected
F tephrina and F cf. tephrina report that they are caught
in fish traps set off the edge of the continental shelf. The
topography of the upper slope in that area is too rough
for trawling and probably unattractive to spiny lobsters,
which usually live in warmer, shallower waters, perhaps
explaining why the area has been avoided by shrimp and
lobster fishermen.
Fasciolaria tephrina has also been reported from off
Isla de San Andres, Colombia (e.g., Miloslavich et ah,
2010; Snyder, 2013: 128, fig. 14), an island of volcanic
origin separated from the continental shelf and located
off the southeastern Nicaraguan coast (Figure 17). How¬
ever, inspection has revealed several shell characters that
differentiate the San Andres population from F tephrina.
The Saji Andres population is described here as a new
species and compared with F tephrina and F cf. tephrina.
THE NAUTILUS, Vol. 130, No. 3
Page 128
MATERIALS AND METHODS
Specimens studied are located at the Museu de Zoologia
da Universidade de Sao Paulo, Brazil (MZSP), the
Academy of Natural Sciences of Drexel University,
Philadelphia (ANSP), the William G. Lyons study collec¬
tion (LC) and the author’s collection (EFG). Shell sizes
are reported in millimeters (mm), measured to nearest
0.1 mm with electronic digital calipers. Unless otherwise
specified, measurements are of shell length (height),
measured as the distance between the apex and the
abapical end of the siphonal canal. Shells were weighed
with a Mettler Toledo PB metric scale model PB 3002-S
and measured to the nearest 0.01 grams.
SYSTEMATICS
Family Fasciolariidae Gray, 1853
Genus Fasciola ria Lamarck, 1799
Tvpe Species: Murex tulipa Linnaeus, 1 758, by monotype.
Fasciolaria delicatissima new species
(Figures 1-7)
Fasciolaria tephrina. — Mallard and Robin, 2005: 8, pi. 4;
Robin, 2008: 207, fig. 9; Miloslavich et ah, 2010:
Table S6, no. 925 (in part); Snyder, 2013: 128,
fig. 14 (in part) ( non Fasciolaria teplirina de Souza
(2002), nee Snyder (2013: 128, fig. 15)).
Diagnosis: Shell large, capacious, with very thin walls
and paucispiral protoconch; shell color off-white with
many thin, brown, uneven and sometimes interrupted
spiral lines on spire and body whorl; lines not extending
onto tan siphonal process.
Description: Shell large, up to 280 mm in length
(Rosenberg, 2009), 105 mm wide, fusiform, semi-glossy,
with swollen, rounded whorls and uncommonly thin
walls. Protoconch (Figure 7) white, large, height approx¬
imately 3.5 to 4 mm, smooth, with one whorl; first half of
whorl rounded, second half swollen; protoconch axis not
deviating from that of teleoeonch. Teleoconeh with
7 rounded whorls, first two whorls shouldered to periphery,
then straight (Figure 7); remaining whorls progressively
more convex, body whorl swollen, globose. Suture deep,
bordered anteriorly by a strong cord wrinkled by minute
axial threads. Axial sculpture of numerous microscopic
threads showing at start of teleoconeh, approximately 30
on first whorl and 60 on second whorl; axial swelling
appearing at periphery of second whorl, developing some
16 low nodes (Figure 7); nodes disappearing on later
whorls; minute axial threads continuing on surface of shell
through last whorl, becoming stronger on siphonal canal.
Spiral sculpture subtly developing at beginning of first
whorl, approximately 8 strong threads showing on second
whorl, 5 threads on shoulder, which will become strong
cords on later whorls; three threads below periphery
disappearing on later whorls; spiral threads on early
whorls creating a somewhat reticulate sculpture as they
cross axial elements; two adapical threads strongest,
wrinkled by axial elements. Siphonal canal straight or
rather sinuous, long, approximately 30 to 35% of shell
length. Aperture widely oval, with many faint interior
lirae best detected by rubbing a fingernail or pin across
the surface; width approximately 20% of shell length.
Parietal wall smooth, without callus; columella with two
strong, oblique anterior plications, posterior one some¬
what bifurcate; plications inconspicuously continuing as
two or three long axial elements in different degrees of
strength along dorsal side of siphonal canal (Figure 5).
Outer lip thin, with 5 denticles at posterior end reflect¬
ing terminations of pre-sutural cords. Shell color off-
white with uneven peneiled-Iike brown spiral lines that
terminate at beginning of siphonal canal and that show
through to inside aperture; columellar area and siphonal
canal light tan, canal with sporadic dark brown blotches
in mature specimens.
Tvpe Material: Holotype (Figures 1-2), ANSP 450737,
277 x 1 05 mm, north of Cayos de San Andres, Caribbean
Sea, Colombia, trawled, 350 m,; paratype 1, ANSP 466083
(Figure 3), 182 mm, with operculum; Paratype 2, EFG
31404 (Figures 4-6), 170.1 mm,. All from type locality.
Other Material Examined: Two shells, LC unnum¬
bered, 177.4 and 188.1 mm, trawled north of Cayos de
San Andres, Colombia, depth 350 m; one shell, LC
unnumbered, 87.1 mm, Golfo de Morrosquillo, Colombia.
Type Locality: North of Cayos de San Andres,
Caribbean Sea, Colombia, 350 m.
Distribution: North of Cayos de San Andres, Caribbean
Sea, Colombia, trawled, 350 m; Golfo de Morrosquillo(P).
Etymology: From the Latin adjective delicatus , -inn,
meaning delicate; used in the superlative to denote the
unusual lightness and fragile nature of the shell.
Discussion: Fasciolaria delicatissima new species can
he differentiated from most congeners by its unusually
thin walls, its color markings and its paucispiral
protoconch. The new species has been confused with
F. tephrina de Souza, 2002 (Figures 10-13), and F. cf.
tephrina (Figures 8-19, 14-16). Fasciolaria delicatissima
shares with F. tephrina the large, capacious shell, the
paucispiral protoconch, the deep-water habitat and rela¬
tive geographic proximity. However, F. tephrina has a
protoconch that deviates from the main axis, whereas the
new species lacks that character; its protoconch is differ¬
ently shaped, with a less convex first half of the whorl and
a more bulbous second half (compare Figure 7 with
Figures 9, 12 and 16). This difference is possibly what
causes the protoconch of F. tephrina to be “deviated.”
Early teleoconeh whorls of F. delicatissima are differ¬
ently sculptured, with more spiral elements and more
numerous (16 vs. 10), weaker axial elements than those
of F tephrina (compare Figure 7 with Figures 9, 12,
E.F. Garcia et al., 2016
Page 129
Figures 1-9. Fasciolaria species. 1-7. Fasciolaria delicatissima new species, north of Cayos de San Andres, 350 m. 1-2. Holotype,
ANSP 450737 (image copyright from Femorale with permission), length 277 mm, width 105 mm. 3. Paratype ANSP 466083
(image copyright from Femorale with permission), 182 mm, specimen with operculum. 4-7. Paratype, EFG 31404, length 164.9 mm,
width 64.5 mm. 8-9. Fasciolaria cf. tephrina, EFG 31298, length 165.9 mm, width 45 mm, near Honduras-Nicaragua border,
in fish trap, 300 in.
and 16). Moreover, the shell surface of the new species is
covered with minute axial threads evident under low
magnification (Figure 6), its parietal area lacks an enam¬
eled wall, and the lirae inside its aperture are determined
by tact; they are visually undetected. The two plications
of F. tephrina are limited to the columella, but those
of F. delicatissima continue as ridges over the siphonal
canal (Figure 5).
The shell of F. delicatissima is much thinner than that
of F. tephrina. A 165.9 mm specimen of F. cf. tephrina
(FFG 31298; Figures 8-9) weighs 46.06 grams, whereas
a similar 164.9 mm specimen of F. delicatissima
Page 130
THE NAUTILUS, Vol. 130, No. 3
Figures 10-16. Fasciolaria species. 10-13. Fasciolaria tephrina de Souza, 2002. 10-12. Holotype MZSP 35048, length 187.4 mm,
width 73.8 mm, north of Quita Sueno Bank, 14°40' N, 81025' W, in 480 m. 13. EFG 31107, length 220 mm, width 85 mm, near
Honduras-Nicaragua border, in fish trap, 300 m. 14-16. Fasciolaria cf. tephrina, EFG 30890, length 230 mm, width 90 mm, near
Honduras- Nicaragua border, in fish trap, 300 m.
(Paratype 2, EFG 31404; Figures 4-7) weighs only
23.16 grams.
The multilineate pattern and coloration of the shell of
F. delicatissima is consistent in all specimens examined,
differing from the white of F tephrina (Figures 10-13)
and the blotched pattern F. cf. tephrina (Figures 8-9,
14-16). The six specimens of F. cf. tephrina that were
studied, all collected empty, clearly show the character¬
istic blotched markings. Even the most eroded speci¬
mens have darker brown coloration at the tip of the
anterior canal.
Although color intergrades between F. tephrina and
F. cf. tephrina have not been seen, no differences between
them are evident except coloration and a stronger enam¬
eled parietal shield in the white form. One may theorize
that because all specimens of F. tephrina have been
E.F. Garcia et al., 2016
Page 131
Figure 17. Bathymetric chart of the seafloor off Honduras
and Nicaragua.
collected empty and the type material is in poor condition,
shells of the white form could represent bleached material.
However, several better-preserved specimens of the white
form have come to light since the original description
(e.g.,Yidi and Sarrmento, 201 1, Figure 493; also Figure 13,
herein) and they show no indications of coloration.
The bathymetry at the eastern edge of the Honduran
continental shell and slope is rugged and complex
(Figure 17). Quita Sueno Bank is not far from the edge
ol the continental shelf, but the abrupt benthic topography
between them may lend itself to the development of small
allopatrie populations, particularly with direct developers
such as species of Fasciolaria. Fishermen may spend
weeks at sea and obtain shells from several non-contiguous
populations and as they range over the continental slope,
but such bycatch may be landed in a single box, making
precise origins of the shells difficult to ascertain.
We question the “Golfo de Morrosquillo” locality for
F. delicatissima . The Golfo is much too shallow to sup¬
port the species, which otherwise has not been reported
despite intensive trawling for shrimp there for at least
four decades. Data available for all of tbe other
F delicatissima specimens indicate that they were
obtained by trawls; at least one specimen was collected
alive (Figure 3), which confirms the efficacy of that
collecting method. This implies that the species lives on
a flat bottom accessible to trawlers, and that would be at
the base of the escarpment (Figure 17). Conversely,
Fasciolaria tephrina and F. cf. tephrina have been col¬
lected exclusively “crabbed” in fish traps, the seafloor not
being conducive to trawling. The latter two forms do not
seem to intergrade, but until more precise catch data
become available, they should be regarded as belonging
to a single species.
AC K N OWLE DC M E NTS
Our thanks to Dr. Geerat Vermeij for reading the
manuscript, to Jose and Marcus Coltro of Femorale for
allowing us to use their copyrighted images of Fasciolaria
‘'tephrina” (Figures 1-3), and to Drs. L.K.S. Simone and
Carlo M. Cunha (MZSP) for providing the images of the
holotype of F. tephrina.
LITERATURE CITED
Crosse, H 1876. Sur une variete du Valuta musica Linne.
Journal de Conehvliologie 24: 163-166, pi. 5.
de Souza, P.J.S. 2002. A new bathyal Fasciolaria (Mollusca:
Caenogastropoda) from the southwestern Caribbean.
Zootaxa 49: 1-7.
Gray, J.E. 1853. On the division of ctenob ranch ous gasteropodus
Mollusca into larger groups and families. Annals and Maga¬
zine of Natural History, Second Series, 1 1(62): 124—133.
Lamarck [J.P.B.A.]. 1799. Prodrome d une nouvelle classification
des eoquilles comprenant une redaction appropriee des
caracteres generiques et etablissement dime grand nombre
de genres nouveaux. Memoires de la Societe d’Histoire
NatureUe de Paris I: 63-91, table.
Linnaeus, C. 1758. Systema naturae per regna tria naturae:
secundum classes, ordines, genera, species, cum characteris-
tibus, differentiis, synonymis, locis. Vol. 1. Editio decimal,
refonnata. Laurentii Salvii, Holmiae lii + 824 pp.
Mallard, D. and A. Robin. 2005. Fasciolariidae. Museum du
Coquillage, Les Sables d'Olonne. 27 pp., 70 pis.
Miloslavich, P, J.M. Diaz, E. Klein, J.| Alvarado, C. Diaz, |
Gobin, et al. 2010. Marine biodiversity in the Caribbean;
regional estimates and distribution patterns. PLoS ONE
5(8): el 1916. dot: doi 10. 1371/journal. pone. 001 1916.
Petuch, E.J. 1988. Neogene history of tropical American
mollusks. Coastal Education & Research Foundation:
Charlottesville, Virginia, [vi] + 217, 39 pis.
Robin, A. 2008. Encyclopedia ol marine gastropods. Xenophora
and ConchBooks, Hackenheim, 480 pp.
Rosenberg, G. 2009. Malacolog 4.1.1: A Database of Western
Atlantic Marine Mollusca. [WWW database (version 4.1.1)]
URL http://www. malacolog.org/.
Snyder, M.A. 2013. Description of Fusilaria garciai new
genus, new species (Gastropoda: Fasciolariidae: Fasciolariinae)
from the western Caribbean Sea. The Nautilus 127:
125-129.
Yidi, E. and V. Sarmiento. 2011. Colombian Seashells from
the Caribbean Sea. LInformatore Piceno, Ancona, 384 pp.,
167 pis.
A tumbling snail (Gastropoda:
Vetigastropoda: Margaritidae)
In August 2015, the NOAA ship Okeanos Explorer con¬
ducted deep-sea studies in the northwestern Hawaiian
Islands, which now are within the Papahanaumokuakea
Marine National Monument. The ship deployed the
remotely operated vehicle Deep Discoverer (D2 KOV),
whose live video feed was shared with researchers on
shore via satellite transmission.
On 5 August 2015, the D2 HOY was exploring angular
basalt blocks and sediment patches on the steep inner
slope of Maro Crater, an unusual 6 km-wide crater east
of Maro Reef (25.16° N, 169.88° W, 2998-3027 m). The
cameras recorded what appeared to he a fish attacking or
being attacked by some other unidentified animal. When
the ROY cameras were zoomed in on the encounter, the
twisting elongate structure though to be a fish was deter¬
mined to actually be the elongate foot of a gastropod
mollusk (Figures 1, 2). No potential predator or prey
could be seen in subsequent view, so it seems likely that
the snail reacted to the close presence of the ROV. The
mollusk first moved horizontally before falling down,
retracting the foot and resting among the rocks. The
camera was equipped with red strobe lights that set a
scale of 10 cm. Because the camera’s focal length
changed, it is difficult to estimate how far the snail
moved but 3 m seems to be a reasonable guess.
Examination of the shell of the mollusk (Figure 3)
showed that it belonged to the genus Gaza Watson, 1879.
Species of Gaza are among the larger gastropods to be
found on the continental shelf and upper slope. They can
be recognized by their size (to 40 mm), ivory color with a
golden sheen, deep umbilicus, uncalcified operculum,
and lack of a noticeable periostracum. Most specimens
have been obtained by trawls on mud bottoms.
The mollusk recorded by the D2 ROV was not col¬
lected for verification. Gaza daedala Watson, 1879 is the
only species of this genus known from the central or
western Pacific. The only report for which a specimen is
known is that of the holotvpe, collected at 19" 10' S,
178°10' E (off Kandavu, Fiji), 1100 m. Robert Moffett
and Christopher Kelley (pers. comm.) informed me that
a specimen of G. daedala was collected off Kauluoa
Point, Big Island of Hawaii (19.34° N, 155.91° W, 600-
803 m, 24 August 1988, submersible vehicle Pisces,) but
the specimen has been lost. Severns (2011) reported G.
daedala from 330 m off Oahu but the material on which
that report was based also is missing. The mollusk in the
photographs has an iridescent shell with radiating lines,
as reported for G. daedala (Simone and Cunha, 2006).
The width of the shell, based on the camera’s scale dots,
seems to have been close to 40 mm, with the extended
foot as much as 100 mm. The characteristic covered
umbilicus can only partially be seen in the photograph,
so the identification remains uncertain. Hickman (2012)
noted that species of Gaza from the Gulf of Mexico
might be associated with chemosvnthetie eommuni-
ties, but the mollusk in tire photographs was living on
manganese-encrusted basalt.
Hickman (2003; 2007) reported “foot thrashing” as an
escape response to predators and in the laboratory by
“mechanical disturbance” in the trochoidean gastropods
Umbonium vestiarium (Linnaeus, 1758), I.sanda coronata
A. Adams, 1854, and other species of the family
Solariellidae. The observations provided here are the first
of such behavior in Gaza spp. and among the few reports
on behavior of non-cephalopod deep-sea mollusks.
Supporting material: video is posted at http://
oceanexplorer.noaa.gov/okeanos/explorations/exl504/logs/
dive4/dive4.html.
I thank Carole Hickman of the University of California,
Berkeley, for her generous assistance with informa¬
tion on behavior and identification of the tumbling
snail. Bob Moffett, National Oceanographic and Atmo¬
spheric Administration; and Chris Kelley, Hawaii Under¬
sea Research Laboratory, for more information on
specimens of G. daedala , and anonymous reviewers
for helpful comments.
The photographs and video were collected under the
auspices of the National Oceanographic and Atmospheric
Administration Off ice of Ocean Exploration and Research,
2015 Hohonu Moana.
LITERATURE CITED
Hickman, C.S. 2003. Functional moqrhology and mode of
life of lsanda coronata (Gastropoda: Trochidae) in an
Australian macrotidal sandllat. In: Wells, F. E., D.l. Walker,
and D.S. Jones (eds.) The marine flora and fauna of
Dampier, Western Australia. Western Australian Museum,
Perth, pp. 69-88.
Hickman, C.S. 2007. Nocturnal swimming, aggregation at
light traps, and mass spawning of scissurellid gastropods
(Mollusca: Vetigastropoda). Invertebrate Zoology 126:
10-17.
Hickman, C.S. 2012. A new genus and two new species of
deep-sea gastropods (Gastropoda: Vetigastropoda: Gazidae).
The Nautilus 126: 57-67.
Severns, M. 2011. Shells of the Hawaiian Islands. IKAN
Unterwasser-Archiv, Frankfurt, Germany, 1021 pp.
Simone, L. H. and C.M. Cunha, 2006. Revision of genera
Gaza and Callogaza (Vetigastropoda, Trochidae), with
M.K. Wicksten, 2016
Page 133
Figures 1-3. Gaza sp. 1, 2. Individual tumbling with foot extended. 3. Close-up detail Photos courtesy of NOAA Oflice of Ocean
Exploration and Research, 2015 Hohonu Moana Expedition.
description of a new Brazilian species. Zootaxa 1318:
1-40.
Watson, R.B. 1879. Mollusca of the H.M.S. ' Challenger” expe¬
dition. 111. Troehidae, xaz., the genera Seguenzia , Basilissa,
Gaza, and Bembix. Zoological Journal of the Linnean
Society, London 14, pp. 586-605.
Mary K. Wicksten
Department of Biology
Texas A&M University
College Station, TX 77843-3258 USA
[email protected]
THE NAUTILUS 130(3): 134-136, 2016 Page 134
Research Note
The authorship of Turrilatirus craticulatus
(Gastropoda: Fasciolariidae: Peristemiinae)
The specific name of the Indo-West Pacific gastropod
Turrilatirus craticulatiis generally has been attributed
to Linnaeus, 1758 in the original combination Murex
craticulatiis, but the correct attribution is to Gmelin,
1791, in the combination Voluta craticulata.
In the tenth edition of Si/stema naturae, Linnaeus
(1758: 755, sp. 495) introduced Murex craticulatiis without
reference to a figure or designated locality. Linne (1767:
1224-1225, sp. 569) repeated the description, added a
phrase “ Turbo angulatus,” cited a figure by Rondelet
(1555: 89) and added the locality “A/. Mediterranean
Rondelet’s figure shows a shell with about four angular
spire whorls plus a body whorl and a relatively long
siphonal canal.
Born ( 1 778; 1 780) and Chemnitz (1780) treated a differ¬
ent shell as M. craticulatiis. Born (1780) and Chemnitz
(1780) each re-described what they believed to be
M. craticulatiis Linnaeus and provided new figures, still
accepting its Mediterranean locality. Richardson et al.
(1979: 165) noted that Chemnitz figs 1382 and 1383 were
cited by Born both in 1778 and 1780. That incongruity of
dates was explained by Kohn (1964: 153), who provided
evidence that Born received plates from Martini at least
two years before they were published by Chemnitz.
Chemnitz expressed uncertainty regarding the name, con¬
cluding that although the original description did not fit
exactly, Linnaeus had intended it for a shell that Chemnitz
recognized. The shell they described and figured, now
well-known as an Indo-West Pacific species, is not the
species that Rondelet figured, nor is it the one that
Linnaeus described.
Gmelin (1791: 3554, sp. 105) repeated the Linnaean
description of the Mediterranean M. craticulatus, again
citing the Rondelet figure. Some pages earlier, however
(1791: 3464), he provided a separate account for a new
species, Voluta craticulata . The impetus for classification
in Voluta apparently traces to Schroter (1783: 284), who
cited “Martini iv [Chemnitz, 1780] tab. 1382 & 1383;
Lister, [1688] Hist. Conch, pi. 919. fig. 13. mala. pi. 967,
fig. 22; Seba [1758], pi. 50, figs 55 & 56, pi. 51, figs 31 &
32; Knorr [1764], pi. 3, fig. 6" in a taxon lie called Voluta
sp. Chemnitz had assigned these figures to M craticulatus
Linnaeus, but Gmelin cited all of them for craticulata.
By these actions, Gmelin recognized that the species
described by Linnaeus was not the shell figured by
Chemnitz and Born.
From the start, Gmelin s distinction was misunder¬
stood. Schreibers (1793: 121, 229) treated M. craticulatus
and V. craticulata, citing correct figures for the latter hut
attributing both names to Linnaeus. Rtiding (1798) asso¬
ciated M. craticulatus (not V craticulata) “Gmel.” with
the Chemnitz figures and reclassified the species in
Fusus. Dillwyn (1817) retained M. craticulatus Linnaeus
hut added V. craticulata Gmelin as a synonym. Lamarck
(1822) placed craticulatus "Lin. Gmel. p. 3554” in
Turbinella and cited “ Voluta craticulata Gmel. p. 3464”
as a junior synonym, as did Deshayes (1832), who cited
the same figures cited by Gmelin. The species was
maintained as T. craticulata by Schubert and Wagner
(1829) and Kiener (1840), both of whom attributed the
name to Lamarck as first user of the combination, and
by Anton (1838) and Reeve (1847), who attributed the
name to Linnaeus.
Deshayes (1843: 386) headed an account “ Turbinella
craticulata Lamk.” but made clear in synonymy and in a
footnote that the name’s author was Linnaeus. Deshayes
(1843: 457) later suggested that some characters mentioned
in the brief original description of Murex craticulatus
seemed to fit a variety of the Mediterranean species Fusus
syracusanus (Linnaeus, 1758). This comment appeared in
a footnote wherein Deshayes considered hut did not
accept the Linnaean name as an earlier name for Fusus
strigosus Lamarck, now Fusinus rostratm (Olivi, 1792).
Clearly, Deshayes was uncertain about the identity of the
Linnaean name.
In another footnote, Deshayes (1843: 386) noted: “Sous
ce nom de craticulata MM. Schubert et Wagner ont
decrit et figure une espece tres distincte du veritable
Murex craticulatus de Linne et des autres auteurs.” Simi¬
larly, Dodge (1957: 188) noted that Schubert and Wagner
(1829: 103) had followed Lamarcks (1822) synonymy for
Turbinella craticulata , hut their figures showed an entirely
different shell that he could not identify. Both Deshayes
and Dodge overlooked the fact that the figures were for
T. craticulata Lamarck “var. b” of Schubert and Wagner
(1829: pi. 227, figs 4023 & 4024), which Anton (1838)
later renamed T. wagneri.
Kiister in Kuster and Kobelt (1844: 21) cited for
T. craticulata both M. craticulatus Gmel. Lin. p. 3554
and V. craticulata Gmelin p. 3464 but also cited for it
the figures by Lister, Seba, Knorr, and Chemnitz (but not
Rondelet), and attributed authorship to Gmelin.
Hanley (1855) remarked that the shell figured by
Kiener (1840) ( i.e ., the Indo-West Pacific craticulatus)
“answers very' correctly to the description by Linnaeus,”
but he soon changed his mind. Hanley (1856: 134) speci¬
fied that M. craticulatus of Wood (1828), the T. craticulata
of Kiener and others, was based on V. craticulata
Gmelin, indicating his belief that the name applied to
a different species.
Kobelt (1876a: 22) also attributed the name to Gmelin
but mistakenly cited for it p. 3554, Gmelins page for
M. craticulatus Linnaeus. Kobelt (1876b: 54) mistakenly
cited “craticulatus Lam.” ( non Linnaeus) as a junior
W.G. Lyons and M.A. Snyder
Page 135
synonym of Latirus turrittis (Gmelin, 1791), another
Indo-Pacific species of Turrildtirus Vermeij and Snyder,
2006. Kobelt in Kiister and Kobelt (1876: 1 18; 1877: 58)
attributed T. craticulata to Gmelin, without mention
of Linnaeus.
Tryon (1880: p. 93, pi. 69, fig. 159) identified Latirus
craticulatus (Linnaeus, 1758) as the Recent Indo-West
Pacific species. Tryon’s assignment was unanimously
followed until Dodge (1957) reviewed the Linnaean
name. Dodge agreed that M. craticulatus Linnaeus and
V. craticulata Gmelin applied to separate species, but he
said that attribution of the name craticulatus Linnaeus to
the Indo-West Pacific species had been so consistently
accepted that it would be confusing to change it. Never¬
theless, he recommended that, for a sound nomencla¬
ture, the Linnaean name should be regarded as a nomen
dubium. On this we concur.
Dodge thought it difficult to disregard Gmelin’s con¬
siderable synonymy but maintained that Gmelin only
increased the confusion begun by Linnaeus because the
description of V. craticulata is “equivocal,” the shell size
given by Gmelin is “somewhat too long,” and no locality
is given. Consequently, he advocated that Gmelin s name
be ignored and craticulatus auct. be attributed to
Dillwyn (1817), who had provided a new description for
M. craticulatus ‘Linnaeus’ and cited the above mentioned
appropriate figures. Nevertheless, all mentions of the
Indo-Pacific species published since Dodge have ignored
his recommendations. Most recently, Marais and Kilbum
(2010: 128) correctly cited the original combination for
the species as V. craticulata but attributed the name
to Linnaeus.
No evidence supports the association of the Linnaean
descriptions, the figure he cited, or the Mediterranean
locality with the Indo-West Pacific species. There is no
specimen of Mu rex craticulatus in the Linnaean collec¬
tion (Hanley, 1855), nor was the species contained in the
Museum Ulricae (Dodge, 1957), so no type material
is available.
The remedy that Dodge proposed is impractical.
Dillwyn s description and figures he cited may have been
appropriate but he, like Born, presented the species as
M. craticulatus Linnaeus, with Gmelin’s name as a syno¬
nym, so Dillwyn’s M. craticulatus is a junior primary
homonym of the Linnaean name (ICZN, 1999: 59; Arti¬
cle 57.2), as are those of Born, Chemnitz, etc. The
Chemnitz name is also unavailable as it was published in
a rejected work (ICZN, 1987: 319). Conversely, Gmelin
distinguished V. craticulata from M. craticulatus Linnaeus
by providing separate accounts in different genera for
the two species, by providing a separate description for
the new species, and by citing figures wherein its iden¬
tity is unmistakable. Clearly, the first available name
for the Indo-West Pacific species is Valuta craticulata
Gmelin, 1791.
For a discussion of the complex synonymic history
of Murex craticulatus “Linnaeus” Brocchi, 1814, a
Mediterranean muricid, see Houart (2001: 82-84).
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Anton, 1I.E. 1838. Verzeichniss der Conchylien welche sich in
der Sammlung von Hermann Eduard Anton befinden.
Eduard Anton, Halle, xvi, 1 10 pp.
Born, I 1778. Index Rerum Naturalium Musei Caesarei
Vindobonensis, Pars Prima: Testacea. Verzeichnis der
naturlichen Seltenheiten des k. k. Naturalien Cabinets
zu Wein. Erster Theil: Schalthiere. Kraus, Vindobonae
[Vienna], [xlii], 458, [82] pp., 1 pi.
Born, I. 1880. Testacei Musei Caesarei Vindobonensis, quae
jussu Mariae Theresiae Augustae disposuit et descripsit.
Kraus, Vindobonae [Vienna], xxxvi, 442, [15] pp., 18 pis.
col., text illustrations.
Brocchi, G.B. 1814. Conchiologia fossile snbapennina con
osservazioni geologiche sugli Apennini vicentin, e sul
suolo adiacente. Stamperia Reale, Milano, 2 vols. [1: 1-
240, 2: 241-712, pis. 1-16],
Chemnitz, J.H. 1780. Neues systematisehes Conchylien Cabi¬
net 4: [24], 1-344, vignettes 37-41, pis. 122-159.
Deshayes, G.P 1832. Encyclopedic Methodique. Histoire
Naturelle des Vers. Tome troisieme (3). Agasse, Paris,
pp. 595-1 152.
Deshayes, G.P. 1843. Histoire Naturelle des Animaux sans
Vertebres; denxieme edition. Revue et augmentee de
notes presentant les faits nouveaux, dont la science s’est
enrichie jusqu’a ee jour. Tome neuvieme. Histoire des
mollusques. J.B. Bailliere, Librairie de l’Academie Royale
de Medicine, Paris and London, 728 pp.
Dillwyn, L.W. 1817. A descriptive catalogue of Recent
shells, arranged according to the Linnean method
with particular attention to the synonymy. John and Arthur
Arch, Cornhill, London. 2 volumes. I: i-xii, 14580; 2: 581-
1092, [29],
Dodge, H. 1957. A historical review of the mollusks of
Linnaeus. Part 5, the genus Murex of the class Gastropoda
Bulletin of the American Museum of Natural History 113:
73-224.
[Gmelin, |.F. 1791], Vermes. In: Caroli a Linnaei, systema
naturae per regna tria naturae. Editio decirna tertia,
Aueta, Reformata. Lipsiae (Leipzig), 1(6), pp. 3021-3910.
Hanley, S.C.T. 1855. Ipsa Linnaei Conchylia. The shells
of Linnaeus, determined from his manuscripts and
collection. Williams and Norgate, London, 556 pp.,
5 pis.
Hanley, S.C.T. 1856. Index Testaceologicus, an illustrated cata¬
logue of British and foreign shells . . . by W. Wood ... A
new and entirely revised edition, with ancient and modem
appellations, synonymy, localities, etc. Willis and Sotheran,
London, xx + 234 pp., 46 pis.
Houart, R. 2001. A review of the Recent Mediterranean and
northeastern Atlantic species of Muricidae. Evolver, Rome,
227 pp.
ICZN (International Commission on Zoological Nomencla¬
ture). 1987. Official lists and indexes of names and works
in zoology. International Trust for Zoological Nomencla¬
ture, London, 366 pp.
ICZN (International Commission on Zoological Nomencla¬
ture). 1999. International Code of Zoological Nomencla¬
ture. Fourth Edition. International Trust for Zoological
Nomenclature, London, xxix + 306 pp.
Kiener, L.C. 1840-1841. Species General et Iconographie des
Coquilles Vivantes, VI. Genre Turbinelle ( Turhinella ,
Lamarck). Rousseau et Bailliere, Paris, pis. 1-21 (1840);
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THE NAUTILUS, Vol. 130, No. 3
Kobelt, W, 1876a. Catalog der gattung Turbinella Lam.
Jahrb tidier der Deutsdien Malakozoologisdien Gesellschaft
3: 18-29.
Kobelt, W. 1876b. Die Muriciden des Rothen Meeres.
Jahrbiicher der Deutschen Malakozoologisdien Gesellsdiaft
3: 38-60.
Kobelt, W. 1877 in 1876-1878. Illustrirtes Conchylienbuch.
Erster Band. Verlag von Bauer 6c Raspe, Niirnberg,
pp. 1 — 40, pis. 1-10 (November 1876); 41-64, pis. 11-20
(November-December 1877); 65-88, pis. 21-30
( February- March 1878); 89-104, pis. 31-40 (May 1878);
I -XVI, 105-144, pis. 41-50 (May 1878).
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unbewehrten Purpurschnecken. Erste Halfte: Turbinella
und Fasciolaria. In Abbildungen nach der Natur mit
Beschreibungen, angerangen von Dr. H.C. Kiister, fortgesetzt
und vollendet von Dr. W. Kobelt. Systematisches
Conchylien-Cabinet von Martini und Chemnitz, neu
herausgegeben (series II), 3(3a): 1-164, pis. 1-32, 9a, 9b,
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cimeliorum e tribus regnis naturae quae olim collegerat.
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in quibus verae piscium effigies expressae sunt. Quae in
tota Piscium historia contineatur, indicat Elerchus pagina
nona et decima. Unito: Universae aquatilium Historiae
parsaltera cum veris ipsorum Imaginibus. Volume II: De
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pp. (12), 242, 1555; 450 text figs.
Schreibers, K. 1793. Versueh einer vollstandingen
Conchylienkentnis nach Linnes System. Joseph Eblen
von Kurzbeck, Vienna, Erster Band, 447 pp.
Schroter. | S. 1783. Einleitung in die Conchylienkenntniss,
nach Linne. Erster Band, von den Schnecken. Johann
Jacob Gebauer, Halle, Germany, i-xxxii. 1-860, (1-2),
pi. 1-3.
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Conchylien-Cabinet- Cabinet angefangen von Martini
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accurata descriptio, et iconibus artificiosissimis expressio,
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1-212, pis. 1-38.
William G. Lyons
4227 Porpoise Drive SE
St. Petersburg, FL 33705 USA
and
Academy of Natural Sciences of Drexel University
Philadelphia, PA 19103 USA
[email protected]
Martin Avery Snyder
Academy of Natural Sciences of Drexel University
Philadelphia PA 19103, USA
and
Museum National d’Histoire Naturelle
Paris, FRANCE
[email protected]
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