THE NAUTILUS
Ql~
HO!
N3H
IZ
Volume 129, Number 1
March 19, 2015
ISSN 0028-1344
A quarterly devoted
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THEf?NAUTILUS
Volume 129, Number 1
March 19, 2015
ISSN 0028-1344
CONTENTS
Jose ph H. Hartman New viviparid gastropods from the end Cretaceous and ear!y
Paleogene of the Williston Basin, USA and Canada . 1
M.E. Chrpa Shell shape variation within a population of Astarte borealis
A.O. Oleinik (Schumacher, 1817) (Bivalvia: Astartidae) from Camden Bay,
northern Alaska: a study using elliptical Fourier analysis . . . 23
Jeffrey H.R. Goddard Reviving a cold case: two northeastern Pacific dendrodorid
Angel Valdes nudibranchs reassessed (Gastropoda: Opisthobranchia) . 31
Sponsored in part by the State of
Florida, Department of State,
Division of Cultural Affairs and the
Florida Council on Arts and Culture
CULTURE
BUILDS
FLORIDA
FLORIDA DEPARTMENT of STATE
DIVISION of CULTURAL AFFAIRS
THE NAUTILUS 129(1): 1-22, 2015
Page 1
New viviparid gastropods from the end Cretaceous and early
Paleogene of the Williston Basin, USA and Canada
Joseph H. Hartman
University of North Dakota
Harold Hamm School of Geology and Geological Engineering
81 Cornell Street Stop 8358
Grand Forks, ND 58202 USA
[email protected]
ABSTRACT
Three new species of Viviparidae (Caenogastropoda) are
described and illustrated from the Williston Basin of North
Dakota, Montana, and Saskatchewan. The new Campeloma,
with strong shoulder development, is separated from others
in geologic time by recognizing its distinctive morphological
features. Campeloma acroterion new species occurs primarily
in Lancian North American Land-Mammal Age (NALMA)
strata, survives the Cretaceous-Paleogene extinction event,
and ranges into the late Puercan NALMA (end Cretaceous
to lower Paleocene; Hell Creek Formation into Tulloek Member
of Fort Union Formation and equivalents). Campeloma nebrascense
whitei, a larger and more rounded, but shouldered species,
appears as a derivative species in younger rocks, followed
and partially coeval with C. n. nebrascense, which is with¬
out shouldering. Vivipams purgatorius new species and
V codomorphus new species are species of discovery. Neither
species are common and the few specimens collected have
been assigned to more common taxa. Viviparus purgatorius
ranges from the Puercan to Torrejonian (lower and middle
Paleocene; Tulloek Member and Ravenscrag Formation), and
V codomorphus may be only known from the Torrejonian
NALMA (middle Paleocene; upper part of the Ludlow Member
of the Fort Union). These species of Vivipams appear derived
from more sculptured forms of V. thompsoni also present
in Lancian strata of the Lance and Hell Creek Formations.
With the regression of the Cannonball Sea and the expansion
of Laramidia in the early late Paleocene, viviparids radiate,
becoming far more numerous, diverse, and morphologically vari¬
able until die end of die Laramide Orogeny (~mid Eocene) with
North America as one continental landmass.
INTRODUCTION
This paper introduces new viviparid species that have
been recognized by the author for some time (Hartman,
1984). Publication of a monograph on fossil Viviparidae
is under construction, but still too far off to delay naming
of a number of taxa. The continental molluscan record of
the Cretaceous and Paleocene of Laramidia is more
diverse than typically known. Three new viviparid species
add to the known species richness during intervals of
geologic time that are, in part, less populated by gastro¬
pods. These species are also in ehronostratigraphic suc¬
cession in the Williston Basin of North Dakota, Montana,
and Saskatchewan. They include Campeloma acroterion
(Lancian into Puercan NALMA), Vivipams purgatorius
(Puercan into Torrejonian NALMA), and Vivipams
codomorphus (Torrejonian into Tiffanian NALMA), rang¬
ing from latest Cretaceous to early late Paleocene
(Maastriehtian to Selandian). These species occur in
the Hell Creek Formation, and Tulloek, Ludlow, and
Tongue River Members of the Fort Union Formation
in the United States, and the Ravenscrag Formation in
Canada (Figure 1).
MATERIALS AND METHODS
The taxa introduced are based on examination of fossils
from more than 1100 continental molluscan localities
(see Appendix 1; e.g., Hartman, 1984, Appendix 3, locality
register; Hartman, 1998, fossil localities; Hartman and
Roth, 1998, Appendix, Bighorn Basin locality register).
This effort originally comprised the dissertation studies of
die author (Hartman, 1984), but now includes many years
of subsequent field and museum studies (e.g., Hartman,
1989, 2004; Hartman and Kihm, 1992; Hartman and
Roth, 1998; Hunter et al, 1997; Hartman and Kirtland,
2002; Scholz and Hartman, 2007; Hartman et al., 2014).
Many of the specimens examined were collected by
others (e.g., Saskatchewan record) and are deposited at
institutions (see list under Abbreviations) visited by the
author. Important primary documentation was available
for many localities at institutional archives. When pos¬
sible, specimens were borrowed for additional study to
compare with in-house specimens. In addition, several
thousand photographs were taken of museum speci¬
mens to augment the borrowed material, and for use
in measuring specimens. Only a few measurements of
Cretaceous or Paleogene continental mollusks were
available from the literature. Most of these measure¬
ments are in reference to types, and do not provide a
Page 2
THE NAUTILUS, Vol. 129, No. 1
most Cretaceous and lower Paleogene strata. Chart based on Lund et al. (2002), Hartman et al. (2005) and is revised using
geochronologic and other data from Murphy et al. (2002), Clechenko et al. (2007), Secord (2008), LeCain et al. (2014), Ogg et al.
(2014), Vandenberghe et al. (2014).
basis for a quantitative analysis of shell form, plus the
methods by which the measurements were taken are
virtually always unknown (e.g., specimen orientation;
see measurement methods below). One of the purposes
in redescribing previously named taxa (Hartman, 1984)
was to provide a more rigorous quantitative database upon
which to compare fossil viviparid morphologies. The quan¬
titative measurement parameters used here to describe the
introduced species (summarized in later tables) are used in
diagnoses, descriptions, tables, mid graphs in the system¬
atic part of this report (and Hartman, 1984). Measurement
types (and their abbreviations) are given in Table 1. Mea¬
surement parameters are illustrated in Figure 2. Indi¬
vidual shell measurements are given in Appendix 2.
Abbreviations and Notes: W = number of whorls;
SD = standard deviation; min. = minimum; max.
maximum: n = number of specimens; loc. = number of
localities. K/Pg = Cretaceous-Paleogene, La = Lancian,
Pu = Puercan, To = Torrejonian, Ti = Tiffanian, Cf =
Clarkforldan, Wa = Was at chian, NALMA = North
American Land-Mammal “Ages.” HC = Hell Creek,
I^A = Lance, LU = Ludlow, TU = Tullock, LE = Lebo,
TR = Tongue River, FU = Fort Union, RA = Ravenserag.
AMNH-FI = American Museum of Natural History,
Fossil Invertebrates, New York; CSC = Geological Survey
of Canada, Ottawa; MCZ = Museum of Comparative
Zoology (Harvard University), Cambridge; UND-PC =
University of North Dakota, Paleontological Collection,
Grand Forks; UND-JHII = Hartman collection at UND;
UCMP = University of California Museum of Paleontol¬
ogy, Berkeley; UMPC = University of Minnesota, Paleon¬
tological Collection, Minneapolis; USNM-PAL = U.S.
Table 1. Measurement Abbreviations.
J.H. Hartman, 2015
Page 3
PAW - >
PWI=MWI - >
PSW2
y\
PSH1
v
y v
PSH2
PBH
PAH
PHI
MHI
FAW -
Coiling Axis
#W = 3.9
Figure 2. Gastropod Measurement Abbreviations. Figures 2A-D represent three standard photographic views from which speci¬
men measurements were acquired (see Table 1 for abbreviations). 2A. Apertural. 2B. Right lateral, growth-line angle (GLA) is
verticil! (one-dashed line) in species of Campeloma. 2C, Apertural flush. 2D. Apical, number of whorls (revolutions) counted with
use of a protractor. 2A and 2C. Campeloma acroterion. 2B. C. nebrascense nebrascense. 2D. Vivipams purgatorius.
National Museum of Natural History, Invertebrate Pale¬
ontology Collection, Washington, DC. See also abbrevia¬
tions in Appendix 1 (Locality Register).
All United States quadrangles are USGS 7.5-minute,
1:24,000, topographic maps, NAD27CONUS datum, with
20 ft (6.1 m) contour intervals. All Canadian maps are
Natural Resources Canada (was Department of Energy,
M ines and Resources), 15x30 minute, 1:50,000, topo¬
graphic maps, NAD27 datum, with 25 ft (7.6 m) con¬
tour intervals.
Two Appendices with supplementary data to this
work are posted online at http://nautilus.shellmuseum
.org. Appendix 1 is a Locality Register describing in
detail the localities discussed herein. Appendix 2 con¬
sists of three tables with measurements of all specimens
of the three species presented in this study.
Explanation of Figure 1
The correlation of strata presented in Figure 1 uses
a Hell Creek Formation duration of 1.8 million years
(Wilson, 2005). Although no single value can be correct,
the Fox Hills Formation in the western part of the
Williston Basin is approximated as the top of the Jeletzkytes
nebrascensis ammonoid zone (Landman et ah, 2007;
Vandenberghe et al., 2014). PETM (= Paleoeene- Eocene
Thermal Maximum) and the K/Pg I A (= Cretaeeous-
Paleogene Iridium Anomaly, or similar evidence, e.g.,
fern spike) are located approximately along a transect
from Bismarck to Marmarth (see Hartman et ah, 2002,
for citations; Clechenko et ah, 2007; Hartman et ah,
2014). The approximate locations and biohorizons of
type localities in Figure 1 are indicated by “?” on vertical
local biostratigraphic range lines. RPu2 (= Ravenserag
Formation, Puercan 2, etc.) symbols are overlays of the
approximate NALMA known or interpreted from the
Ravenserag Formation in the Cypress Hills of south¬
ernmost Saskatchewan (see text). Figure 1 notes include:
1) Campeloma acroterion is described from the upper¬
most Cretaceous Hell Creek Formation in its type
area of Hell Creek (Table 3), north of Jordan, Montana.
C. acroterion is also known from the lowermost Paleoeene
Page 4
THE NAUTILUS, Vol. 129, No. 1
Tullock Member (Fort Union Formation) in the western
p;irt of the Williston Basin. 2) C. acroterion is not well
known from the lowermost Paleocene of easternmost
Montana or North Dakota. C. acroterion is known from
a number of other uppermost Cretaceous formations in
the USA and Canada. 3) Viviparus purgatorius is
described from the lowermost Paleocene Tullock Member
in McCone County, Montana (see Table 7). V. purgatorius
is also known from the Bear Member (Fort Union For¬
mation) in the Crazy Mountains Basin, Montana, and
from the lower part of the Ravenserag Formation in
Saskatchewan. 4) V. codomorphus is described from
the Ludlow Member (Fort Union Formation) between
the tongues of the Boyce and Three V Tongues of the
Cannonball Formation in the Little Missouri River
valley. Slope County, North Dakota. 5) The type locality
of V. codomorphus is from the east flank of the Cedar
Creek Anticline in what the author interprets as
uppermost part of the Ludlow Member in easternmost
Montana. 6) V. codomorphus also occur at an ill-defined
location in the vicinity of Wibaux, Wibaux County,
Montana. This occurrence might place it in the Tongue
River Member. 7) The Fkalaka Member (Fort Union
Formation) does not occur between the Ludlow and
Tongue River Members (Fort Union Formation) along
the traverse of this profile (see Vuke et ah, 2007).
STRATIGRAPHY
The litho- and chronostratigraphy of the Cretaceous-
Paleogene boundary-interval strata in the northern Great
Plains and intermontane basins of the United States and
Canada are subjects of continual refinement (see citations
above and Figure 1). Field notes and other unpublished
information were used to augment collection and/or
published accounts of species records to better geo¬
graphically plot and place specimens in relative strati¬
graphic position. All type localities were examined by
the author, with the help of others, on more than one
occasion. Stratigraphic and sedimentologic data were
collected along with other elements of the local fauna.
The three taxa described are presented in stratigraphic
succession. Campeloma acroterion is known primarily
from the uppermost Cretaceous Hell Creek and Lance
Formations and also from formational equivalents. It
also is one of the few species in the northern Great
Plains known to cross the K/Pg boundary, with occur¬
rences in the Tullock Member of the Fort Union
Formation. Viviparus purgatorius is primarily known
from localities in the Tullock Member and is well
documented from the lower and middle part of the
Ravenserag Formation in Saskatchewan. The strati¬
graphic horizon of the type series of V codomorjdms
is the least well documented, in occurring at the top of
a large, grass-covered hill on the flank of an anticline.
Vuke et al. (2003) mapped this silerete-bearing section
as Tongue River Member, but previously mapped the
area as equivalent to the upper Ludlow Member (Vuke
et al., 1986). The author is inclined to map it as Ludlow
Member (see later discussion below and with Figure 1).
In North Dakota, V. codomorphus occurs between the
Boyce and Three V Tongues of the Cannonball Forma¬
tion in the upper part of the Ludlow Member (above the
T Cross lignite of Moore, and below the silerete horizon
at the top of the member [Hartman, 1993]).
A NALMA can be inteqmeted for each species. The
holotype of Campeloma acroterion is Lancian and ranges
to Puerean 2-3 (Figure 1, note 1). The species duration
is estimated to be about 3 million years. The holotype of
Viviparus purgatorius is Puerean 2-3 and likely ranges
to mid Torrejonian (Figure 1, note 3). The duration of
the species is also estimated to be about 3 million years.
The holotype of V. codomorphus , if the specimens are
from lowermost part of the Tongue River Member, is
lower Tiffanian (see Figure 1, notes 4, 5, 6). The holo¬
type may be approximately equivalent in age to the Ollie
North local fauna, which has yet to receive study to
assign a NALMA. Lithomarker correlation studies by
the author place the Ollie North about 20 m below the
silerete-White-Marker-Bed used to demarcate the top of
the Ludlow Member (Belt et al., 1984). The duration of
the record of V. codomorphus on the basis of mammals is
estimated at about 2.5 to 3.0 million years (depending
on lithostratigraphic interpretation).
SYSTEM ATICS
Class Gastropoda Cuvier, 1797
Subclass Orthogastropoda Ponder and Lindberg, 1996
Superorder Caenogastropoda Cox, 1960
Order Architaenioglossa Haller, 1892 [paraphyletie]
Superfamily Viviparoidea Gray, 1847 *
*Suprafamilial classification after de Bruyne (2003) and
Bouehet and Rocroi (2005) after The Taxonomicon
compilation (see sources therein).
Family Viviparidae Gray, 1847
Subfamily Lioplaeinae Gill, 1863
Campeloma Rafinesque, 1819
Etymology: campeloma (Greek), a bending , and (Greek)
a margin, “in exact keeping with the sigmoid character
of the aperture of all the species of the genus” (Call,
1883, p. 605).
Type Species: Campeloma crassula Rafinesque, 1819
(available; see, ICZN, 1999, Opinion 1931).
Campeloma acroterion new species
Figures 3-22
Diagnosis: Campeloma acroterion is distinguished from
other fossil species of Campeloma by the regular occur¬
rence of a well-marked revoking shoulder and shelf
(Fig. 15) a typical example of shoulder and shelf with
defined lip). The shelf is characteristically nearly per¬
pendicular to the axis of coiling (Figure 23), and the
shell material forming the shelf and shoulder is very
J.H. Hartman, 2015
Page 5
Hgures 3-22. Campeloina acroterion new species. All specimens were coated with ammonium chloride for photography. 3-6. USNM-
PAL 374598 [UND-JHH S0942], holotype (LI 151); 3) apertural. 4. right lateral. 5. Basal. 6. abapertural. 7-10. USNM-PAL 374600
[UND-JHH S0945], paratype-b (LI 151). 7. Apertural. 8. right lateral. 9. Basal. 10. abapertural. 9-12. USNM-PAL 374601 [UND-
JHH S0946], paratype-c (LI 151). 11. apertural. 12. right lateral. 13. Basal. 14. abapertural. 13-16. USNM-PAL 374599 [UND-JHH
S0943], paratype-a (L1151). 15. Apertural. 16. right lateral. 17. Basal. 18. abapertural. 19. UMPC 13602 [UND-JHH S0953],
paratype-d (L1151). 19. apertural. 20. UMPC 13603 [UND-JHH S0961], paratype-e (LI 151). 20. apertural. 21, 22. UCMP 37428,
paratype-f (LI 151). 21. Apertural. 22. basal.
Page 6
THE NAUTILUS, Vol. 129, No. 1
much thickened. Shouldering, with tendencies towards
shelf development, are common in Paleocene speci¬
mens of C. nebrascense, exemplified by those specimens
assigned to C. n. whitei. This subspecies is slightly larger
than C. acroterion and characteristically has a more
sloping shoulder (Figure 23), which is also more vari¬
ably expressed within a population.
Description: Shells medium in size, commonly rang¬
ing from 25 to 30 mm in maximum height, a few frag¬
mentary specimens indicate the species might attain
heights of up to 35-40 mm (Table 2); maximum number
Table 2. Vivipams acroterion. Measurement Summary (in min).
C. n. whitei
C. n. nebrascense
Figure 23. Outline comparison of Campeloma species (each
pair representing apertural and right lateral hews; not to scale).
23A, B. C. acroterion (L0053, S0217). 23C, D. C. nebrascense
whitei (L0349, UMPC 13609). 23E, F. C. nebrascense nebrascense
(L0023, UMPC 13604).
Table 3. Stratigraphic distribution of Campeloma acroterion
(examined specimens only).
Stratigraphic Unit
State/Province Localities (Lnos)
Ravenscrag Formation
Saskatchewan L0220, L0436
Ludlow Member, Fort Union Formation
North Dakota L0184?, L0185?
Lebo Member, Fort Union Formation
Montana L1412(LE?), L1420(LE?)
Tullock (Bear) Member, Fort Union Formation
Montana L0128
Tullock Member, Fort Union Formation
Montana L0010B, L0027, L0902(TU?), L0924,
L0925, L2951
Hell Creek Formation
Montana L0017A, L0020, L0021, L0053, L0182,
L0419?, L0560B, L0586A, L5087P,
LI 147, LI 149?, LI 150, L11511* *,
LI 152, LI 153, L1356, L1415, L2373,
L2949, L3960, L4321
North Dakota L0155, L0157, L0158, L0159?, L0162,
LG 163, L0164?, L0167?, L0169?,
L0170?, L0175, L0183?, L0923
Lance Formation
Wyoming L552B, L734, L735?, L738, L744, L880,
L3483, L3486?, L4289, L4292,
L4296, L4298, L4299
Code: ? = Questioned identification based on inadequate
preservation. (FM?) = Uncertain formational assignment.
*Type locality.
j.H. Hartman, 2015
Page 7
of whorls observed 7.4, with many specimens greater
than 6.2 whorls; measurements below are for “mature”
specimens greater than or equal to 6 whorls; average
width-to-maximum height ratio 0.60 (1 SD = 0.03, min. =
0.55, max. = 0.64, n = 7, loc. = 3); shell wall thickness
variable, ranging from relatively thin to robust; robustness
accentuated by substantial thickening on shoulders; apical
bp frequently broken. Shells turbiniform, subovately to
elongately conic, with an average mean spire angle for
specimens with a complete apex 51.7° (average 6.7 W;
1 SD = 3.7, min. = 43.5°, max. = 58.0°, n = 21, loc. = 2),
and for all specimens greater than 6 whorls 48.6° (1 SD =
4.4, min. = 38.0°, max. = 58.0°, n = 64, loc. = 6); spire
elevated, with an average mean spire width to mean
spire height ratio of 1.04 (average of 6.6 W; 1 SD =
0.07, min. = 0.93, max. - 1.14, n = 12, loc. = 2);
average spire height-to-body height ratio 0.51 (1 SD =
0.04, min. = 0.47, max. = 0.59, n = 8, loc. = 3); apical
tip rounded, teloconch whorls moderately convex, basal
periphery rounded; suture moderately impressed, accen¬
tuated by strongly developed shoulder and shelf. Growth
lines opisthoeyrt, weakly sigmoidal, finely to moderately
developed, with minima! coarsening with increasing
shell size. Surface sculpture of multiple revolving lirae
and some striae, varying from weakly to finely devel¬
oped, with the greatest density of lines just below
(abapical) whorl periphery; striae, as revolving lines of
closely spaced punctae and later as grooves, precede
revolving lirae on some specimens; shouldering is pre¬
ceded by one or two revolving lirae; a pair of lirae often
define the shoulder, with the area between the lirae
becoming much thickened and accentuated, producing
a shelf approximately perpendicular to coiling axis; shelf
sometimes with a revolving trough, which can be defined
by revolving lirae and striae; a revolving shallow sinus
can also occur abapical to the shelf; when the shelf is
more oblique to the axis of coiling, its appearance is as a
well-defined, usually rather angular, lip. Umbilicate; as a
narrow, but well defined opening. Aperture ovate, parietal
wall relatively thin; average aperture width to height
ratio measured in plane of aperture 0.75 (1 SD = 0.02,
min. = 0.69, max. = 0.78, n = 9, loc. = 2), measured in
plane of coiling axis 0.79 (1 SD = 0.03, min. = 0.72,
max. = 0.86, n = 14, loc. = 3), with a foreshortened
change in aperture height of about 8%. Individual speci¬
men measurements are given in Appendix 2 (for graphs
of parameters, see Hartman, 1984).
Etymology: acroterion (Greek), a promontory; as in a
projection or extremity; here used in reference to the
development of a well-defined shelf.
Type Specimens: Holotype, USNM-PAL 374598 (UND-
JHH S0942); paratype-a, 374599 (UND-JHH 0943);
paratype-b, 374600' (UND-JHH S0945); paratype-c,
USNM-PAL 374601 (UND-JHH S0946); paratype-d,
UMPC 13602 (UND-JHH S0953); paratype-e, UMPC
13603 (UND-JHH S0961); paratype-f, UCMP 37428;
all from Locality L1151 (see Type Locality and Appen¬
dix 1, Locality Register), Hell Creek Formation, Garfield
County, Montana.
Chresonyms-Nomenclatural Summary: Specimens
assigned to this taxon were most frequently identified
either as Campelonia nebrascense or C. n. ivhitei (see
Previously Illustrated Specimens and Appendix 2 for
examples of previous identifications).
Previously Illustrated Specimens now assigned to
C. acroterion
Figured specimen, USNM-PAL 9029a (L3340)
1883b White, pi. 28, fig. 4a (line drawing) [figured
as C. multilineata and herein identified as
C. acroterion?}.
Figured specimen, USNM-PAL 9029b (L3340)
1883b White, pi. 28, fig. 4b (line drawing) [figured
as C. multilineata and herein identified as
C. acroterion ?].
1883d White, pi. 27, fig. 7 (redrawing of White,
1883b) [reidentified as C. nebrascense ivhitei
by Russell (1931a) and affirmed by Tozer
(1956)].
Figured specimen, GSC 38365 (L0436)
1974 Russell, p. 48, 49, figs. 4a, b [figured as
C. n. ivhitei]; figured specimen, UMPC
12432 (LOO 10b).
1976 Hartman, pi. VII, figs. 10, 11 (unpublished
Master’s Thesis) [figured as C. n. ivhitei ].
Figured specimen, UMPC 12434 (L0027)
1976 Hartman, pi. VII, fig. 12 (unpublished Mas¬
ters Thesis) [figured as C. n. ivhitei ].
Figured specimen, UMPC 12433 (L0027)
1976 Hartman, pi. VII, fig. 15 (unpublished Mas¬
ters Thesis) [figured as C. n. ivhitei].
Figured specimen, UMPC 12435 (L0020)
1976 Hartman, pi. VII, fig. 13 (unpublished Mas¬
ter’s Thesis) [figured as C. n. ivhitei ].
Figured specimen, USNM-PAL 374598 (L1151)
1984 Hartman, pi. 1, figs. 1-4 (unpublished
Ph.D. Dissertation) [figured as holotype
of C. acroterion ].
Figured specimen, USNM-PAL 374600 (L1151)
1984 Hartman, pi. 1, figs. 5-8 (unpublished
Ph.D. Dissertation) [figured as paratype-b of
C. acroterion].
Figured specimen, USNM-PAL 374601 (L1151)
1984 Hartman, pi. 1, figs. 9-12 (unpublished
Ph.D. Dissertation) [figured as paratype-c of
C. acroterion].
Figured specimen, USNM-PAL 374599 (L1151)
1984 Hartman, pi. 1, figs. 13-16 (unpublished
Ph.D. Dissertation) [figured as paratype-a
of C. acroterion].
Figured specimen, UMPC 13602 (L1151)
1984 Hartman, pi. 1, figs. 17 (unpublished
Ph.D. Dissertation) [figured as paratype-d of
C. acroterion].
Page 8
THE NAUTILUS, Vol. 129, No. 1
Figured specimen, UMPC 13603 (LI 151)
1984 Hartman, pi. 1, figs. 18 (unpublished
Ph.D. Dissertation) [figured as paratvpe-e of
C. acroterion].
Figured specimen, UCMP 37428 (L1151)
1984 Hartman, pi. 1, figs. 19, 20 (unpublished
Ph.D. Dissertation) [figured as paratype-f of
C. acroterion].
Type Locality (LI 151): The type locality was discov¬
ered by Harley Garbani (Los Angeles County Museum)
as part of the search for dinosaurs and mammals in Hell
Creek country. J.H. Garbani and N.T. Greenwald col¬
lected a specimen of the type series of Campeloma
acroterion from “Harley’s shell bed” on July 7, 1976
(paratype-f, UCMP 37428; field number - JDA-76-7/
7-4; UCB locality number - D-7274). The author was
given directions to the locality by Garbani and studied
and collected the holotype, paratypes, and measured
specimens on: 1979 (July 26-27, UM; paratype-a, b, c,
d, e); 1982 (July 26, Hartman, UM; holotype); 1991
(July 20; Hartman, W.D. Peck, and D.E. Heinen, UND);
2002 (July 18; Hartman; R. Mattison, Wellesley College;
S. Uthus, UND); and 2008 (July 22; A.E. Bogan, D.P.
MeCollor, UND-EERC; K. Voegele, Concordia College;
and others).
Locality LI 151 occurs in the East Ried Coulee unit
(see Hartman et ah, 2014) of the Hell Creek Formation
at approximately 27.4 m (90 ft) (Hartman, 1998) below
the Z coalbed (after Archibald, 1982) that approximates
the top of the formation with the overlying Tulloek
Member of the Fort Union Formation (Hartman et ah,
2014). The locality is at an elevation of about 823 m
(2700 ft) on a north-trending promontory, near the
center of the south edge of the NW14 SW'A NE14
sec. 5, T. 20 N., R. 37 E., Trumbo Ranch Quadrangle
(1971), Garfield County, Montana (see map, Figure 24).
General Distribution: Recognizably distinctive speci¬
mens of Campeloma acroterion are found throughout
the Lance and Hell Creek Formations in the northern
Great Plains (Figure 24). Specimens are frequently a
quantitatively important element of the faunule and are
often the dominant gastropod. Similar specimens, with
distinctively shelved and shouldered whorls, also occur
in the overlying Fort Union (Tulloek, Bear, and lower
Ludlow Members [Tulloek Formation after others and
Frye, 1967]) and basal Ravenscrag Formations. The
poorly preserved specimens previously reported from
the upper part of the Livingston Group of the Crazy
Mountains Basin of south-central Montana are now
reassigned to horizons within the Hell Creek and Fort
Union Formations (Table 3; revised from Hartman,
1984, table 24).
Uppermost Cretaceous records from the Hell Creek
Formation in Custer, Dawson (or Wilbaux), Garfield,
Golden Valley, Meagher, McCone, Rosebud, Wibaux
(or Dawson), and Yellowstone Counties, Montana; from
Bowman, Emmons, Grant, Morton, Sioux, and Slope
Counties, North Dakota; from the Lance Formation in
Niobrara County, Wyoming; from the Paleocene Fort
Union Formation members including the Bear Member
in Wheatland County, Montana; from the Tulloek
Member in Daniels, Garfield, and McCone Counties,
Montana; with uncertainty from the Lebo Member in
Wheatland County; from the Ludlow Member in Slope
County, North Dakota; and from the Ravenscrag For¬
mation in Cypress Hills in southernmost Saskatchewan.
Cadastral location data for examined C. acroterion are
given in Hartman (1984, table 25).
Frye (1967, 1969) lithostratigraphically organized the
Hell Creek Formation into several members. Identifica¬
tion of Frye’s (1967) specimens shows C. acroterion
to occur in the Pretty Butte (L0183?), Huff (L0170?,
L0175), and Fort Rice (L0155, L0157, L0158, L0159?,
L0162, L0163, L0164?, L0167?, L0169?) members.
Carlson (1979; personal communication, 15 June 1981),
Muq)hy et al. (2002), and others questioned the utility
and ability of others to recognize these subdivisions.
Butler (1980), Butler and Hartman (1999), Lund et al.
(2002) and Hartman et al. (2014) suggested that a Hell
Creek Formation architecture exists and could be corre¬
lated in both surface exposures and subsurface logs. The
member contacts proposed by Frye (1967, 1969) may
not be as useful as he hoped, but his lithofaeies have
been recognized significantly beyond the limits of their
type sections (Butler, 1980; Hartman et ah, 2014).
Subfamily Viviparinae Gray, 1847
Viviparus Montfort, 1810
Etymology: viviparus, that brings forth its young alive.
Type Species: Viviparus fliwiorum Montfort, 1810
(= Helix vivipara Linnaeus, 1758) (available, see ICZN,
1959, Opinion 573).
Viviparus purgatorius new species
Figures 25-66
Diagnosis: Viviparus purgatorius is distinguished from
other fossil Viviparinae on the basis of a suite of charac¬
ters including a much greater mean spire angle, much
more inclined growth lines, strongly keeled and erenn-
lated periphery (in adult specimens), and development
of nodes in adult forms. Only V. retusus growth lines
are as acutely inclined to the periphery, but it differs
from V. purgatorius in being much smaller in size for
the same number of whorls, in possessing a depressed
spire, and in lacking significant sculpture (compare
selected Viviparus morphologies, see Figure 67).
Viviparus formosus resembles V. purgatorius in main¬
taining a large mean spire angle for a large shell size,
but it differs from V purgatorius in possessing less
inclined growth lines, and a more elevated spire, in lack¬
ing nodes, and in lacking a crenulated, strongly angular
basal periphery. Viviparus sp. form-W also resembles
V. purgatorius in maintaining a large mean spire angle
J.H. Hartman, 2015
Page 9
Figure 24. Biogeography of Campeloma acroterion localities in the northern Great Plains and intermontane basins ol the USA
and Canada. Upland areas (roughly Laramide uplifts) are shown in pink, with preserved Paleogene and younger sediments preserved
in basins shown in yellow (see Abbreviations.)
for a large shell size, but differs from V. purgatorius
most obviously in possessing a much more rounded basal
periphery, but also in lacking nodes, a erenulated periph¬
ery, and in possessing a larger last whorl expansion ratio.
Other noduliferous species proximal to V. purgatorius
are Paleocene Viviparus codomorjrhus and Cretaceous
V. thompsoni. Viviparus codomorphus differs from
V. purgatorius in possessing a more blunted apex, in
the infrequency of adapical node development (only
one specimen has nodes developed), and in lack¬
ing a erenulated periphery. V. thompsoni differs from
V purgatorius in possessing a distinctive pattern of
nodes and node ridges. Viviparus purgatorius also dif¬
fers from V. thompsoni in lacking any significant basal
surface sculpture (other than spiral lirae) on the latter
whorls of larger specimens.
Description: When this species was originally recog¬
nized and described informally, no undeformed large
specimens of this taxon existed, prohibiting a rigorous,
numerically calibrated description. Essentially complete,
undistorted large specimens were subsequently discov¬
ered by D.L. Lofgren (see Appendix 2).
Shells relatively large in size, attaining a maximum
height of about 35 mm; maximum number of whorls
observed 6.9; measurements given below are for mature
specimens from 5 to 6 greater whorls; shell walls rela¬
tively thin (Table 4). Shells subtrochiform, globosely
Page 10
THE NAUTILUS, Vol. 129, No. 1
mm
|.H. Hartman, 2015
Page 1 1
V meeki
Figure 67. Outline comparison o i Viviparus species (apertural views, not to scale). 67A-C. Blunted apex. 67A. V. campaniformis
(L0070; S0517). 67B. V returns (L0435; USNM-PAL 2155). 67C. V codomorphus (L0049; USNM-PAL 374604). 67D. Pointed
apex, rounded whorls and periphery. 67D. V. leai (L2305; USNM-PAL 2154). 67E-I. Pointed apex, subtrochiform marginal
outline. 67E. V peculiars (LQ349; S0309). 67F. V meeki (L0953; USNM-PAL 374985). 67G. V. meeki, narrower form (L0953;
USNM-PAL 374986). 67H. V.formosus (L0435; USNM-PAL 2159). 671. V purgatorius (L6460, S10441).
conic, with an average mean spire angle of 68.5°
(#W > 5.5: aver #W = 5.7 W, 1 SD = 9.26, min = 61.9°,
max = 75.0°, n =2, loc = 2; #W > 5.0: MSA = 66.6°,
1 SD = 5.1, min. = 61.9°, max. = 73.5°, n = 4, loc. = 3;
#W > 4.6: MSA = 68.8°, 1 SD = 6.5, min. = 81.5°, max. =
60.0°, n = 16, loc. = 6); average last whorl expansion
ratio of 1.88 (1 SD = 0.03, min. = 1.84, max. = 1.90,
n = 4, loc. = 2); apical tip bluntly pointed, apical whorls
not substantially produced; teloconch whorls broadly
convex, with an average mean spire width to mean spire
height ratio of 3.3 (aver of 5.5 W: 1 SD = 1.00, min.
i .52. max. = 1.51, n = 6, loc. = 1); spire height-to-body
height ratio 0.30 (1 SD = 0.04, min. = 0.22, max. =
0.30, n = 3, loc. = 1); slight adapical shouldering proximal
to the suture; upper spire and basal periphery carinate-
angular, with basal periphery often strongly angular;
suture slightly to very slightly impressed. Growth lines
prosocline, very strongly inclined at about 38°( #W > 5.0:
Figures 25-66. Viviparus purgatorius new species. Specimens from Localities LOOlOb and L0214 were coated with ammo¬
nium chloride for photography. 25-28. GSC-IP 76957, hypotype (L0214). 25. apertural. 26. right lateral. 27. basal. 28. apertural.
29. GSC-IP 76958, hypotype (L0214). 29. abapertural (specimen slightly distorted). 30. UMPC 12427 [UND-JHH S0754],
paratype-a (LOOlOb). 30. apertural (specimen distorted). 31-34. USNM-PAL 374614 [UND-JHH S0755], paratype-b (LOOlOb).
31. apertural. 32. right lateral. 33. abapertural. 34. basal (specimen slightly distorted, adaperturally depressed). 35. UMPC 13616
[UND-JHH S0757], paratype-d (LOOlOb). 35. apertural. 36. UMPC 13617 [UND-JHH S1025], paratype-e (LOOlOb). 36. basal
(specimen distorted). 37, 38. UMPC 12428 [UND-JHH S0756], paratype-c (LOOlOb). 37. apertural. 38. right lateral. 39-11. USNM-
PAL 374613 [UMPC 12429; UND-JHH S0753], paratype-f (was unpublished holotype) (LOOlOb). 39. right lateral. 40. abapertural.
41. basal (specimen axially compressed). 42—45. UND-JHH S 10443, figured young specimen, topotype (L6460). 42. apertural. 43. right
lateral. 44. abapertural. 45. apical. 46-50. USNM-PAL requestl [UND-JHH S10441], holotype (L6460). 46. apertural. 47. right lateral.
48. Abapertural. 49. apertural flush. 50. apical. 51-53. USNM-PAL will request2 [UND-JHH S 10445], paratype-g, topotype (L6460).
51. apertural. 52. right lateral. 53. apical. 54—57. USNM-PAL request [UND-JHH S10439], paratype-h, (L6460). 54. apertural
oblique. 55. right lateral. 56. Abapertural. 57. apertural flush. 58-62. UND-PC 16163 [UND-JHH'si0446], paratype-I (L6460).
58. apertural. 59. right lateral. 60. abapertural. 61. apertural flush. 62. apical. 63-66. UND-PC 16164 [UND-JHH S10447], paratype-j
(L6460). 63. apertural. 64. right lateral. 65. apertural flush. 66. apical.
Page 12
THE NAUTILUS, Vol. 129, No. 1
Table 4. Viviparus purgatorius. Measurement Summary
(in mm).1*
*On specimens with > 5 #W.
GLA = 41.2°, 1 SD = 10.1, min. = 31.0°, max. = 53.0°,
n = 5, loc. = 2; nearly straight, but apparently with a slight
adapertural arch proximal to suture; fine-to-moderate
development, becoming coarser on latter whorls of larger
specimens. Sculpture consists, in part, of a pair of slightly
raised revolving ridges of approximately equal promi¬
nence and nearly equally spaced between ad- and abapi-
cal sutures, strongly reminiscent of some specimens of
V meeki; revolving ridges appear to be most clearly
seen on upper spire whorls, becoming more obscure with
increasing shell height; other revolving lirae may be
approximately equal in prominence to the pair of raised
ridges, and may thus obscure the basic sculptural simi¬
larity with V. meeki; Paleogene revolving sculpture is
obscure to absent, but some lirae occur below (abapieal)
basal periphery, are of approximately equal prominence,
and are best seen on smaller specimens; a few specimens
show a slight, basal surface ridge without nodes; basal
periphery keeled and slightly to moderately crenulated on
larger specimens, reminiscent of V. thompsoni; crenula-
tion may be enhanced by lateral distortion, but its exis¬
tence, although seeming to be somewhat variable, appears
definitely to be a feature of this species. Without umbi¬
licus, with thick inner apertural wall and umbilical
area covered by a distinct, relatively wide and thickened
eolumellar callus. Aperture poorly known, ovate, tapers
adapieally, with a rather strong outer lip angulation at
the whorl periphery; parietal lip thins along basal whorl;
aperture plane strongly inclined; average aperture width -
to-height ratio measured in plane of aperture 0.78
(1 SD = 0.09, min. = 0.9, max. = 2.9, n = 4, loc. = 1),
measured in plane of coiling axis 1.62 (1 SD = 0.85,
min. = 0.91, max. = 2.86, n = 4, loc. = 1), with a
foreshortened change in aperture height of about
48%. Individual specimen measurements are given in
Appendix 2 (for graph of parameters, see Hartman, 1984).
Etymology: purgatorius , named after Purgatory Hill
(L0010), McCone County, Montana, the discovery loca¬
tion for the species.
Chresonyms-Nomenclatural Summary
1974 Viviparus formosus (Meek and Hayden): Russell,
p. 46, 47, figs. 3e, f.
1976 Viviparus aff. V thompsoni White: Hartman,
p. 116-119, 176, pi. VII, figs. 1-5 (unpublished
M.S. Thesis).
1984 Viviparus purgatorius Hartman, p. 445—454, pi. 8,
figs. 1-24 (unpublished Ph.D. Dissertation).
Previously Illustrated Specimens now assigned to
V. purgatorius
Figured specimen, GSC 38361 (L0224)
1974 Russell, p. 46, 47, figs. 3e, f (figured as
V. formosus) .
Figured specimen, GSC 76957 (L0214)
1984 Hartman, pi. 8, figs. 1^1 (unpublished Ph.D.
Dissertation) [figured as V. purgatorius ].
Figured specimen, GSC 76958 (L0214)
1984 Hartman, pi. 8, figs. 5, 6 (unpublished Ph.D.
Dissertation) [figured as V. purgatorius].
Figured specimen, UMPC 12427 (LOOlOb)
1976 Hartman, pi. VII, figs. 1, 2 (unpublished Mas¬
ter’s Thesis) [figured as V aff. V. thompsoni ]
1984 Hartman, pi. 8, figs. 7, 8 (unpublished
Ph.D. Dissertation) [figured as paratype-a
of V. purgatorius ].
Figured specimen, UMPC 12428 (LOOlOb)
1976 Hartman, pi. VII, figs. 4, 5 (unpublished Mas¬
ter’s Thesis) [figured as V aff. V. thompsoni]
1984 Hartman, pi. 8, figs. 17-20 (unpublished
Ph.D. Dissertation) [figured as paratype-c
of V. purgatorius].
Figured specimen, UMPC 12429 (LOOlOb)
1976 Hartman, pi. VII, fig. 3 (unpublished Master’s
Thesis) [figured as V aff. V. thompsoni]
1984 Hartman, pi. 8, figs. 21-24 (unpublished
Ph.D. Dissertation) [figured as holotype of
V. purgatorius].
Figured specimen, UMPC 13617 (LOOlOb)
1984 Hartman, pi. 8, figs. 9, 10 (unpublished
Ph.D. Dissertation) [figured as paratype-e
of V. purgatorius].
Figured specimen, UMPC 13616 (LOOlOb)
1984 Hartman, pi. 8, figs. 11, 12 (unpublished
Ph.D. Dissertation) [figured as paratype-d of
V purgatorius].
Figured specimen, USNM-PAL 374614 (LOOlOb)
1984 Hartman, pi. 8, figs. 17, 20 (unpublished
Ph.D. Dissertation) [figured as paratype-b
of V purgatorius].
Type Specimens: Holotype, USNM-PAL 611016
(UND-JHH S10441, L6460); paratype-a UMPC 12427
(UND-JHH S0754, LOOlOb); paratype-b, USNM-PAL
J.H. Hartman, 2015
Page 13
374614 (UND-JHH S0755, LOOlOb); paratype-c, UMPC
12428 (UND-JHH S0756, LOOlOb); paratype-d, UMPC
13616 (UND-JHH S0757, LOOlOb); paratype-e, UMPC
13617 (UND-JHH S1025, LOOlOb); paratype-f, USNM-
PAL 374613 (UMPC 12429, UND-JHH S0753, LOOlOb);
paratype-g, USNM-PAL 611017 (UND-JHH S10445,
L6460); paratype-h, USNM-PAL 611018 (UND-JHH
S 10439, L6460); paratype-I, UND-PC 16163 (UND-
JHH S10446, L6460); and paratype-j, UND-PC 16164
(UND-JHH S 10447, L6460) from the Tullock Member,
Fort Union Formation, McCone County, Montana (see
Type Locality and Appendix 1, Locality Register).
Reference Specimens: Hypotypes, GSC-IP 76957,
76958, Locality L0214; Ravenscrag Formation, Big Muddy
Valley, on Minton map (1974), Saskatchewan.
Type Locality (L6460): The type locality of Viviparus
purgatorius was discovered August 12, 1994 by Donald
L. Lofgren (Raymond M. Alf Museum) and the holotype
and paratypes-h and -j were collected from the Race¬
track Ridge locality on August 12, 1994, by D.L. Lofgren
and W.A. Clemens (University of California-Berkeley)
(field 94DLL8-12-2, Alf 194161), along with other mea¬
sured specimens (see Appendix 2). The specimens were
subsequently donated to the UND-PC collections.
Paratype-i was collected by Hartman on September 16,
1995 (L6460b; [a-e sites were recognized). Paratype-g
was collected on July 27, 2000 by J.H. Hartman,
D. Lamb, D.L. Lofgren, A.C. Raser, G.C. Winsinger.
These and other specimens were collected from the
Tullock Member of the Fort Union Formation about
36.6 m above base of the MCZ coalbed of Lofgren
(1995, McGuire Creek Z) representing here the base
of the formation, marking the contact with the underlying
Hell Creek Formation. The locality is at an elevation of
about 742.8 m (2437 ft) in the W V6 NW14 NEV4 sec. 16,
T. 21 N., R. 43 E., on the Nelson Creek Bay Quad. (1973)
in McCone County, Montana (see map, Figure 68).
Reference Locality (LOOlOb): Discovery location para-
types (a-f) of Viviparus purgatorius were collected by
R.E. Sloan and J.H. Hartman in 1974 from Purgatory
Hill at an elevation of about 751.3 m (2478 ft), located
in SE14 NE14 SEJ/4 NW14 SW!4, sec. 36, T. 23 N., R.
43 E., Bug Creek Quad. (1973), McCone County,
Montana. Locality LOOlOb is located on the north-facing
exposures near the top of the butte in a north-south
trending channel capped by a sandstone monolith. The
channel cut is stratigraphically about 30.5 m above the
MCZ coalbed representing the base of the Tullock
Member, Fort Union Formation (see also Bell, 1965,
measured section E). Locality LOOlOb is also the a
source of late Puerean mammals interpreted to be Pu2-3
NALMA reported by Van Valen and Sloan (1965) and
frequently thereafter (see Hartman, 1978; Archibald, 1987;
Lofgren, 1995) (see Figure 68).
Literature Distribution: Viviparus purgatorius was
reported by Hartman (1976) as V. aff. V. thompsoni (see
Chresonyms) from the Tullock Formation in McCone
County (LOOlOb), Montana. The only other previous
report of specimens assignable to this taxon were made
by L.S. Russell (in Fraser and others, 1935; Russell,
1974) from the Ravenscrag Formation in southernmost
Saskatchewan. Russell’s identifications, now assigned
to V. purgatorius , included V. forrnosus, V leichji, and
V. meeki (— V. trochiformis) .
General Distribution: Viviparus purgatorius was known
primarily from distorted and/or incomplete specimens
throughout its range in northeastern Montana in the
Tullock Member in McCone (LOOlOb, L0027?) and
Garfield (L2951) Counties; southern Saskatchewan in
the Ravenscrag Formation (L0209?, L0212?, L0214,
L0217?, L0219, L0220, L0223?, L0224, L0225, L0229?,
and L0436); and from a queried occurrence in south-
central Montana in the Bear Member (Fort Union For¬
mation) in Wheatland County (L0128; see Hartman and
Krause, 1993); and from a queried occurrence in the
Tongue River Member of the Fort Union Formation in
Musselshell County (LI 458?) (Figure 68; Table 5) (see
Hartman, 1984, table 60 for a cadastral record of occur¬
rences). The record from the Tongue River Member
may be referable to V meeki , which, if true, would
restrict the age of V purgatorius to early Paleocene
(~Pu2-3; LOOlOb) in Montana. The occurrences of
V. purgatorius in the Ravenscrag Formation stratigra¬
phically range from a documented earl)’ Paleocene locality
(Pu2; L0220) in the lower part of the Ravenscrag For¬
mation in the area of Anxiety Butte, to an unresolved
Paleocene age in the upper middle part of the forma¬
tion just above the Willowbunch coal in the Big Muddy
Valley area. On the basis of palynological data. Sweet
(1978) stated that the strata of the Willowbunch coal
zone may be correlative with the Estevan or Boundary
coal sequence in the Estevan area, which would prob¬
ably make the occurrence of V. purgatorius in the
Big Muddy Valley at least mid Paleocene (Ti2) (see
Whitaker and others, 1978, pi. 2, Regional Correlation
Sections A-A')-
Discussion - Viviparus purgatorius : Although rec¬
ognizably distinct, this species was first found plastically
deformed. The style of deformation and preservation
of Tullock Member and Ravenscrag Formation speci¬
mens was very similar suggesting similar shell structure.
V. purgatorius could be identified on the basis of the
features mentioned in the Diagnosis and Description,
but the deformed specimen variable whorl convexity,
growth line angle, and mean spire angle could make
positive identification difficult. The discovery of numer¬
ous undistorted specimens at Locality L6460 at Racetrack
Ridge confirmed the interpretation of the proposed spe¬
cies (Hartman, 1984) and provided measurements not
previously available (Appendix 2).
Probably all of the Ravenscrag Formation speci¬
mens identified by Russell (in Fraser and others, 1935;
Russell, 1974) as V. forrnosus and V leichji , and many
THE NAUTILUS, Vol. 129, No. 1
Page 14
» 52°) 11Q°
45 | Cypress
: HiHs^-, a]
C T ° W, d
! ° '
RA
. . 102°|
Turtle Mountains
50 100 mi
50 100 km
Biogeography of Viviparus
purgatorius study
area localities
Single-Species (Study Area Maps)
type Locality
positive identifications
queried identifications
cf. and cf? identification
positive unevaluated identifications ... g
queried, cf. and cf.? unevaluated identifications □
aff identifications .
aff. unevaluated identifications . q
104°03'1
Figure 68. Biogeography of Viviparus purgatorius localities in the northern Great Plains and intermontane basins of the USA and
Canada (see Figure 24 for map notes).
Table 5. Stratigraphic distribution of Viviparus codomorphus
(examined specimens only).
Stratigraphic Unit State Localities (Lnos)
Fort Union Formation undivided
Montana L0936
Ludlow (Upper) (Slope) Member, Fort Union Formation
North Dakota L4232A, L4232B
Ludlow Member, Fort Union Formation
Montana L00491*, L0939
*Type locality; Area interpreted by Vuke et al. (2003) to be
Tongue River Member (see text).
identified as V. meeki (= V trochiformis), are referable
to V. purgatorius. The specimens assigned to these
younger Paleocene taxa are all very similar and are at
once distinguished from V. purgatorius by possessing
an umbilicus. They also differ in possessing a more
pointed apex, less inclined growth lines, and a less
angular periphery.
Small and incomplete specimens of V. purgatorius could
be difficult to separate from either V. campaniformis or
V. retusus, as all three species have a blunt apex and
a more or less carinate periphery. Small specimens of
these species should not be positively identified unless
larger specimens are present in the population. The
adult morphologies are, however, quite dissimilar.
J.H. Hartman, 2015
Page 15
Discussion - Viviparus cf. V. purgatorius: All of the
specimens assigned to V cf. V. purgatorius are from the
Ravenscrag Formation (Localities L0214 and L0218)
and appear to represent a morphology distinct from
V purgatorius. Viviparus cf. V. purgatorius differs from
V. purgatorius in being notably more striate, particularly
on the last whorl of larger specimens, in possessing a
less angular periphery with increasing shell size, and
in having a more impressed suture. They are similar in
possessing multispiral lirae on upper spire whorls, in
lacking an umbilicus, and in possessing strongly inclined
growth lines. All of the larger specimens assigned to
V. cf. V. purgatorius are lacking apical whorls. V. leai
is the only other Paleocene viviparid characterized by
revoking striae or punctae, but unlike V. purgatorius, it
is umbilieate.
Viviparus codomorphus new species
Figures 69-92
Diagnosis: Viviparus codomorphus is distinguished
from other fossil Viviparinae on the basis of a suite of
characters including its blunted apical tip (Figure 69),
absence of an umbilicus, and mean spire angle for a
given number of whorls. Viviparus codomorphus is
also represented by an atypical specimen (Figure 91,
UMPC 13611) displaying adapical nodes and a basal
surface ridge. Species with similar morphology are
V. campaniformis and V meeki (see outline comparisons,
Figure 67).
Viviparus campaniformis differs from V. codomorfdms
in its greater rounding of apical whorls, more flanged
and carinate appearance on upper spire whorls, smaller
mean spire angle and last whorl expansion ratio in larger
specimens, in attaining a greater shell height for the
same number of whorls, and somewhat more inclined
growth lines. V. meeki differs from V. codomorphus in
having a pointed apex, attaining an equivalent or some¬
what smaller size for a greater number of whorls, and in
possessing an umbilicus.
Description: Shell medium in size, commonly ranges
from 23 to 28 mm in maximum height, rarely exceeding
30 mm; maximum number of whorls observed 5.8, with
most specimens ranging from 5.0 to 5.5; measurements
given below are for “mature” specimens greater than or
equal to 5.0 whorls; average width-to-maximum height
ratio 0.82 (1 SD = 0.04, min. = 0.72, max. = 0.90, n =
28, loe. = 4); shell walls moderately thick, frequently
robust (Table 6). Shells suhtrochiform, broadly conic,
with an average mean spire angle for specimens with a
complete apex of 62.0° (average of 5.4 W; 1 SD = 3.0,
min. = 56.5°, max. = 71.0°, n = 26, loe. = 5), and for all
specimens greater than 5.0 whorls of 60.5° (1 SD = 3.3,
min. = 53.5°, max. = 71.0°, n = 46, loc. = 5); spire some¬
what elevated, with an average mean spire width to mean
spire height ratio of 1.34 (average of 5.4 W; 1 SD = 0.08,
min. = 1.25, max. = 1.54, n = 22, loc. = 5); last whorl
expansion ratio of 1.65 (1 SD = 0.06, min. = 1.54, max. =
1.80, n = 46, loc. = 5); spire height-to-body height ratio
0.38 (1 SD = 0.06, min. = 0.28, max. = 6.48, n = 30,
loc. = 5); apical tip slightly blunted, rounded at extreme
tip, teloconeh whorls broadly convex, with a slight but
consistent oblique shoulder proximal to suture; upper
spire whorl periphery angular and carinate, becoming
more subangular with increasing shell size, but usually
maintaining evidence of a peripheral keel; suture slightly
to moderately impressed. Growth lines prosoeline, with
an average growth-line angle of about 23° (approximately
determined: mean = 22.7°, 1 SD = 4.2, min. = 15.0°,
max. = 31.0°, n = 25, loc. = 5); nearly straight across
midwhorl, although frequently with a slight adapertural
arch and usually curved over slight shoulder proximal to
suture; finely to moderately developed, becoming some¬
what more coarse with increasing shell size. Sculpture
consists of a pair or more of slightly raised ridges and very
obscure to absent secondary revolving lirae and striae;
peripheral carina on upper spire, often accentuated by a
slightly adapical revolving stria; revolving sculpture below
whorl periphery apparently rare or absent; on one speci¬
men (not obviously developed on any others) with par¬
ticularly well-developed sculpture, low elongate nodes
are developed on the adapical spiral ridge and are
expressed only on the last third of the last whorl of
5.5 whorl specimens; node axes appear to parallel the
revolving ridge; on the basal surface of the same speci¬
men, one rather wide nodular ridge is developed. With¬
out umbilicus, umbilical area covered by a thickened,
but not wide, columellar lip. Aperture broadly ovate,
tending towards subquadrate depending on the degree
to which the peripheral angularity is expressed on the
outer lip; parietal lip not appreciably thinned; average
aperture width-to-height ratio measured in plane of
aperture 0.89 (1 SD = 0.04, min. = 0.83, max. = 0.99,
n = 28, loc. = 4), measured in plane of coiling axis 1.20
(1 SD = 0.11, min. = 1.04, max. = 1.45, n = 28, loc. = 4),
with a foreshortened change in aperture height of about
29%. Individual specimen measurements are given in
Appendix 2 (for graphs of parameters, see Hartman, 1984).
Etymology: codomorjdms (Greek), codono (kodon),
bell; morpho (morphe), form, shape; the shape of a bell.
Type Specimens: Holotype, USNM-PAL 374602 (UND-
JHH S0892); paratype-a USNM-PAL 374602 (UND-
JHH S0200); paratype-b, USNM-PAL 374602 (UND-JHH
S0202); paratype-c, USNM-PAL 374606 (UND-JHH
S0894); paratype-d, UMPC 13610 (UND-JHH S0900);
and paratype-e, UMPC 13611 (UND-JHH S0984) are all
from Locality L0049 (see Type Locality and Appendix 1,
Locality Register), Tongue River Member (after Vuke
et ah, 2003; Ludlow Member after Hartman, 1993; and
suggested herein) of the Fort Union Formation, Fallon
County, Montana.
Chresonyms-Nomenclatural Summary
1921 Viviparus ( Paludotrochus ) trochiformis : Cossmann,
pi. V, figs. 1, 2.
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THE NAUTILUS, Vol. 129, No. 1
Figures 69-92. Viviparus codomorphus new species. All specimens were coated with ammonium chloride for photography.
69-72. UMPC 13610 [UND-JHH S0900], paratype-d (L0049). 69. apertural. 70. right lateral. 71. Basal. 72. abapertural.
73-76. USNM-PAL 374603 [UND-JHH S0200], paratype-a (L0049). 73. apertural. 74. right lateral. 75. basal. 76. abapertural.
77-80. USNM-PAL 374605 [UND-JHH S0894], paratype-c (L0049). 77. apertural. 78. right lateral. 79. basal. 80. abapertural.
81-84. USNM-PAL 374602 [UND-JHH S0892], holotype (L0049). 81. Apertural. 82. right lateral. 83. basal. 84. abapertural.
85-88. USNM-PAL 374604 [UND-JHH S0202], paratype-b (L0049). 85. apertural. 86. right lateral. 87. basal. 88. abapertural.
89-92. UMPC 13611 [UND-JHH S0984], paratype-e (L4232). 89. Apertural. 90. right lateral. 91. Basal. 92. abapertural (note
node development on basal periphery).
J.H. Hartman, 2015
Page 17
Table 6. Viviparus codomorphus Measurement Summary
(in mm).1*
Table 7. Stratigraphic distribution of Viviparus purgatorius
(examined specimens only).
Stratigraphic unit
State/Province Localities (Lnos)
Tongue River Member, Fort Union Formation
Montana L1458?
Tullock (Rear) Member, Fort Union Formation
Montana L0010B, L0027?, L2951, L64601* *
Montana (Bear) L0128?
Ravenscrag Formation
Saskatchewan LQ2Q9?, LQ212?, L0214, L0217?,
L0219, L0220, L0223?, L0224,
L0225, L0229?, L0436
Code: ? = Questioned identification based on inadequate
preservation.
*Type locality.
1984 Viviparus codomorphus Hartman, p. 331-340,
pi, 3, figs. 9-32 (unpublished Ph.D. Dissertation).
Previously Illustrated Specimens now assigned to
V. codomorphus
Figured specimen, repository and locality not known.
1921 Cossmann, pi. V, figs. 1, 2 (figured as Viviparus
trochiformis) .
Holotype, USNM-PAL 374602 (L0049)
1984 Hartman, p. 575, pi, 3, figs. 21-24 (unpub¬
lished Ph.D. Dissertation) [figured as holotype].
Paratype, USNM-PAL 374603 (L0049)
1984 Hartman, p. 575, pi, 3, figs. 13-16 (unpublished
Ph.D. Dissertation) [figured as paratype-a],
Paratype, USNM-PAL 374604 (L0049)
1984 Hartman, p. 575, pi, 3, figs. 2.5-28 (unpublished
Ph.D. Dissertation) [figured as paratype-b],
Paratype, USNM-PAL 374605 (L0049)
1984 Hartman, p. 575, pi, 3, figs. 17-20 (unpublished
Ph.D. Dissertation) [figured as paratype-c].
Paratype, UMPC 13610 (L0049)
1984 Hartman, p. 575, pi, 3, figs. 9-12 (unpublished
Ph.D. Dissertation) [figured as paratype-d].
Paratype, UMPC 13611 (L4232)
1984 Hartman, p. 575, pi, 3, figs. 29-32 (unpublished
Ph.D. Dissertation) [figured as paratype-e].
Type Locality (L0049): Mr. Marshall E. Lambert,
longtime curator of the Carter County Museum, Ekalaka,
Montana, very kindly provided the necessary information
and encouragement to relocate a locality he collected in
August 1960, but for which he had only vague location
information: “North of Baker” locality, 20 to 22 miles
[32.2-35.5 km] north of Baker on Highway 7 on a conical
butte. The locality was known to locals and the landowner,
as it was originally excavated for chicken feed. The loca¬
tion is near the top of a very broad, largely grass-covered,
and silcrete-boulder-strewn butte. The specimens were
collected from a shallow pit at the north end, now nearly
grassed over. The shell-bed thickness is only partially
exposed, but is greater than 1-m thick when excavated
and may extend over 60 m on a north-south trend.
The type series of Viviparus codomorphus was col¬
lected on July 8, 1977 by Hartman and j.A. Milske
(UM) at an elevation of about 1027 m (3370 ft) from the
Shell Butte locality (L0049), located near the center of
the NE!4 NEVa SW!4 sec. 13, T. 10 N., R. 59 E„ Shell
Butte Quad. (1981), Fallon County, Montana (Figure 93).
The locality is mapped as Tongue River Member (Vuke
et ah, 2003), and is provisionally placed by the author
at about 46 m (150 ft) above the top of the underlying
Hell Creek Formation. The nearest available exposures
of the Fort Union-Hell Creek formational contact are
located about 2 km to the soudiwest. The exact strati¬
graphic placement of the locality depends on a more
precise knowledge of the amount and change of dip in
the strata as the beds flatten to the east on the east flank
of the Cedar Creek Anticline. Stratigraphic studies by
the author suggest that the locality may actually occur in
the Ludlow Member (Hartman, 1993). Vuke et al. (1986)
previously mapped the Shell Butte area as FU3 and cor¬
related it with the Slope Formation of North Dakota.
Measured and study specimens were also collected
on May 3, 1978, by Hartman, L. Doull, W. Hughes,
L. Huntley, A. Krafft, E. Olson, A. Rautman, S. Russell,
A. Snodgrass, and M. Timmerman; June 30, 1978, by
Hartman and R.C. Holtzman; August 22, 1988, by
Hartman; and July 6, 7, 1989, by Hartman, D.W. Krause,
T. Kroeger, J. Hunter, G. Buckley, RE Lemelin, J.A.
Quintana, and K.A. Zarembev.
Reference Locality (L4232): Locality L4232 (Brown
Ranch locality) was discovered by the author July 28,
1982, and is one of the few continental molluscan locali¬
ties with abundant snails and/or bivalves that form a
shell-layer in the Ludlow Member of the Fort Union
Formation in North Dakota. Locality L4232 was found
to occur over a significant area of the School Section
drainage (L4232a-e) at the same horizon between the
Page 18
THE NAUTILUS, Vol. 129, No. 1
52T [10'. ......
45 [ Cypress
Hills
pr i.
[ EMT
|_(TR1s Ludlow of author) 0 50 100 km
Biogeography of Viviparus
codomorphus study
area localities
104' 031
Figure 93. Biogeography of Viviparus codomorphus localities in the northern Great Plains of the USA (see Figure 24 for map notes)
“Lower and Upper Coal Pair” and the Boyce and
Three V Tongues of the Cannonball Formation (Belt
et al., 2005) in the upper part of the Ludlow Member
(Slope Formation of others) on the Williams Lake Quad.
(1979), Slope County, North Dakota. Measured and
other specimens (Appendix 2) were collected at the
time of discovery and on July 9, 1986, by Hartman and
K.D. Enebo, and others; June 19, 1987, by Hartman,
S.G. Strait, R. Crane; and June 23, 1988, by Hartman,
D.W. Krause, T.C. Rae, and J.P. Hunter.
General Distribution: Viviparus codomorphus is pres¬
ently known from just a few localities. These occurrences
may be stratigraphically restricted to the upper part of the
Ludlow Member of the Fort Union Formation or may
range into the lowest part of the Tongue River Member
(Figure 1; Table 7). All of the occurrences are adjacent to
the Cedar Creek Anticline in far-western North Dakota
(Slope County, L4232) and easternmost Montana (Carter
and Fallon Counties, L0939 and L0049) to the Miles City
Arch (Wibaux County, L0936), Montana (see Hartman,
1984, cross section F-8, L4232a, b). This limited dis¬
tribution is mapped on Figure 93 (see Hartman, 1984,
table 42 for cadastral data). Although amenable to bulk
processing, no mammal teeth were found at L0049 after
considerable effort and picking. The Brown Ranch local
fauna includes mammal teeth from depauperate sites
between the Boyce and Three V Tongues of the Cannon¬
ball Formation. A Tol or To2 is presently the best age
estimate for this interval of the Ludlow Member (Hunter
J. H. Hartman, 2015
Page 19
and Hartman, 2004). The age of the lowermost part of
the Tongue River Member in Montana is uncertain, but
is unlikely to be younger than Ti2 NALMA (see Figure 1).
Discussion - Viviparm (Paludotrochus) codomorphus:
Viviparus codomorphus is the type species of the genus-
group taxon Paludotrochus Cossmann. It was introduced
to encompass the Late Cretaceous and Paleogene
viviparid taxa in the Western Interior (specifically, the
Laramie, s.L, of Montana). Cossmann (1921) identified
the type species as Viviparus meeki (then V. trochiformis) .
However, his description and illustrations clearly indi¬
cate that Paludotrochus was based on a taxon other than
V. meeki. Cossmann’s description and illustrated speci¬
men are assignable to V. codomorphus, thereby recog¬
nizing this taxon as the type-species of Paludotrochus.
Viviparus codomorphus is a relatively uncommon if
not rare taxon, but is locally abundant. V codomorphus
is presently known only from exposures near the Cedar
Creek Anticline in easternmost Montana and west¬
ernmost North Dakota (Little Missouri River valley)
(Figure 93). Specimens of this species are present in
some of the museum collections examined during the
course of this study (e.g., CSC general collection).
Most museum labels record the following information:
Montana (or Dawson County, Montana), Fort Union
Formation, Wards. I suggest as a possibility' that speci¬
mens of this species were acquired by Ward’s Natural
Science Establishment, Inc., Rochester, New York, and
were sold to various institutions and maybe to Cossmann.
Interestingly enough, the Ward’s specimens were most
often identified as V. trochiformis {— V. meeki).
DISCUSSION
The Laneian viviparid fauna (Figures 1, 67) is distinctive,
but was not recognized so by earlier paleontologists. In
the Williston Basin, the Laneian fauna is largely charac¬
terized by C. acroterion and V. thompsoni. The former
can occur by the thousands at some locations, while the
latter is always relatively few in numbers. Other species
of Viviparus are also present in the Laneian of Montana
and North Dakota and await further treatment. They,
too, are also relatively uncommon, with the North
Dakota specimens preserved as sideritic steinkerns
(Justham et ah, 2007).
The Puercan (early Paleocene) viviparid fauna is
based on relatively few localities, but additional records
are available (with limited temporal control) from studies
in the Bear Formation of the Crazy Mountains Basin,
Montana (Hartman, 1989). Bear Formation localities
are interpreted as lower Paleocene based on mamma¬
lian studies by Buckley (Hartman et ah, 1989; Buckley,
1995). The discovery of the Racetrack Ridge locality con¬
firmed the distinctiveness of the Viviparus purgatorius,
and provided an additional local fauna for comparison.
Other localities of importance include Purgatory Hill
(L0010) in Montana and Pine Cree Park (L0437) in
Cypress Hills, southwestern Saskatchewan, from the
basal strata of the Ravenscrag Formation. All of the
localities below the Pine Cree Locality are interpreted
to be early Paleocene. Locality L0220 occurs in the
lowermost strata of the Ravenscrag Formation and is
probably the oldest Paleocene locality of this report.
Few Torrejonian viviparid-bearing localities are known
from the Williston Basin (Hartman, 1984). The two best-
known localities (L0049 and L4232) provide very well
preserved specimens of Viviparus codomorphus and asso¬
ciated fossils, indicating their distinctiveness. Subsequent
work has found a large number of Torrejonian-age locali¬
ties in the Crazy Mountains Basin. Although most mate¬
rial is plastically deformed, viviparid diversity appears to
remain low although specimens are common. Possible
occurrences of Viviparus codomorphus in the lowermost
part of the Tongue River Member may indicate an upper
Torrejonian or lower Tiffanian age, depending on loca¬
tion. In Saskatchewan, in the valley of the Big Muddy,
the age of the Ravenscrag Formation may be at least
mid Paleocene (Ti2) (Sweet, 1978; Craig Scott, Geological
Survey of Canada, verbal communication, 2014).
Three new species viviparid species herald the spe¬
cies and evolution to follow in their respective genera.
All three species appear to have existed for about the
same duration (3 million years). Campeloma acroterion
is beginning part of a succession of three species of
Campeloma over about a 12 million-year period that
may be traced back to C. vetula (Meek and Hayden) of
the Campanian Judith River Formation of north-central
Montana. Viviparus purgatorius and V. codomorphus
may or may not give rise to the radiation of Viviparus
taxa seen in the northern Great Plains later in the
Paleocene. Viviparid taxa are a common if not domi¬
nant macrogastropod element of primarily fluvial faunas
during the changes in sea level effecting Laramidia
during the Late Cretaceous. The taxa have a similar, if
maybe reduced, overall disparity and diversity in early
and mid Paleocene faunas with multiple incursions of
the Cannonball Sea. With regression of the Cannonball
Sea in the early late Paleocene (Hartman et al., 1999),
viviparids radiate, becoming far more numerous, diverse,
and morphologically variable until the end of the
Laramide Orogeny (about mid Eocene). The species
introduced here, along with others that may exist, fore¬
shadow the success of Viviparus in the late Paleocene in
North America.
ACKNOWLEDGMENTS
This project was originally supported, in part, by the
University of Minnesota (Doctoral Dissertation Fellow¬
ship and Doctoral Dissertation Special Grant, Graduate
School; Department of Geology and Geophysics scholar¬
ships); Research Grant, Geological Society of America;
Grant-in-Aid of Research, Sigma Xi; research funding
from the New Mexico Bureau of Mines and Mineral
Resources; resources of the Minnesota Geological
Page 20
THE NAUTILUS, Vol. 129, No. 1
Survey, Smithsonian Institution Archives, U.S. National
Museum, and U.S. Geological Survey (Denver, Washington,
D.C.). Additional funding was provided by the NSF,
U.S. Bureau of Mines, and U.S. Department of Energy.
Generosity of time, expertise, and/or resources were
given by J.H. Hanley and N.F. Sohl (USGS); F.M.
Swain and R.E. Sloan (University of Minnesota); M.J.
Copeland and T.E. Bolton (Geological Survey of Canada),
F.D. Holland, Jr., and A.M. Cvancara (University of
North Dakota); C.G. Carlson (North Dakota Geological
Survey) (see also Hartman, 1984). More recent assis¬
tance and considerations were kindly given by A.E.
Bogan (North Carolina Museum of Natural Sciences),
W.A. Clemens (University of California- Berkeley), G.H.
Groenewold (Energy & Environmental Research
Center), J. Horner (Museum of the Rockies), D.W.
Krause (State University of New York-Stony Brook),
D.L. Lofgren (Raymond M. Alf Museum), and G.P.
Wilson (University7 of Washington). Acknowledgment
is gladly given to all the ranchers, who kindly provided
access to their property for study and fossil collection
and to those that continue to do so. Many field crews
were involved in collecting fossils and in support of
these activities; their camaraderie is gratefully noted
(see Appendix 1). R.D. Butler improved the manuscript
with his thoughtful edit.
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THE NAUTILUS 129(1 ):23-30, 2015
Page 23
Shell shape variation within a population of Astarte borealis
(Schumacher, 1817) (Bivalvia: Astartidae) from Camden Bay,
northern Alaska: a study using elliptical Fourier analysis
M.E. Chrpa
Department of Geology and Geophysics, MS 3115
Texas A&M University
College Station, TX 77843 USA
[email protected]
A.O. Oleinik
Department of Geosciences
Florida Atlantic University
777 Glades Road
Boca Raton, FL 33431 USA
[email protected]
ABSTRACT
Shells in the bivalve genus Astarte are known for variable
morphology and polymorphism within living and fossil species.
Astarte borealis (Schumacher, 1817), the most common living
species, is recognizable and common in the mid- to high-
latitude North Pacific, Arctic, and North Atlantic oceans, and
has been previously subdivided into several subspecies and
varieties based on variations in overall shell shape. A collec¬
tion of 641 recent specimens of A. borealis from Camden Bay,
northern Alaska, with intact outlines was analyzed for variabil¬
ity of the shell shape within a population of the species. The
analysis has important implications for morphological studies
of recent and fossil bivalve mollusks. Bivariate analysis of
length vs. height and morphometric analysis of shell outline
determined variants within a population of A. borealis, and
were compared to Pliocene A. borealis. The computer pro¬
gram SHAPE version 1.3 (Iwata and Ukai, 2002) uses elliptic
Fourier coefficients of shell outlines to evaluate and visualize
shape variations. The multivariate outline analysis indicates
that intraspecific shell variation in A. borealis is based upon a
modal shape that grades into other shapes, rather than grade
between two or more end-forms.
Additional Keywords: Multivariate outline analysis
INTRODUCTION
It has long been suggested that the outlines of objects,
mollusk shells in this case, are of great significance to
visual recognition and are therefore important for clas¬
sification purposes (Scott, 1980). The outline measure¬
ments, however has not been widely used in taxonomic
studies, in favor of more traditional distance measure¬
ments of height, length, and width, and relative geomet¬
ric locations of certain well-defined characters, known
as “landmarks”. Part of the problem was the lack of suit¬
able instrumentation for precise outline measurements.
Advances in computer technology and digital image
analysis have taken outline data collection and process¬
ing to a new level. Computer-based techniques of
shell shape quantification are particularly important
for fossil species, where other diagnostic characteristics,
such as those in soft tissues, are unavailable for study.
One of the common problems arising from attempts
to identify large number of similarly shaped individ¬
uals is the recognition of patterns of variability within
the population(s) that can be used as a basis for precise
taxonomic identification. As a test case for outline shape
analysis, we selected the high-latitude bivalve Astarte
borealis. Recent and Pliocene specimens from Alaska
were evaluated. Bivalves of the genus Astarte Sowerby,
1816 are also notorious for the conservatism in shell
shape that made them particularly suitable for this study
(Dali, 1920).
The bivalve genus Astarte is known from as early as
the Lower Jurassic in northern Siberia (Zakharov, 1970).
Recent species of Astarte are common in circumpolar
panarctic waters and are known for polymorphism
(Zettler, 2001). One of the most abundant recent spe¬
cies is Astarte borealis (Schumacher, 1817). The species
is known to have a high degree of shell shape variability
and has been often been called a “species complex”,
by various authors owing to numerous subspecies and
named varieties (Zettler, 2001, 2002; Ockelmann, 1958;
Petersen, 2001). Although there are several synonyms
attributed to this polymoiphic species, there have been
few detailed morphological studies done. Qualitatively,
the shell varies from ovate to subtrigonal and quadran¬
gular within tire genus. Among other characteristics, these
qualitative terms are often found in the descriptions of
Astarte species (Dali, 1903, 1920; Coan et ah, 2000). A
more precise definition of the shell shape is necessary in
an attempt to better quantify tire shape of the shell. One
of tire ways to examine the amount of variation and evalu¬
ate the possible presence of different shape morphs with
gradation, for both recent and fossil Astarte species, is to
use statistical techniques to study shell shape variability.
The objective of this study was to quantitatively deter¬
mine the degree of morphological variability in shell shape
Page 24
THE NAUTILUS, Vol. 129, No. 1
within a population of recent Astarte borealis, from a
single location in Arctic Alaska, based on a large sample
size. The hypothesis for this study is that a population
of extant A. borealis in Camden Bay, northern Alaska,
shows a high degree of variation in shell outline morphol¬
ogy. The variation is attributed to a central form grading
into a range of variants rather than the presence of two or
more distinct forms. An unbiased clustering around the
central form will indicate that any separation of morpho¬
logic forms that could potentially be defined within the
population is artificial. Existence of at least two distinct
shape “clusters” within the population will be suggestive
of quantifiable morphological variability in shell shape.
The secondary objective was to compare the results of
the outline analysis of modern A. borealis to Pliocene
A. borealis from Alaska. That was done to test how a
completely random small sample will compare with our
dataset. We had chosen a fossil, instead of recent, subset
to make samples both spatially and temporally different.
This is the first comprehensive study of the shell
outline variation within a single population of Astarte
borealis based on a large number of specimens. Although
A. borealis has been studied in comparison to other
Astarte species (Gardner and Thompson, 1999; Ockelmann,
1958; Saleuddin, 1965, 1967, 1974; Schaefer et al., 1985;
Selin, 2007; Skazina et ah, 2013; Zettler M.L., 2002),
comparative studies using the shell outline to determine
the degree and type of variability within A. borealis from
the same population have never been attempted. The
study is important in demonstrating the potential sig¬
nificance of a new technique for understanding the shell
shape variation in recent A. borealis and has significant
implications for systematic studies of both recent and
fossil species.
MATERIALS AND METHODS
More than 700 specimens of recent Astarte borealis
were collected in July of 2005, along a 5.5 km stretch
of gravel beach at Camden Bay, North Slope of Alaska
(Figure 1).
Twenty-three fossil specimens of Astarte borealis used
in this study were obtained on loan from the California
Academy of Sciences, Department of Invertebrate Zoology,
and Geology fossil collections. These specimens were
collected from the Pliocene marine facies of the Milky
River formation at Sandy Ridge section, Alaska Peninsula
(Marincovich et al., 2002) (Figure 1). Thirteen specimens
from this collection were used for the distance measure¬
ment analysis based on completeness with respect to the
distance measurements needed. Eleven specimens were
used for the outline analysis and were chosen based on
degree of outline completeness.
All recent shells were sorted to separate left from right
valves, then left valves were retained and sorted again
for elimination of specimens without complete outlines.
Valves with large chips or partial outlines due to break¬
age were removed from the study sample. A total of
641 recent left valves and 11 Pliocene specimens were
used for this study. Left valves of recent specimens were
chosen, with no preference over right valves, to eliminate
duplication of a specimen in the quantitative analyses.
All left valves were soaked in 10% sodium hypo¬
chlorite solution for approximately 48 hours to remove
periostracum, rinsed with de-ionized water, and dried.
Fossil specimens were unaltered for this study. Due
to limited number of specimens, both left and right
valves of fossil Astarte borealis were used when com¬
plete outline was visible.
Figure 1. Map showing location of collection sites in Alaska. 1. Camden Bay, North Slope, recent Astarte borealis. 2. Sandy Ridge,
Pliocene A. borealis.
M.E. Chrpa and A.O. Oleinik, 2015
Page 25
Figure 2. Outline of the shell of Astarte borealis showing
basic measurements. H: height; L: length; W: width.
Recent shells were prepared for photography by coat¬
ing the exterior surface with water-based tempera paint
to remove any image interference from rust staining on
the surfaces. The shells were photographed with digital
camera and processed in Adobe Photoshop 7.0 to retain
the outline of the image. The fossil shells had no pro¬
cessing prior to digital imaging, and images of any right
valves were reflected geometrically to represent left
valves. The last step was possible due to the equivalve
nature of A. borealis shell (Saleuddin, 1965).
The length to height (L/H) and width to height (W/H)
ratios were calculated for one third of recent specimens
that were used for outline analysis. These 225 valves were
randomly selected for the distance measurements, as well
as 13 fossil specimens. Regarding the measurements,
length (L) is the longest distance from front to back
edge; height (H) is the distance from the limbo to edge;
and width (W), is the longest distance of the valve in
a lateral plane across the valve (Figure 2). The linear
measurements were taken using a Mitutoyo Absolute
Digimatic digital caliper, and were made at a resolution
of millimeters to the hundredth place (0.01 mm).
The multivariate outline analysis was performed using
SHAPE ver. 1 .3, a software package for Quantitative Eval¬
uation of Biological Shapes Based on elliptic;)! Fourier
descriptors, developed by Iwata and Ukai (2002). The
variation in valve outline shape, a case of a closed contour,
is characterized bv the Elliptic Fourier descriptors (EFDs),
which are obtained by decomposing a curve into a sum of
harmonically related ellipses (Kuhl and Giardina, 1982).
First, the chain code was obtained from the processed
digital images, using the ChainCoder program (Iwata
and Ukai, 2002). The program converts the full color
image to black and white by splitting the image into
three colors with gray scale, converting the image with
clearest contrast to black and white. The noise is reduced
and the closed contour of the valve was extracted by edge
detection which is described as a chain code. Chain code
Length (mm)
Figure 3. H/L relationship for recent and Pliocene shells of Astarte borealis.
Page 26
THE NAUTILUS, Vol. 129, No. 1
is a coding system for describing geometrical information
about contours using numbers from 0 to 7, indicating
direction as measured counterclockwise from X axis of
X-Y coordinate system to represent the position of each
successive point in relation to the previous (Kuhl and
Giardina, 1982). The area of each valve was also recorded.
The normalized Elliptic Fourier descriptors (EFDs)
were calculated in Chc2Nef program of the SHAPE
ver. 1.3 package (Iwata and Ukai, 2002). The program
obtained chain code to calculate normalized EFDs fol¬
lowing the procedures suggested by Kuhl and Giardina
(1982). The EFDs are normalized to be invariant with
respect to size, rotation, and starting point and are based
on the first harmonic ellipse that corresponds to the con¬
tour information’s first Fourier approximation. The EFDs
are used to find the principal components of tire shape
variation. The principal component analysis of the normal¬
ized EFDs was accomplished in the PrinComp program
PCI
PC2
PC3
PC4
PCS
PC6
PC7
-2S.D.
Mean +2S.D.
Figure 4. The PrinPrint visualization of the mean shape and the +2 and —2 standard deviation from the mean in outlines for
die first 7 significant principal components. The first column shows the outlines of +2 and —2 standard deviation superimposed on
the mean form.
M.E. Chrpa and A.O. Oleinik, 2015
Page 27
of the SHAPE package (Iwata and Ukai, 2002) to effi¬
ciently summarize the information contained in these
coefficients for easier interpretation. The principal com¬
ponent analysis converts a set of observed correlated
variables and by means of orthogonal transformations,
produces a set of values that are linearly uncorrelated
variables or principal components (Rohlf and Archie,
1984). The first principal component produced has the
highest possible variance with respect to the data set. Each
successive principal component has the highest possible
variance with respect to the preceding component. Prin¬
cipal components were then visualized on a chart.
The visualization of principal component analysis was
done through the PrinPrint program. The program out¬
puts the shape variation accounted for by the largest
principal components. Following the procedure in Iwata
and Ukai (2002), the coefficients of the EFDs are calcu¬
lated such that the score for a particular principal com¬
ponent is equal to +2 or —2 times the standard deviation
from the mean, the square root of the eigenvalue of the
particular component and the scores of the remaining
components are zero. The coefficients are used to exe¬
cute an inverse Fourier transform and create contour
shapes that are visual representations of the data. That
visual output is helpful in interpreting the variation asso¬
ciated with each principal component.
RESULTS AND DISCUSSION
According to Selin (2007), shell proportions can be a
reliable parameter for differentiating between species.
The results of bivariate analysis for length to height (L/H)
and width to height (W/H) ratios are shown in Table 1.
The L/H ratio is 1.16 and is comparable to the pre¬
vious result of 1.15 obtained by Zettler (2001), also fall¬
ing between the ratio of 1.28 obtained by Oekelmann
(1958) and 1.10 by Selin (2007). The plot of height and
length measurements for both recent and fossil speci¬
mens is shown on Figure 3. The correlation coefficient
for height to length ratio is R2 = 0.7393 for the recent
(two-tailed P value is less than 0.0001) and R2 = 0.7363
for the fossil specimens (two-tailed P value equals 0.0027),
which suggest a very close similarity of shape between
recent and fossil specimens. The outline analysis shows
the areas where variability in shell outline was found as
well as the proportion of variance that could be attrib¬
uted to each component of variation among the popula¬
tion. The results of the statistical computation using
the SHAPE PrinComp package and the first seven princi¬
pal components are shown in Figure 4.
Principal components (PC) that represent variance in
outline were calculated from the symmetric and asym¬
metric aspects of the shell. The PrinComp package gives
results based numbers of harmonics used in the calcula¬
tion; for this analysis, 77 principal components were pro¬
duced. Here, only the first 10 principal components are
discussed, as coefficients with small variance and covari¬
ance values are generally not important for explaining
Table 2. Eigenvalues and contribution of principal compo¬
nents of Astarte borealis in this study.
the observed morphological variations. Table 2 has first
10 PC of the recent A. borealis outline analysis as com¬
puted by PrinComp. The 10 PCs account for about
94.94% of the variance found in the population, and the
first three PCs represent 83.21% of variance.
The PCs were plotted against one another to show the
concentration of variance in outline shapes (Figure 5).
The greatest variation is represented by the first PC, the
second greatest variation by the second PC and so forth.
The distribution of variation can be visualized by plot¬
ting the PCs against one another, since the first two PCs
comprise the most variation; they are plotted with the
PC f on the x-axis and the PC 2 on the y-axis. Figure 5
shows the first and second principal components plotted
with the outline contours drawn by the PrinPrint pro¬
gram, as well as the first and third principal components.
As shown on Figure 5, PC 1 accounts for 47.42% of the
variance found in the recent specimens and based on
tlie visualization this is representative of relative shell
height. From the positive to the negative standard devia¬
tions, the shell length only varies about 1% whereas the
shell height varies about 15%. The positive standard
deviation has a height to length ratio of 1 .09, the mean
shape ratio is 1.17 and the negative standard deviation
has a ratio of 1.26. The mean ratio of 1.17 is consistent
with the bivariate analysis results of length to height
ratios (Table 1). PC 2 accounts for 22.74 % of the vari¬
ance and represents the position of the umbones with
respect to the central line. The mean shape has an umbo
nearly on the midline, the positive deviation has the
umbo quite dorsal to the midline and the negative devi¬
ation has the umbo slightly dorsal of the midline. PC 3
accounts for 13.05 % of the variance and is representa¬
tive of the overall shell shape. The positive deviation has
a rounded, subquadrate figure ranging to the negative
valve with a compressed subtrigonal shape. This is con¬
sistent with the various descriptions available in litera¬
ture. No correlations occurred when PC 2 and PC 3
were plotted against one another.
The shell outline was analyzed for 1 1 fossil specimens
and results were plotted over the recent graphs for com¬
parison (Figure 6). Fossil specimens fall entirely within
the range of the recent specimens data. Since sample
size is different, 641 recent samples and 11 Pliocene
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THE NAUTILUS, Vol. 129, No. 1
0.1 -i
-0.06 -
-0.08 -
Principal Component 1
o o o
Figure 5. Plots of principle components. A. PC 1 vs. PC 2 of Astarte borealis with reconstructed contours. B. PC 1 vs. PC 3 of
A. borealis with reconstructed contours.
fossil samples, one-way Kruskal and Wallis ANOVA was
used to test the significant of differences (p < 0.0001)
between the recent and fossil A. borealis PC 1 results
(Kruskal and Wallis, 1952). This is interpreted as an
indication that the morphological variability of fossils is
similar to that of the recent specimens, however not a
direct indicator of correspondence.
Existing paleontological descriptions of fossil and
recent species of Astarte suggest great similarity in shape
among species. Use of elliptical Fourier analysis is an
attempt to show that the outline of fossil clams match
the modern and fall into the same distribution.
Morphologic variability of shell is one of the main cri¬
teria used for identification of fossil species of bivalve
M.E. Chrpa and A.O. Oleinik, 2015
Page 29
Principal Component 1
Figure 6. Principal Component 1 vs. Principal Component 2 lor recent and Pliocene A. borealis.
mollusks. It is, therefore, important to understand the limits
of morphologic variability within the same species and
population of bivalve mollusks, particularly the ones lack¬
ing prominent and diagnostic morphologic characteristics.
Outline shape analysis clearly indicates that Astarte
borealis has a high morphological variability at the spe¬
cies level and even within a single population, which can
be a potential source of confusion during species identi¬
fication, especially with the use of qualitative descriptive
parameters for shell shape. The bivariate comparison
analysis between Pliocene and recent populations shows
that there is continuity in the species and that the varia¬
tion has been a characteristic of the species for a long
time. The multivariate outline analysis indicates that the
intraspecific variation is based upon a common shape
that grades into other shapes evenly or a correlated popu¬
lation centered on a common “central” form. This study
does not support the idea that the variation within certain
species is a continuum between two end forms. For the
studied population of Astarte borealis , allocation to forms
and varieties would be at best problematic. Separation of
fossil species in the absence of a large specimens sample
should not be based solely on the shell outline and
should include other diagnostic characteristics, such as
external sculpture, details of the hinge, and features of
the shell margin.
The study of geographical distribution of species by
Zettler (2002) emphasized and confirms the polymor¬
phism of A. borealis. The polymorphism of the species
has been attributed to non-pelagic reproduction that
causes the eggs to attach to die substrate near the par¬
ents (Bernard, 1979; Ockelman, 1958). Since there is a
lack of diversity in reproduction due to limited genetic
mixing and a greater chance of isolation in the popu¬
lation, the variation could accelerate depending upon
environmental conditions such as substrate composition,
salinity, temperature, and nutrients. The slight differ¬
ences in environment may influence the changes in the
shell height to length ratio or perhaps the ventral margin
that affect a certain portion of the population but do not
separate it from the species. Changes can probably be
detected in shell thickness and overall size, which could
be a function of water temperature. That, in turn would
be a function of changing climate.
ACKNOWLEDGMENTS
The specimens were collected during a field trip
sponsored by the National Science Foundation grant
PLR 0425103.
LITERATURE CITED
Bernard, F. 1979. Bivalve Mollusks of the Western Beaufort
Sea. Natural History Museum Los Angeles County, Con¬
tributions in Science 313, 80 pp.
Coan, E.V., R Valentieh-Scott, and F. B. Bernard. 2000. Bivalve
Seashells of Western North America: Marine Bivalve Mol¬
lusks from Arctic Alaska to Baja California. Santa Barbara
Museum of Natural History Monographs Number 2, Studies
in Biodiversity Number 2, 764 pp.
Dali, W.H. 1903. Synopsis of the family Astartidae, with a
review of the American species. Proceedings of the U.S.
National Museum 26 (1342): 933-951.
Dali, W.H. 1920. Pliocene and Pleistocene fossils from the
Arctic coast of Alaska and the auriferous beaches of Nome,
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Norton Sound, Alaska. U.S. Geological Survey Professional
Paper 125C: 23-37.
Gardner, J. and R. Thompson. 1999. High levels ol shared
allozyme polymorphism among strongly differentiated con¬
generic clams of the genus Astarte (Bivalvia: Mollusca).
Heredity 82: 89-99.
Iwata, H. and Y. Ukai. 2002. SHAPE: A computer program
package for quantitative evaluation ol biological shapes
based on elliptic Fourier descriptors. Journal of Heredity
93: 384-385.
Kruskal, W.H. and W.A. Wallis. 1952. Use of ranks in one-
criterion variance analysis. Journal of the American Statis¬
tical Association 47(260): 583-621.
Kuhl, F. and C. Giardina. 1982. Elliptic Fourier features ol a
closed contour. Computer Graphics and Image Processing
18: 236-258.
Marincovich, L., K.B. Barinov, and A.E. Oleinik. 2002. The
Astarte (Bivalvia: Astartidae) that document the earliest
opening of Bering Strait. Journal ol Paleontology 76:
239-245.
Ockehnann, W.K. 1958. The zoology of East Greenland: Marine
Lamellibranchiata. Meddelelser om Gr0nland 122 (4):
1-256.
Petersen, G. H. 2001. Studies on some Arctic and Baltic Astarte
species (Bivalvia, Mollusca). Meddelelser om Grpnland,
Bioscience, 71 pp.
Rohlf, F.J. and J.W. Archie. 1984. A comparison of Fourier
methods for the description of wing shape in mosquitos
(Diptera: Culicidae). Systematic Zoology 33: 302-317.
Saleuddin, A.S.M. 1965. The mode of life and functional
anatomy of Astarte spp. (Eulamellibranchia). Proceed¬
ings of the Malacological Society of London 36: 229-257.
Saleuddin, A.S.M. 1967. Notes on the functional anatomy of
three North American species of Astarte, A. undata Gould,
A. g astanea Say and A. esquimalti Baird. Proceedings of
the Malacological Society of London 37: 381-384.
Saleuddin, A.S.M. 1974. An electron microscopic study ol the
formation and structure ol the periostraeum in Astarte
(Bivalvia). Canadian Journal of Zoology 52: 1463-1471.
Schaefer, R., K. Trutsclder, and H. Rumohr. 1985. Biometric
studies on the bivalves Astarte elliptica, A. borealis and
A. montagui in Kiel Bay (Western Baltic Sea). Helgoland
Marine Research 39: 245-253.
Scott, G. 1980. The value of outline processing in the biometry
and systematics of fossils. Palaeontology 23: 757-768.
Selin, N.I. 2007. Shell form, growth and life span ol Astarte
arctica and A. borealis (Mollusca: Bivalvia) from the sub-
tidal zone of northeastern Sakhalin. Russian Journal of
Marine Biology 33: 232-237.
Skazina, M., FI. Sofronova, and V. Khaitov. 2013. Paving the
way for the new generations: Astarte borealis population
dynamics in the White Sea. Hydrobiologia 706: 35^49.
Zakharov, V.A. 1970. Late Jurassic and Early Cretaceous
Bivalves of the North of Siberia and their ecology. Family
Astartidae. Transactions ol the Institute of Geology and
Geophysics SB Academy ol Sciences USSR, Nauka, Moscow,
143 pp. [in Russian]
Zettler, M. 2001. Recent geographical distribution of the Astarte
borealis species complex, its nomenclature and bibliography
(Bivalvia: Astartidae). Schriften zur Malakozoologie 18: 1- 14.
Zettler, M. 2002. Ecological and moqdiological features ol the
bivalve Astarte borealis (Schumacher, 1817) in the Baltic
Sea near its geographical range. Journal of Shellfish
Research 21: 33-40.
THE NAUTILUS 129(l):31-42, 2015
Page 31
Reviving a cold case: two northeastern Pacific dendrodorid
nudibranchs reassessed (Gastropoda: Opisthobranchia)
Jeffrey H.R. Goddard
Marine Science Institute
University of California
Santa Barbara, CA 93106-6150 USA
[email protected]
Angel Valdes
Department of Biological Sciences
California State Polytechnic University
3801 West Temple Avenue
Pomona, CA 91768-4032 USA
[email protected]
ABSTRACT
Ten nominal species of dendrodoridid nudibranchs are known
from the NE Pacific Ocean, including the Gulf of California.
However, disagreement surrounds the taxonomic status of
Doriopsilla nigromaculata (Cockerell in Cockerell and Eliot,
1905) and Doriopsilla rowena Marcus and Marcus, 1967, includ¬
ing the correct generic placement of the former. To resolve
this disagreement, we examined type specimens, the original
descriptions, and unpublished materials in the James Lance
Collection at the California Academy of Sciences and conclude
that Doriopsilla nigromaculata is: (1) a member of Dendrodoris;
(2) not synonymous widi the valid species Doriopsilla rowena-,
and (3) a senior synonym of Dendrodoris behrensi Millen and
Bertsch, 2005. Like other members of the genus, Dendrodoris
nigromaculata has a centered anus, smooth dorsum, delicate
wavy mande edge, and possesses both ptyaline and esophageal
glands. It is translucent white widi chocolate brown blotches, the
larger of which are usually clustered into three or four groups
centered mid-dorsally. In contrast, Doriopsilla rowena has an
off-center anus, a papillate and densely spiculate dorsum with a
stiff margin, and lacks both ptyaline and esophageal glands. Its
scattered brown decks and larger, round concentrations of
opaque white distinguish it dorsaOy, and notal spicules include
rods and forks. Doriopsilla rmuena grows to 12 mm long and has
been found in La Jolla, California, the Pacific coast of Baja
California, and the northern Gulf of California soudi to Panama.
Dendrodoris nigromaculata grows to 27 mm and is known from
Monterey, California soudi to the San Benitos Islands, Baja
California. Bodi species have large eggs and ametamoqihic
direct development, but small eggs indicating planktotropbic
development have also been observed in D. rcnvena from Jalisco,
Mexico, suggesting D. rowena may constitute a cryptic species
complex or display poecilogony.
Additional Keywords: Dendrodorididae, Nudibranchia,
nomenclature
INTRODUCTION
Ten nominal species of dendrodoridid nudibranchs are
currently recognized from the NE Pacific Ocean, includ¬
ing the Gulf of California (Behrens and Hermosillo,
2005): Dendrodoris azineae Behrens and Valdes, 2004;
Dendrodoris behrensi Millen and Bertsch, 2005; Dendrodoris
fumata (Riippell and Leuchart, 1831); Dendrodoris stohleri
Millen and Bertsch, 2005; DoriojtsiUa albopunctata (Cooper,
1863); Doriopsilla gemela Gosliner, Schaefer and Millen,
1999; Doriopsilla janaina Marcus and Marcus, 1967;
Doriopsilla nigromaculata (Cockerell in Cockerell and
Eliot, 1905); Doriopsilla rowena Marcus and Marcus,
1967; and Doriopsilla spaldingi Valdes and Behrens,
1998. However, an important taxonomic disagreement
remains in the modern literature. C amach o- Garcia et al.
(2005) considered Doriopsilla rowena a valid species,
whereas Behrens and Hermosillo (2005) regarded it as a
junior synonym of Doriopsilla nigromaculata. The sug¬
gested synonymy of D. nigromaadata and D. rowena was
raised as a possibility by James Lance in Keen (1971: 830;
cited as Doriopsilla nigromaadata ) and again by McDonald
(1983: 171; cited as Dendrodoris nigromaadata). How¬
ever, if Camaeho-Gareia et al. (2005) were correct, and
Doriopsilla rowena is valid, then D. nigromaculata, which
was originally described based on a single specimen col¬
lected by Cockerell in La Jolla, California, has either
remained unknown since its original description and
should be regarded as a nomen dubium, or has since been
described under another name.
A contributing factor to this disagreement is that
historically the distinctions between Dendrodoris and
Doriopsilla have been confusing, including for species
from the northeast Pacific Ocean (reviewed by Steinberg
1961; Valdes and Ortea 1997; Gosliner et al. 1999). How¬
ever, Valdes et al. (1996) and Valdes and Ortea (1997)
provided clear anatomical and morphological criteria
separating the two genera, and recent phylogenetic anal¬
yses support this separation (Valdes and Gosliner 1999,
Valdes 2003). Here, we use these criteria and the exam¬
ination of type specimens to first establish the correct
generic placement of D. rowena and D. nigromaculata,
ruling out the synonymy of these two species. Then we
compare their original descriptions with those of other
species of dendrodoridids known from the region to
Page 32
THE NAUTILUS, Vol. 129, No. 1
show that (1) Doriopsilla rowena is valid, and (2) that
one species described recently from the region is in fact
a junior synonym of Dendrodoris nigromaculata. In both
steps we also draw from extensive materials in the James
R. Lance Collection at the California Academy of Sci¬
ences in San Francisco (hereafter, Lance Collection).
These constitute new evidence not available prior to
Lance’s death in 2006 and shed light on what had effec¬
tively become a taxonomic cold case.
MATERIALS AND METHODS
To establish the generic placement of Doriopsilla
nigromaculata and Doriopsilla rowena, we extracted
from their original descriptions information on four mor¬
phological and two anatomical characters which taken
together can be used to separate Dendrodoris from
Doriopsilla (Table 1). We corroborated this information
by examining the type specimens of both species, which
were obtained, respectively, from The Natural History
Museum, London (NHMUK) and the U.S. National
M useum of Natural History (USMNH). We also exam¬
ined Lance’s specimens of D. rowena in the Invertebrate
Zoology and Geology collection at the California Acad¬
emy of Sciences (CASIZ) and used his description of D.
rowena in Keen (1971), as well as his unpublished notes,
35mm photographic slides, and illustrations in the Lance
Collection. Lance kept many of these materials orga¬
nized by species in folders, which contain information
on the morphology, color, anatomy, egg masses, and
development of living specimens of most of the
dendrodoridids known from the northeast Pacific Ocean.
The folder for D. rowena in particular supplements
Marcus and Marcus’s original (1967) description of
D. rowena, which was based on preserved material.
Where appropriate, we refer to species folders in the
Lance Collection by their names; field accounts by num¬
ber, date, and locality; and 35mm slides by date and
locality when possible. To compare D. nigromaculata
with other species of Dendrodoris from the northeast
Pacific Ocean, we used information from our examina¬
tion of the type specimen, as well as from the Lance
Collection, Behrens and Hermosillo (2005), Millen and
Bertsch (2005), and Goddard (2005). Finally, we used
the online searchable database of the California Acad¬
emy of Sciences Invertebrate Zoology collection (http://
research.calacademy.org/redirect?url = http://research
archive.calacademy.org/research/izg/iz_coll_db/index.asp)
to obtain additional and otherwise unpublished locality
information to further document the known geographic
distribution of these species. Here, we used information
for specimens identified by established authorities famil¬
iar with the nudibranch fauna of the region. These are
referenced below by CASIZ and the corresponding cata¬
log numbers.
RESULTS
Description oe Relevant Materials in the Lance
Collection
1. Species Folder: Doriopsilla rowena contains:
(1) A hand-written description, accompanied by pencil
sketches, of this species with the manuscript name
“Doriopsilla puertecitensis” (later, Lance wrote the name
D. rowena in red ink on this description). The pencil
sketches are based on living specimens (CASIZ 182606)
collected 20 March 1965 from 4.4 mi south of Puertecito,
Baja California, and include dorsal and ventral views of an
adult, details of the notum and notal spicules, and penis.
The description also includes a sketch and brief descrip¬
tion of an egg mass laid by a 12 mm adult. (2) Six separate
pen and ink illustrations, all labeled as Doriopsilla rowena
and clearly based on the above pencil sketches. (3) A
sheet labeled ‘'Doriopsilla rowena - Field Account Data”
for sites in the Gulf of California. The dates listed include
years from 1954 to 1979, and out of 10 sites listed record
D. rowena only from Puerto Penasco. (4) One clear plastic
sheet holding 35 mm slides from 1966 and 1969, the
earliest labeled first as “Dendrodoris sp.” and later as
“Doriopsilla rowena or D. nigromacidata?”
2. Species Folder: Doriopsilla nigromaculata con¬
tains: (1) A list of specimens found at South Casa Reef
and Windansea Reef, both in La Jolla, California. (2) A
lined sheet with dorsal and ventral sketches of an adult
from La Jolla. The notation “F.A. 181”on this sheet refers
to Field Account 181, which was for 25 June 1967 at
South Casa Reef, La Jolla. This sheet is first labelled as
“Brown spotted Doriopsilla,” with “? D. nigromaculata”
later added in red ink. (3) A note stating “D. nigromaculata
Published sketch in Opis. News. 14(8): 29 of hatching.”
(4) A three page typed description of this species with the
Table 1. Select diagnostic characters distinguishing Dendrodoris from Doriopsilla. Based on Valdes et al. (1996) and Valdes and
Ortea (1997).
J.H.R. Goddard and A. Valdes, 2015
Page 33
Figures 1-6. Type specimens. 1. Dendrodoris nigromaculata (NHMUK 1904.7.7.1). Photo by Harry Taylor. 2-6. Doriopsilla
roxvena (USMNH 678413). Photos by Chris Meyer.
heading “Doriopsilla nigromaculata (Cockerell and Eliot,
1905) Figs. 1-7.” The figures for this description and
legend are on a separate sheet and are a composite of
copies of the six pen and ink illustrations from the Species
Folder lor Doriopsilla rowena, plus a map of California
with a single red ink dot showing “Distribution in California.”
The description is based on the four specimens Lance
found in La Jolla on 25 June 1967 but refers to the speci¬
men from La Jolla described by Cockerell and Eliot
70 years earlier. (5) Two sheets with sketches and notes
on egg masses laid by specimens collected at Windansea
Reel in 1968, San Quintin in 2001, and South Casa Reef
in 1998. (6) Two sheets containing 35 mm photographic
slides, one with images of adult specimens, the other with
images of egg masses and hatching juveniles.
3. Species Folder: White Porostome Spotted con¬
tains: (1) A sheet titled “Crenulate dorid” with sketches
in pencil of two specimens and a brief description,
including dimensions of 24 x 5 mm and 27 x 5 mm.
This sheet is undated but the dimensions of the larger
specimen, combined with information in (2) below indi¬
cate that these specimens were collected in either 1961
from the Coronados Islands or in 1963 from Point Loma,
San Diego. (2) A hand-written description of this species
with the manuscript name “Dendrodoris barbarensis,”
based on a single specimen collected from 8 m depth at
Naples Reef, Santa Barbara County 30 Oct 1966, two
specimens collected at 30 m depth 1.6 km south of South
Coronado Island by Nan Limbaugh on 22 Apr 1961, and
two specimens collected intertidally at Point Loma by
Wesley Farmer on 29 Oct 1963. (3) A typed sheet with
information on three specimens collected intertidally at
Lunada Bay, Palos Verdes Peninsula, by William Jaeekle,
20 Apr 1983. A note indicates that one of these speci¬
mens laid an egg mass on 4 May 1983. (4) Pencil notes
and sketches of an egg mass laid by an individual col¬
lected from South Casa Reef, La Jolla, 20 July 1974, and
sketches of the subsequent direct development to hatch¬
ing juveniles from that egg mass.
4. Field Account 181 (June 25, 1967, South Casa Reef,
La Jolla) contains, in addition to the annotated list of opis-
thobranchs found on that date by Lance (and Barbara
Good), contains notes, from three separate dates, on die
development of embryos in an egg mass deposited by
Doriopsilla rowena (cited as “doriopsillids,” with a later nota¬
tion as “Doriopsilla nigromaculata ?”) collected on that date.
Page 34
THE NAUTILUS, Vol. 129, No. 1
Figures 7-10. Living adult Doriopsilla rowena. 7. Percebu, Gulf of California, Baja California, 7 April 1966 (35 mm slide in Lance
Collection, Species Folder: Doriopsilla rowena). 8. Total length 9.2 mm, San Quintin, Baja California, 16 November 2001 (35 mm
slide in Lance Collection, Species Folder: Doriopsilla nigromaculata) . 9. South Casa Reef, La Jolla, California, 25 June 1967 (35 mm
slide in Lance Collection, Folder: Doriopsilla nigromaculata). 10. Total length (in MgCl2) 6.7 mm, Lindo Mar, Bahia de Banderas,
Jalisco, Mexico, 26 Feb. 2006 (CASIZ 174055; Goddard and Hermosillo, 2008, as Doriopsilla nigromaculata).
Genus-Level Taxonomy
Doriopsilla nigromaculata. In the original descrip¬
tion of D. nigromaculata Cockerell and Eliot (1905)
described the mantle of a single preserved specimen as
“smooth, not tubereulate,” with a “rather narrow” mar¬
gin. The latter is consistent with the delicate margin char¬
acteristic of living specimens of Dendrodoris (Table 1) and
is visible in the type specimen (NHMUK 1904.7.7.1) as
upturned, crenulate and soft in appearance (Figure 1).
Cockerell and Eliot (1905) mentioned “a number of glis¬
tening white spicules. . .imbedded in the skin” of the man¬
tle and described them as “mostly fairly straight long rods,
but some of the smaller ones are bent and have an irreg¬
ular outline.” The spicules are not described as being
regularly or densely arrayed and are visible in the type
specimen where the mantle tissue had been carefully
scraped away, presumably by Eliot in his original exami¬
nation of the specimen (Figure 1). The position of the
anus can be inferred as centered based on their descrip¬
tion of the gill plume as “set in a semicircle open
behind.” Internally, they described a “follieulate” mouth
gland “with a fairly long duct,” which is clearly a ptyaline
gland (e.g., illustrations in Valdes et ah, 1996; Millen and
Bertsch, 2005). No esophageal glands were mentioned,
but these are minute (Valdes et al. 1996; Millen and
Bertsch, 2005, Figure 2) and likely would not have gar¬
nered attention. All of these traits, as well as the extreme
anterior and posterior position of the rhinophores and
gills, respectively, in the type specimen, are consistent with
contemporary descriptions of species of Dendrodoris and
clearly indicate the correct generic of D. nigromaculata is
Dendrodoris (Table 1).
Doriopsilla rowena. Marcus and Marcus (1967)
described D. rowena based on seven preserved syntypes,
five of which remain (USMNH 678413), and notes by
the collector on the color of the living animals, which
were collected in Puerto Penasco, Sonora, Mexico, in
the northern Gulf of California (Figures 2-6). They
described the mantle as “smooth,” with evidence of
large, but dissolved, diagonally crossed spicules in the
“connective tissue of the back." They noted that “the
thick anal papilla and the renal pore in front of it lie to
the left of the branchial tuft,” clearly indicating the
eccentric position of the anus. The anterior part of the
alimentary tract is illustrated in their figure 62B, and
does not show either ptyaline or esophageal glands. Else¬
where, Marcus and Marcus (1967: 99) state that the
absence of a ptyaline gland is one characteristic of
Doriopsilla. Marcus and Marcus (1967) did not describe
any dorsal tubercles in D. roivena. Plowever, James Lance,
who found and observed living specimens of this species
from Puerteeitos, Baja California, and Puerto Penasco,
both in the northern Gulf of California (Figures 1 1—12),
illustrated minute, spiculate dorsal tubercles, which was
confirmed with examination of Lances original specimens
(CASIZ 182606). Lance also illustrated the pattern and
J.H.R. Goddard and A. Valdes, 2015
Page 35
11
Figures 11-13. Doriopsilla roivena, details of notum; all
Irom Lance Collection, Species Folder: Doriopsilla roivena.
11. 1x1 mm detail of notal surface. 12. Notal tubercle with
spicule detail. 13. Notal spicule pattern.
shapes of the notal spicules (Figure 13), which consist of
diagonally crossed rods and forks (Lance Collection, Spe¬
cies Folder: Doriopsilla roivena). Taken together, these
traits indicate that D. roivena is indeed a Doriopsilla,
as originally described by Marcus and Marcus (1967)
(Table 1). This was further confirmed with the examination
of the syn types (Figures 2-6), which are relatively flat and
wide animals covered with small spiculate tubercles and an
eccentric anus. All these traits are consistent widi the diag¬
nosis of Doriopsilla by Valdes and Gosliner (1999).
Species-Level Taxonomy
Doriopsilla roivena. Based on the above morpho¬
logical and anatomical differences, Doriopsilla roivena
cannot be a synonym of Dendrodoris nigromaculata.
Doriopsilla roivena differs externally from all other con¬
geners known from the region in its small adult size
(up to 12 mm total length) and unique color pattern,
consisting of small reddish-brown flecks scattered over the
dorsum, as well as larger, round concentrations of opaque
white, frequently arranged in longitudinal series, against a
ground color of off white to pale yellow, pink or orange
(Marcus and Marcus, 1967; Lance in Keen 1971; Behrens
and Ilermosillo, 2005, cited as Doriopsilla nigromaculata ;
Camaeho-Garefa et al., 2005) (Figures 2-10). Doriopsilla
roivena is therefore a valid species.
Dendrodoris nigromaculata. Cockerell and Eliot
(1905) based their original description of D. mgromaculata
on a single specimen, 10 mm long preserved, that was
collected in July 1901 from La Jolla, California (Figure 1).
The color of die preserved specimen was described as
“yellowish-drab with a slight inclination to lilac in places,”
with “a double border of black spots round the dorsal
margin, and a few larger black blotches symmetiieally dis¬
posed, one in front of the rhinophores, two behind them,
two in die middle of die back, and five in front of the
branchiae.” These larger blotches are still visible in the
type specimen (Figure 1). Two nominal species of
Dendrodoris from die northeast Pacific Ocean have dark
spots against a pale background: D. behrensi Milieu and
Bertsch, 2005 (Figure 17) and D. .stohleri Millen and
Bertsch, 2005. In D. stohleri die black spots are relatively
uniform in size and scattered over the entire dorsum, save
its margin (Millen and Bertsch, 2005). However, compari¬
son of the color patterns originally described for D.
nigronuicidata and D. behrensi reveals a virtually identical
match (Table 2). Indeed, the only discrepancy between the
descriptions of these two species is that Millen and Bertsch
(2005: 195) state that D. behrensi lacks mantle spicules.
However, in the companion paper to Millen and Bertsch
(2005), Goddard (2005) illustrated and described “slightly
curved, spindle-shaped spicules” arranged in a lattice
in D. behrensi recently hatched (Figure 21) from egg
masses hud by adults (Figure 17) included as paratypes
of D. behrensi by Millen and Bertsch (2005). Lance was
also familiar with this species and observed egg masses
and hatching juveniles virtually identical to those
described by Goddard (2005) (Lance Collection, Spe¬
cies Folder: White Porostome Spotted) (Figures 18-20).
Moreover, Lance described the adult body of this
species as “very slightly spiculose with notal margins
non spiculose” (Lance Collection, Species Folder: White
Porostome Spotted). The density of notal spicules there¬
fore decreases as juveniles grow into adults, which
could make the spicules easily overlooked in living adult
specimens. A similar phenomenon was recently docu¬
mented by Sanehez-Tocino et al. (2014) for some chromo-
dorid nudibranchs.
Page 36
THE NAUTILUS, Vol. 129, No. 1
Table 2. Comparison of original descriptions of external color of a single preserved Dendrodoris nigromaculata and living
Dendrodoris behrensi by Cockerell and Eliot (1905) and Millen and Bertsch (2005), respectively.
With no significant differences between their original
descriptions, Dendrodoris behrensi is therefore a junior
synonym of Dendrodoris nigromaculata .
SYSTEM ATICS
Based on the taxonomic results described above, a new
systematic arrangement and list of synonyms is proposed
for D. rowena and D. nigromaculata.
Family Dendrodorididae O’Donoghue, 1924
Genus Doriopsilla Bergh, 1880
Doriopsilla rowena Marcus and Marcus, 1967
(Figures 2-16)
Doriopsilla rowena Marcus and Marcus, 1967: 205-
207; Keen, 1971: 830; Poorman and Poorman, 1978: 373;
Bertsch and Kersbtch, 1984: 267; Valdes and Ortea, 1997:
253; Gosliner et al., 1999: 209; Valdes and Gosliner, 1999:
338-340; Camacho-Garcia et al., 2005: 80; Goddard and
Hermosillo, 2008: 87; Angulo-Campillo, 2005: Table 2.
Dendrodoris (?) nigromaculata [non Cockerell in
Cockerell and Eliot, 1905] - Behrens, 1980: 58.
Dendrodoris nigromaculata [non Cockerell in
Cockerell and Eliot, 1905], — Steinberg 1961: 59; Sphon
1972: 61; McDonald and Nybakken, 1980: 52; Lance,
1982: 29; McDonald, 1983: 170-171; Behrens, 1991: 71;
Angulo-Campillo, 2003: Table 2; Goddard, 2004: 1959,
1963; Goddard, 2005: 206.
Doriopsilla nigromaculata [non Cockerell in Cockerell
and Eliot, 1905], — Behrens and Hermosillo, 2005:
88; Lance Collection, Species Folder: Doriopsilla
nigromaculata. California Academy of Sciences and
Goddard, 2013: worksheets for Ladera St., False Point,
Windansea, and So. Casa Reef (data from unpublished
field accounts, no page numbers).
Type Material: Doriopsilla roivena - Syn types: 5
specimens, Puerto Penasco, Sonora, Mexico (USMNH
678413).
Anatomy: The anatomy of D. rowena was described
by Marcus and Marcus (1967), with additional details
presented by Valdes and Gosliner (1999). Further, as
noted and illustrated by Lance, the notum is densely
spieulate with rods and forks (Figures 12-13, 16).
External Morphology: The external morphology of
D. roivena was described by Marcus and Marcus (1967),
with additional details presented by Lance (Lance
Collection, Species Folders: Doriopsilla rowena and
Doriopsilla nigromaculata ) and Lance in Keen (1971).
Living adults reach 12 mm in length. The dorsum is
flecked with dark brown, has larger, round concentra¬
tions of opaque white, typically in longitudinal series,
and is covered by minute, spieulate tubercles (Figures 7-
12). The ground color varies from whitish to yellow to
pinkish orange.
Development: Lance obtained egg masses and
observed development of Doriopsilla rowena from
La Jolla (Figures 14-16). The egg ribbons were tightly
coiled and laid flat, rather than on edge like most other
dorid nudibranchs, and contained large eggs, encap¬
sulated singly, that developed into hatching juveniles
(Figure 15) in more than 26, but less than 38 days, at an
ambient temperature of approximately 20° C (Lance
Collection, Species Folder: Doriopsilla nigromaculata-.
Field Account 181, South Casa Reef, 25 June 1967).
Recently hatched juveniles possess the same notal spic¬
ule complement of rods and forks as adults (Figures 13
and 16). Lance sketched another egg mass, laid by a
specimen 9.5 mm long collected from Windansea Reef
in June 1968. This egg mass was also laid flat and had 4
whorls in a closed spiral and a total diameter of 5 mm, as
indicated by a scale bar. Based on that scale bar, the eggs
illustrated by Lance measured approximately 215 pm in
diameter. A note on Lance Field Account 181 indicates
that the embryos did not develop a shell. Taken together,
this information indicates that D. rowena from La Jolla
have ametamorphic direct development (Bonar 1978;
Goddard 2004). Lances sketch and notes on an egg mass
laid by a D. roivena from San Quintin, Baja California
are also consistent with this mode of development
J.H.R. Goddard and A. Valdes, 2015
Page 37
Figures 14-16. Doriopsilla rowena, egg mass and hatching juveniles; all from 35 mm slides in Lance Collection, Species Folder:
Doriopsilla nigromaculata. 14. Egg mass from South Casa Reef, 26 Apr. 1998. 15. Hatching juveniles, no date or locality (35 mm
slide processed Sep. 1974). 16. Spicule detail, hatched juvenile. South Casa Reef, no date (35 mm slide processed Sep. 1974).
(Lance Collection, Species Folder: Doriopsilla
nigromaculata) .
A specimen of D. rowena (Figure 10; CASIZ 174055),
6.7 mm long, collected from Nayarit, Mexico laid a flat
egg ribbon of 6 turns in a closed spiral 6 mm across,
virtually identical to the egg masses described above
from La Jolla (JG, unpublished data). However, the eggs
from Nayarit averaged 97.4 pm in diameter, indicating
planktotrophic development (Goddard and Hermosillo,
2008). Lance described similar egg masses laid by speci¬
mens of D. roivena from the northern Gulf of California
but mode of development cannot be inferred based on
any of the information included (Lance Collection, Spe¬
cies Folder: Doriopsilla rowena).
Geographic Distribution: Doriopsilla roivena is
known from the northern Gulf of California to Panama,
(Marcus and Marcus 1967; Keen 1971; Poorman and
Poorman 1978; Bertsch and Kerstitch 1984; Camacho-
Gareia et al. 2005; Goddard and Hermosillo 2008;
Angulo-Campillo 2005; CASIZ 171209, 171229), the
Galapagos Islands (CASIZ 78390, 78408), and La Jolla,
California to El Campo, near Punta Eugenia, Baja
California Sur (Lance Collection, Species Folder:
Doriopsilla nigromaculata; CASIZ 71519; Behrens anti
Hermosillo 2005) (Figure 22).
Remarks: As evidenced in the Lance Collection (Spe¬
cies Folder: Doriopsilla roivena), Jim Lance started to
Page 38
THE NAUTILUS, Vol. 129, No. 1
describe D. rowena under the manuscript name “D.
puerteeitensis,” based on three specimens he collected at
Puertecitos, on the Baja California side of the northern
Gulf of California. Sometime following the publication of
Marcus and Marcus (1967), Lance added in red ink the
name D. rowena on his original pencil sketches of these
specimens. Later he executed a set of undated pen and
ink illustrations, some of which are reproduced here in
Figures 1 1-13 and labeled them all as Doriopsilla rowena
(these illustrations were likely made for the monograph
on Panamic opisthobranehs that Lance intended but
never completed). In the drawing of the dorsal view of
an adult (not reproduced here) Lance depicted slx
thin, widely spread, bipinnate gills and a centered anus.
However, examination of his original specimens from
Puertecitos (CASIZ 182606) reveal more tightly clustered
gills and an eccentric anus, indicating that Lances draw¬
ing in these respects was purely schematic.
On 25 June 1967 at South Casa Reef in La Jolla, Lance
found four specimens of a small (up to 9 mm long)
“doriopsillid” that he first referred to in his notes and
illustrations as the “Brown-spotted Doriopsilla ” but later
changed in red ink to “?D. nigromaculata” (Lance Collec¬
tion, Field Account 181 and Species Folder: Doriopsilla
nigromaculata). He eventually found this species at three
more sites on the outer coast of San Diego County, as well
as in Bahia San Quintin, Baja California, and in his field
accounts for these sites consistently used the name
Doriopsilla nigromaculata for this species (Lance Collec¬
tion; California Academy of Sciences and Goddard 2013).
Lance’s Species Folder for Doriopsilla nigromaculata
includes a typed description, titled Doriopsilla
nigromaculata (Cockerell and Eliot, 1905), which was
based on the four specimens from La Jolla, California in
June 1967. This description is accompanied by and ref¬
erences a composite figure, also labeled as Doriopsilla
nigromaculata , but which was comprised solely of his
earlier illustrations (eventually labelled as D. rowena) of
“D. puerteeitensis” from the Gulf of California (some of
which are reproduced here in Figures 11-13). In the text
of this description Lance states “it is likely that the pres¬
ent species and an unnamed one, frequently encoun¬
tered in the Gulf of California, will eventually prove to
be nonspecific.” This indicates that Lance wrote this
description after June 1967, but prior to receiving a copy
of Marcus and Marcus (1967), and also suggests that
after receiving the latter, he considered D. rowena a
junior synonym of D. nigromaculata. Further, as men¬
tioned above, hatching juveniles from La Jolla possess
the same notal spicule complement of rods and forks
as adults from the Gulf of California (Figures 13 and
16). Although morphologically the specimens from La
Jolla are very similar to D. rowena from the Gulf of
California, Lance never applied the name D. nigromaculata
to the latter, and in conversation with at least one colleague,
maintained that the two were probably distinct (T.M.
Gosliner, personal communication to JG, 12 Dec 2014).
Lance (1982) illustrated a hatching juvenile, labeled as
“ Dendrodoris nigromaculata (Cockerell in Cockerell and
Eliot, 1905)”, the name used by California workers dur¬
ing this time period for what we have shown here to be
Doriopsilla roivena. Lance noted publication of this illus¬
tration on a separate sheet in his Species Folder
“Doriopsilla nigromaculata”. However, comparison of
this illustration with his sketches of direct development
in his species folder “White Porostome Spotted" reveals
it to be copied from a sketch for that species, which we
demonstrate here to be Dendrodoris nigromaculata.
Although their egg masses and spicule complement
as hatching juveniles are different, both species have
ametamorphic direct development (see below), and
Lance (1982) was probably mainly making a statement
about developmental mode in the one. The unexpected
twist is that the binomial used in the figure caption turns
out to be accurate for the species actually illustrated.
The specimens described and identified as D. rowena
by Bertseh and Aguilar-Rosas (1984) from El Tomatal,
on the Pacific coast of Baja California, are, based on their
larger size (up to 30 mm long) and color pattern, actually
Diaulula aurila Marcus and Marcus (1967), which is
common in that region (Bertseh et al. 1999, cited as “Sal
y pimenta” (Salt and pepper [dorid]); Goddard and
Schickel 2000, cited as Discodoris sp. 1 of Behrens
1991; personal observations).
Genus Dendrodoris Ehrenberg, 1831
Dendrodoris nigromaculata (Cockerell in Cockerell
and Eliot, 1905)
(Figures 1, 17-21)
Doridopsis vidua (?) [non Bergh, 1878], — Cockerell and
Eliot, 1905: 40^1.
Doris nigromaculata Cockerell in Cockerell and Eliot,
1905: 40-41.
Doridopsis nigromaculata (Cockerell in Cockerell and
Eliot, 1905). — Cockerell, 1908: 106.
Dendrodoris vidua [non Bergh, 1878] . — O'Donoghue,
1926: 212.
Dendrodoris nigromaculata (Cockerell in Cockerell and
Eliot, 1905). — O’Donoghue, 1926: 213; Steinberg,
1961: 59. Lance, 1982: 29.
Dendrodoris sp. Lee and Brophy, 1969: 20.
Dendrodoris sp. Behrens, 1980: 100; Behrens and
Gatewood, 1986: 139, 142.
Dendrodoris sp. b McDonald and Nybakken, 1980: 54;
McDonald, 1983: 171.
Dendrodoris sp. 3 Behrens, 1991: 72; Goddard, 2004:
1957, 1959, 1963.
Dendrodoris behrensi Millen and Bertseh, 2005: 189-
199; Goddard, 2005: 201-211; Behrens and Hennosillo,
2005: 86; California Academy of Sciences and Goddard,
2013: worksheets for Hill St. and So. Casa Reef.
Type Material:
Doris nigromaculata - Holotype: La Jolla, California,
July 1901 (NHMUK 1904.7.7.1), dissected by Sir C.
Eliot in 1905.
J.H.R. Goddard and A. Valdes, 2015
Page 39
Figures 17-21. Dendrodoris nigromaculata. 17. Living adult, 22 imn long, from Bahia Falsa, Baja California, May 2001. Included
as a paratype of Dendrodoris behrensi by Millen and Bertsch (2005). 18. Adult, no date or locality (Lance Collection, Species Folder:
White Porostome Spotted). 19. Egg mass, sketch labelled as South Casa Reef, La Jolla, 20 July 1974, but Lance Field Accounts
indicate adults were from Hill Street, San Diego, 19 July 1974 (Lance Collection, Species Folder: White Porostome Spotted).
20. Recently hatched juvenile, Aug. 1974 (pencil sketch in Lance Collection, Species Folder: White Porostome Spotted). 21. Juve¬
nile, 570 pm long, four days after hatching, June 2001. From egg mass laid by adult from Bahia Falsa, Baja California, May 2001
(Goddard 2005, as Dendrodoris behrensi).
Dendrodoris behrensi - Holotype: California, 14 Feb
1988 (CASIZ 69303). Paratypes: California, 14 Feb
1988 (CASIZ 171658); San Quintfn, Mexico, 27 May
2001 (CASIZ 171659), San Quintrn, Mexico, 27 May
2001 (CASIZ 171660).
Anatomy; The anatomy of Dendrodoris nigromaculata
was described by Cockerell and Eliot (1905) and Millen
and Bertsch (2005). As noted by Cockerell and Eliot
(1905), Lance (Lance Collection, Species Folder: White
Porostome Spotted), and Goddard (2005), notal spicules
are sparse and consist of straight to slightly curved rods
only, some of which are irregular in outline (Figures 1,
20-21).
External Morphology: The external morphology of
D. nigronmculata was described by Cockerell and Eliot
(1905) and Millen and Bertsch (2005), with additional
details presented by Lance (Lance Collection, Species
Folder: White Porostome Spotted). Adults grow to
27 mm long and are distinguished externally by their trans¬
lucent white ground color and chocolate brown blotches.
Page 40
THE NAUTILUS, Vol. 129, No. 1
the larger of which are usually clustered into three or
four groups centered mid-dorsally, and the smaller of
which are scattered toward the edges of the dorsum
(Figures 17-18).
Development: As described by Goddard (2005) and
Lance (Lance Collection, Species Folder: Wliite Porostome
Spotted) Dendrodoris nigromaculata has arnetamorphie
direct development in short, stout egg ribbons laid on edge
in a loose coil of only a turn or two (Figure 19). Juveniles
(Figures 20-21) hatch after an embryonic period of 38 days
at 16-19° C and are about 510 microns long.
Geographic Distribution: Dendrodoris nigromaculata
is known from the Monterey Peninsula, California south
to the San Benitos Islands, Baja California (McDonald,
1983; Behrens and Gatewood, 1986; Millen and Bertseh,
2005) (Figure 23).
Remarks: Bergh (1878) described the species Doiiojms
vidua based on specimens collected from Tahiti, French
Polynesia. The illustrations of the live animal (Bergh,
1878: pi. 1, figs. 17-20) represent an elongate Dendrodoris
with a white background color and numerous black spots
all over the dorsum, larger near the center of the animal,
and small dorsal tubercles. Based on the body shape and
coloration, D. vidua is most likely a senior synonym of
Dendrodoris elongate Baba, 1936. Cockerel and Eliot
(1905) tentatively reported this species from California as
Doridopsis vidua (?), based on a single specimen collected
v
-Puerto Pertasco, Son.
San Quintin. BC' — -ipercebu, qc
-PuerteCitos. BC
El Campo. BCS— ^ ' - . Bahia de Los Angeles. BC
• A
Guaymas. Sort.
< \
-v>- iS.la CerTalvo. BCS
Punta Pcrico. BCS- — ' . _
T — Mazatten. Sin
GULF OF
MEXICO
Jj
If
I,
&
I \
\ . (
% 7
PACIFIC OCEAN
CARIBBEAN
SEA
Pacific coast of Costa Rica
Islas Secas. Panama
x
Isla Uva, Panama / ‘
Figure 22. Map showing collection localities of Doriopsilla
rowena.
Pacific Grove, CA
Carmel Bay, CA
Naples Reef. Santa Barbara Co.. CA
Jk
i'
Point Dume, Malibu, CA
San Miguel Island, CA—
Anacapa Island, CA W
Santa Cruz Island, CA
Santa Catalina Island, CA y
San Clemente Island, CA
Bahia Falsa, San Quintin, BC
Islas San Benito, BC-
PACIFIC OCEAN
— Santa Monica Bay, CA
Redondo Beach, CA
. A- Palos Verdes Peninsula, CA
t Jy — Laguna Beach, CA
La Jolla, CA
Point Loma, San Diego, CA
Islas Coronados, BC
\
V/'
Figure 23. Map showing collection localities of Dendrodoris
nigromaculata.
in La Jolla. Cockerel and Eliot (1905) noted some differ¬
ences between their specimen and Bergh s (1878) original
description in several details, including the coloration
and mantle margin width. Cockerel and Eliot (1905) also
inchoated that their record was “suspicious” considering
the geographic distance between the type locality and
California, thus they introduced Cockerell’s new name
Doridopsis nigromaculata in case the specimen was ulti¬
mately proven to belong to a distinct species. As men¬
tioned above, examination of the specimen studied by
Cockerel and Eliot (1905) and therefore the holotype
of D. nigromaculata (NHMUK 1904.7.7.1) revealed
that it is a species of Dendrodoris. The holotype of
D. nigromaculata has three pairs of large dark spots and
several smaller spots irregularly distributed (Figure 1),
very different from the original description of Doriopsis
vidua by Bergh (1878) and references to D. nigromaculata
by other authors, but nearly identical to the original
description of D. behrensi by Millen and Bertseh (2005).
Lance was aware of this species as early as 1961 (or
possibly even 1953) and referred to it in his notes and
illustrations first as the “erenulate dorid,” then as the
“white dendrodorid” or “brown-spotted Dendrodoris”
(Lance Collection, Species Folder: White Porostome
Spotted), and in his field accounts as “ Dendrodoris sp. true
dendrodoris” (19 July 1974, Hill St) and “ Dendrodoris sp.
3 [following Behrens 1991] white porostome” (26 Apr.
1998, So Casa Reef). As evidenced by an undated, hand¬
written description and pen and ink illustration of an adult
(Figure 18), Lance started to formally describe it under
the manuscript name “Dendrodoris barbarensis,” based on
five specimens collected in the 1950s and 60s from Naples
Reef, Santa Barbara County; Point Lima, San Diego; and
tire Coronado Islands. Lances folder for this species con¬
tains no evidence diat he ever associated Cockerell and
Eliots (1905) description of D. nigromaculata widi it.
J.H.R. Goddard and A. Valdes, 2015
Page 41
F.M. MacFarland collected D. nigromaculata on the
Monterey Peninsula in the 1920s (see Millen and Bertsch
2005, Material Examined). However, MacFarland (1966)
does not mention those specimens nor refer to Cockerell
and Eliot’s 1905 description of D. nigromaculata.
DISCUSSION
The taxonomic confusion surrounding Dendrodoris
nigromaculata and Doriopsilla rowena likely has two
main sources, the first being the historical controversy
over the validity of Bergh’s genus Doriopsilla, and the
second being Lance’s overlooking of Cockerell and
Eliot’s description of Dendrodoris nigromaculata as he
was describing “Dendrodoris barbarensis.” Lance was
clearly familiar with Cockerell and Eliot’s 1905 paper,
and in hindsight their description of Dendrodoris
nigromaculata is unequivocal, especially with respect
to color pattern, so how could Lance have ended up
misapplying that name to the smaller and differently col¬
ored species we have shown here to be Doriopsilla
rowena ? The type locality, combined with a view of habi¬
tat fidelity for nudibranehs, may be keys. After a decade of
collecting along the coast of San Diego County Jim Lance
considered himself intimately acquainted with the inter¬
tidal nudibranehs there (see Steinberg 1961: 59), and
must have wondered why he had never found Cockerell’s
D. nigromaculata from 60 years earlier. Thus, in 1967
when Lance did find in La Jolla a dendrodoridid
with brown spots that was new to him, he was primed
to recognize it as Cockerell’s missing Dendrodoris
nigromaculata , overlooking that that name better applied
to his “Dendrodoris barbarensis” which he had already
seen from other locations in southern California. In any
case, once Lance misapplied the name, new workers in
the field followed his lead and considerable authority,
especially with regards to the fauna of southern California
(see Steinberg, 1961; Lance, 1961, 1966). The inertia
gained by this use of nigromaculata subsumed rowena
for decades among California workers, helped muddle
the generic distinctions between Dendrodoris and
Doriopsilla, and steered Millen and Bertsch (2005) away
from considering Cockerell’s nigromacidata when they
described it as Dendrodoris behrensi. Cockerell’s type
specimen of Dendrodoris nigromacidata had effectively
become a cold case, sitting on a shelf half a world away in
the Natural History Museum in London, its type status
unknown to the museum curatorial staff.
Doriopsilla roivena from the Pacific coast of California
and Baja California lay large eggs and have ameta-
morphic direct development. However, based on
the limited information presented by Goddard and
Hermosillo (2008), members of the Panamic population
appears to have planktotrophie development, which is
consistent with a geographic range stretching from the
northern Gulf of California to Panama and the Galapagos
Islands. However, if planktotrophy is confirmed, including
at the type locality of D. rowena in Puerto Pehasco, then
the direedy developing population from California and
the Pacific coast of Baja California likely represents an
undescribed cryptic species. An alternative explanation is
that this species displays poecilogony. Aside from its
rarity, there are no confirmed examples of poecilogony
that include such disparate modes of development, thus
we consider the first hypothesis as the most likely.
Doriopsilla gemela Gosliner, Schaefer, and Millen, 1999
exhibits a similar but geographically reversed variability,
with planktotrophie development in California and direct
development in the Gulf of California (Goddard, 2005;
and see Lance Collection, Species Folder: Gulf Yellow
Porostome), and has recently been found to comprise two
species (Hoover et id., in preparation). Genetic confirma¬
tion of the two forms of D. roivena as separate species
would bring the total number of nominal species of
dendrodoridid nudibranehs in the greater region to eleven.
ACKNOWLEDGMENTS
Terry Gosliner and Liz Kools (CASIZ) loaned us
specimens collected by Jim Lance for examination and
provided us access to the Lance Collection. Andreia
Salvador (NHMUK) and Chris Meyer (USNM) pro¬
vided us with photographs and information on the type
specimens of D. nigromacidata and D. rowena respec¬
tively. Suggestions by Terry Gosliner and Sandra Millen
improved the manuscript substantially.
LITERATURE CITED
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poblaciones de opistobranquios (Mollusca: Opisthobranchia)
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Departamento de Pesquerias y Biologia Marina, Centro
Interdisciplinario de Ciencias Marinas, La Paz, Baja
California Sur, Mexico.
Angulo-Campillo, O. 2005. A four year survey of the opistho-
branch fauna (Gastropoda, Opisthobranchia) from Baja
California Sur, Mexico. Vita Malacologica 3: 43-50.
Behrens, D.W. 1980. Pacific Coast Nudibranehs. Sea Chal¬
lengers, Monterey, 112 pp.
Behrens, D.W. 1991. Pacific Coast Nudibranehs. 2nd Edition.
Sea Challengers, Monterey, 107 pp.
Behrens, D.W. and J. Gatewood. 1986. New opisthobranch
records for the west coast of Baja California. Shells and
Sea Life [Opisthobranch Newsletter] 18: 139-142.
Behrens, D.W. and A. Hermosillo. 2005. Eastern Pacific Nudi-
branchs. Sea Challengers, Monterey, 137 pp.
Bertsch, H. and L. Aguilar-Rosas. 1984. Range extensions of
four species of nudibranehs along the Pacific coast of Baja
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THE0NAUTILUS
Volume 129, Number 2
June 25, 2015
ISSN 0028-1344
CONTENTS
THE NAUTILUS 129(2):43-53, 2015
Page 43
Paleocene and Miocene Thyasira sensu stricto (Bivalvia:
Thyasiridae) from chemosynthetic communities from Japan
and New Zealand
Kazutaka Amano
Department of Geoscience
Joetsu University of Education
Joetsu 943-8512^ JAPAN
amano@juen .ac.jp
Robert G. Jenkins
School of Natural System
College of Science and Engineering
Kanazawa University
Kanazawa City, Ishikawa
920-1192, JAPAN
[email protected]
Crispin T.S. Little
School of Earth and Environment
University of Leeds
Leeds LS2 9JT, UNITED KINGDOM
[email protected]
Kathleen A. Campbell
Earth Sciences Programme
School of Environment, Faculty of Science
University of Auckland
Private Bag 92019, Auckland Mail Centre
Auckland 1142, NEW ZEALAND
[email protected]
Kristian P. Saether
State Key Laboratory of Palaeobiology and Stratigraphy
Nanjing Institute of Geology and Palaeontology
39 Beijing East Road
Nanjing 210008, CHINA
[email protected]
ABSTRACT
A new species of bivalve, Thyasira ( Thyasira ) beui, is described
from lower to middle Miocene hydrocarbon seep deposits from
the North Island of New Zealand. Thyasira (T.) nakazawai
Matsumoto, 1971 is redescribed from lower Miocene seep
deposits in central Honshu of Japan, and T. (T.) sp. from
Paleocene wood-fall communities in eastern Hokkaido is
described as the first Thyasira sensu stricto of this age in Japan.
As the genus Conchocele replaced the niche of Thyasira sensu
stricto at seep sites from the Eocene in Japan, the occurrence of
T. (T.) nakazawai is an exceptional occurrence of this genus at
younger seeps in Japan. In contrast, Conchocele disappeared
from New Zealand waters from the end of the Paleocene,
leaving Thyasira sensu stricto as the sole thyasirid taxon at
New Zealand Cenozoic seep sites.
Additional Keywords: Conchocele, Fossil, hydrocarbon seep,
cold seep
INTRODUCTION
Bivalves within the family Thyasiridae today inhabit
reduced environments from intertidal mudflats to
deep-sea hydrothermal vents. Some thyasirid species
host chemoautotrophic bacteria in their gills, particu¬
larly those toward the larger end of the size range of
the family, and some do not (Dufour, 2005; Oliver and
Levin, 2006; Taylor and Glover, 2010). Most species
within the genus Thyasira have two demibranchs
and symbionts (Oliver and Killeen, 2002; Dufor,
2005). Such chemosymbiotic thyasirids are deep bur-
rowers and mine sulfide deep in the substrate using
their vermiform foot (Dando and Southward, 1986;
Seilacher, 1990; Oliver and Killeen, 2002; Dufour and
Felbeck, 2003). Thyasirid species can extend their foot
up to 30 times the length of the shell (Dufour and
Felbeck, 2003).
The oldest known thyasirid, Cretaxinns hurumi Hryniewicz,
Little, and Nakrem, 2014, comes from uppermost Jurassic
to lowermost Cretaceous seeps in Svalbard. As noted
by Kiel et al. (2008), Thyasira rouyana (d’Orbigny,
1844) from Lower Cretaceous (Valangian-Hauterivian)
rocks in Europe is the oldest species of Thyasira sensu
stricto. Bv the late Early Cretaceous (Albian), spe¬
cies within this subgenus appeared in seep sites in
Hokkaido, northern Japan (Kiel et al., 2008, 2009).
Several species of Thyasira sensu stricto have been
reported from Cenozoic deposits around the Pacific
Rim (Table 1), including Thyasira sensu stricto from
Paleocene carbonates with plant debris in eastern
Hokkaido, Japan, Thyasira nakazawai from Miocene
aceretionary-prism deposits in central Honshu, Japan
(Matsumoto, 1971), and Thyasira sp. from Miocene
seep deposits of North Island, New' Zealand (Campbell
et al., 2008).
Here we formally describe Thyasira sensu stricto
fossils from the Paleocene and Miocene of Japan and
the Miocene from New Zealand. The descriptions extend
knowledge of the fossil species of Thyasira sensu stricto
Page 44
THE NAUTILUS, Vol. 129, No. 2
Table 1. Cenozoic Thyasira sensu stricto from the Pacific Him. Symbols: * maximum length (mm); ** length >height; + distinctly
longer than high, ± subcircular, - distinctly higher than long; *** medial flattened area.
in hydrocarbon seep and wood-fall communities from the
Pacific Rim.
MATERIALS AND METHODS
The fossils used in this study were collected from two
Paleoeene sites in japan and from two Miocene locali¬
ties in New Zealand. We also examined some Miocene
specimens from New Zealand housed at the University
of Auckland and one Miocene species described by
Matsumoto (1971), which is stored at National Museum
of Nature and Science, Tokyo. Details of localities and
associated faunas are as follows:
Eastern Hokkaido, Japan. All specimens were collected
from two carbonate float blocks from the Katsuhira-zawa
(Kl) and Katsuhira-kita-zawa (K2) localities of Urahoro
Town, eastern Hokkaido, Japan (Figure 1). The upper
part of the Katsuhira Formation crops out in this area
and consists of mudstones yielding carbonates that con¬
tain many plant fragments. The carbonates contain¬
ing the thyasirid fossils have probably been eroded out
from the mudstones of this formation, the age of which
has been assigned to the Paleoeene (early Selandian)
(see Amano and Jenkins, 2014). In addition to the thyasirid
bivalves, the carbonates also contain specimens of a
provannid(?) gastropod, a limpet, and Bentharca steffeni
(Amano et ah, 2015). The taxonomic composition of this
fauna suggests that its primary energy source was the
degradation products derived from sunken wood, which
was probably bored by xylophagain bivalves.
Central Honshu, Japan. Thyasira nakazawai Matsumoto,
1971 was the name proposed for specimens collected
from limestone lenses or calcareous mudstones within
turbidites of the Wappazawa Formation (Setogawa Group)
on a branch of the Hakkou River, 1600 m west of
Matsushita, Shimada City (SI) and at Nakadaira, Shimada
City (S2) (Figure 1). The age of the Wappazawa Forma¬
tion has been assigned to the early Miocene (Watanabe,
1988). From the formation, molluscan fossils have been
recovered only from the limestone lenses and calcar¬
eous mudstones (Matsumoto, 1971). Thyasira nakazawai
was collected with Saxolucina ( Megaxinus ) matsmhitai
Matsumoto, 1971 and Pitar matsuraensis (Nagao, 1928)
[= Pliocardia ? sp.J. Based on the fauna and lithofacies of
the limestone lenses and calcareous mudstones, the taxa
from these localities probably inhabited hydrocarbon seeps.
K. Amano et al., 2015
Page 45
Hokkaido;
North Island
4 ...,y
•-K^Japan - Sear*'
South Island
TH
Gisborne
iZke
W&karQ.'n&jfw
| ~~| Pliocene - Quaternary
I | Eocene • Miocene
mid-Cretaceous - Paleocene
KM basement, volcanics, etc
O seep limestone site
£ seep site treated in this paper
HAWK
BAY
Ra^ifiGi®G'e'anl
B.OQKrfil
Figure 1. Localities of the fossil Thy anira sensu stricto described herein.
North Island, New Zealand. Specimens of Thyasira
beui new species were collected from hydrocarbon seep
carbonates from the Moonlight North (MN), Bexhaven
(BX), and Turihaua (TH) localities, north of Gisborne,
North Island, New Zealand (Figure 1). The deposits
belong to the Bexhaven Limestone, which is assigned
to the early to middle Miocene (Campbell et al, 2008).
From MN, Amano et al. (2014) described the vesieoinyid
species Notocalyptogena neozelandica and Pliocardia ?
sp. Saether et al. (2010) described the bathymodioline
mussels, Bathymodiolus (s. 1.) heretaunga , from BX and
MN, and Gigantidus coseli from BX, MN, and TH.
We describe the Thyasira species in this study using
the terminology of Kauffman (1967) and Oliver and
Killeen (2002). All figured and supplementary specimens
are catalogued at the University of Auckland (UOA L),
Joetsu University of Education (JUE) and the National
Museum of Nature and Science (NSM).
SYSTEMATIC PALEONTOLOGY
Family Thyasiridae Dali, 1900 (Dali, 1895)
Genus Thyasira Lamarck, 1818
Subgenus Thyasira Lamarck, 1818
Type Species: Tellina flexuosa Montagu, 1803
Remarks: The subgenus Parathyasira Iredale, 1930 dif¬
fers from Thyasira sensu stricto by having no shell auricle.
Most historical and some recent literature has treated
the taxon Conchocele as a subgenus of Thyasira (e.g.,
Yabe, H. and S. Nomura, 1925; Grant and Gale, 1931;
Krishtofovich, 1936; Slodkewitsch, 1938; Weaver, 1942;
Hickman, 1984; Matsui, 1985; Moore, 1988; Matsui, 1990).
However, Conchocele Gabb, 1866 attains a large size
(max. 165.4 mm in length; Kamenev et al., 2001), has a
thick shell, and lacks an auricle. Therefore, we regard
Conchocele as a genus distinct from Thyasira.
Thyasira ( Thyasira ) nakazatvai Matsumoto, 1971
(Figures 2-7)
Thyasira nakazawai Matsumoto, 1971: 665-666, pi. 3,
fig. 1 5—18, Amano, 2014: 7, fig. 1.
Type Material: Holotype, NSM PM-16922a. Paratypes,
NSM PM -16923, NSM PM-16924, NSM PM-16925.'
Material Examined: Eleven specimens including the
type specimens.
Measurement: See Table 2.
Original Description: “Shell medium in size, thin tri¬
gonal oval, nearly long as high, strongly inflated. Antero-
dorsal border strongly concave, sharply turned to broadly
curved, subangular ventral border forming almost a right
angle; postero-dorsal long, faintly arched passing into
the ventral border forming an obtuse angle. Beak small,
strongly curved forward and situated at about the middle
of the shell. Surface of the shell ornamented with fine
and concentric, but somewhat irregular growth-lines.
Posterior surface depressed from the upper side of the
postero-ventral corner making oblique ridge. A central
part of the shell faintly ridged from the beak to middle of
the ventral border.”
Complementary' Description: On examination of the
material we found that there are some elements of the
Page 46
THE NAUTILUS, Vol. 129, No. 2
Figures 2-7. Thyasira ( Thyasira ) nakazawai Matsumoto. 2. Dorsal view of posterior part of left valve; Paratype; NSM PM-16924;
Lot. SI. 3a, b. Frontal and oblique view of right valve; NSM PM-16910; Lot. S2. 4. Frontal view of left valve; Paratype; NSM
PM-16923; Loc. SI. 5. Frontal view of left valve; Holotype; NSM PM-16922a; Loe. SI. 6. Frontal Mew of left valve; NSM PM-16905;
Loc. SI. 7. Inner surface of right valve; AAS, anterior adductor scar; NSM PM-16909; Loc. SI.
Table 2. Measurements of Thyasira ( Thyasira ) nakazawai Matsumoto.
original description of the species that are incorrect. We
therefore offer here more accurate and complementary
morphological information.
Shell rather large for the genus (maximum 28.3 mm
in length), thin, ovate, slightly longer than high (height/
length ratio = 0.85-0.97; exceptionally 1.03), well inflated
(width/length ratio = 0.70). Anterodorsal margin short,
strongly concave; anterior margin subcircular and gradu¬
ating into arched ventral margin. Second posterior fold
distinct, hut not stronger than first posterior fold; pos¬
terior sulcus rather shallow and narrow; first poste¬
rior fold strong and ridged; submarginal sulcus distinct;
auricle narrow but extending total length of submarginal
sulcus. Lunule moderately depressed. Beak prominent,
prosogyrate, situated at about one-third of shell length.
Shell surface ornamented with fine growth lines. Ante¬
rior adductor scar elongate quadrate and attached to
pallial line; posterior adductor scar indistinct. Inner
surface of shell crenulated by many fine radial lines.
Comparison: Thyasira ( Thyasira ) nakazaivai is simi¬
lar to T. (7’.) tanahei Kiel, Amano and Jenkins, 2008 from
the Upper Cretaceous formations in Hokkaido, sharing
a strongly concave anterodorsal margin and strong and
K. Amano et al., 2015
Page 47
ridged posterior fold. However, T. (T.) nakazawai differs
from the latter species by having a larger (maximum
length of T. (T.) tanabei = 13.5 mm) and more inflated
shell with a smaller anterior adductor scar.
Distribution: Lower Miocene Wappazawa Formation
of the Setogawa Group from the Shizuoka Prefecture,
central Honshu, Japan.
Thyasira (Thyasira) beui new species
(Figures 8-15)
Thyasira sp. — Campbell et al., 2008: 90.
Thyasira sp. nov. — Saether, 2011: 135-138, fig. 5-19.
Diagnosis: Medium-sized Thyasira with suborbicular
shell, shallow lunule, and small auricle. Ventral end of first
posterior fold occasionally angulated.
Description: Shell up to 13.8 mm in length, rather thin,
moderately inflated (width/length ratio = 0.58-0.91), sub-
orbicular (height/length ratio = 0.90-1.17), equivalve and
inequilateral. Antero-dorsal margin broadly arched and
continuing to rounded anterior margin; ventral margin
broadly arched. Auricle small, extending in length two-
thirds along marginal sulcus; first posterior fold sharp,
with ventral end occasionally angulated; posterior sulcus
very shallow; second posterior fold less distinct than first
posterior fold. Beak prominent, prosogyrate and located
around two-fifths of shell length (i.e., at 36-44% of shell
length from anterior margin). Lunule shallow and demar¬
cated by very shallow groove. Shell surface with fine
growth lines. Inner shell surface ornamented by many
fine radial grooves. Pallial line entire, starting from mid¬
point of ventral side of anterior adductor scar. Anterior
adductor scar elongate-quadrate; posterior adductor scar
very small and ovate.
Holotype: UOA L4626 from MN (YI6/f 1054), collec¬
tion AU 15844.
Paratypes: UOA L4627 from MN (Y16/1033), col¬
lection AU 19618; UOA L4628 from MN (Y16/H174),
collection AU 19923; UOA L4629 and L4630 from MN
(Y16/1059), collection AU 19982; UOA L4631 from BX
(Y16/1032), collection AU 19617.
Type Locality: Moonlight North seep carbonates, north
of Gisborne, North Island, New Zealand.
Material Examined: Twenty-two specimens from three
localities (Loe. MN, BX, TH in Figure 1).
Measurements: See Table 3.
Remarks: Thyasira ( Thyasira ) beui is Thyasira sp. in
the compilation of molluscan fossils (in part taken from
Beu and Maxwell (1990)) from New Zealand hydrocar¬
bon seep carbonates in Campbell et al. (2008). Saether
(2011) described and illustrated this species as “ Thyasira
sp. nov. in his unpublished Ph.D. thesis.
Comparison: Thyasira ( Thyasira ) beui shares a promi¬
nent beak and a moderately inflated shell with T. (T.)
motutaraensis Powell, 1935 from the lower Miocene
Motutara deposit west of Auckland, North Island,
New Zealand (see also Beu and Maxwell, 1990). How¬
ever, T. (T. ) motutaraensis can be separated from the
new species by its smaller and higher triangular shell
(length = 7.4 mm, height/length ratio = 1.14). Thyasira
(T.) planata Marwick, 1926 [this name was preoccupied
by Jeffreys, 1882 and a new name, T. (T.) marwicki
is proposed herein] from upper Miocene deposits in
the western part of North Island, New Zealand, can
be distinguished from T. (T.) beui by having a wider
posterior area, a longer marginal sulcus, and a nar¬
rower auricle than that of the new species. Thyasira
(T.) mironovi Kalishevich from the Paleoeene of South
Sakhalin (Klishevieh et al. 1981) is similar to T. (T.) beui
in having a first posterior fold with angular ventral
end. However, T. (T.) mironovi can be separated from
T. (T.) beui by having a less inflated shell, a wider pos¬
terior fold and a weak medial flattened area. Another
species from the Paleoeene of South Sakhalin, T. (T.)
uncinata Kalishevich, can be easily distinguished from
T. (T) beui by having an elongate shell with posteriorly
situated beak. Thyasira (T.) bartrumi Powell, 1935 from
the lower Miocene Motutara deposit is distinctly dif¬
ferent from T. (T.) beui by having a Conchocele-\ike shell
witli beak at anterior one-seventh of shell length and a
medial flattened area. The Recent New Zealand species,
T. (T.) peregrina Iredale, 1930 differs from T. (71) beui by
its smaller shell (maximum length = 10.4 mm), which is
higher than long, and by having a medial flattened area.
Distribution: Lower to middle Miocene Bexhaven
Limestone, north of Gisborne, North Island, New Zealand.
Etymology: Named after Dr. Alan G. Beu who has
made significant contributions to the taxonomy of
Cenozoic fossil faunas from New Zealand.
Thyasira ( Thyasira ) sp.
(Figures 16-17)
Material Examined: Two articulated but imperfect
specimens (JUE nos. 15936, 15937).
Description: Shell rather small in size (9.7-10.1 mm +
in length), thin, ovate, longer than high, well inflated
(width/length ratio = 0.52-0.58). Anterodorsal margin
short, nearly straight; anterior margin subcircular. Second
posterior fold distinct; posterior sulcus rather deep; first
posterior fold wide and ridged; submarginal sulcus dis¬
tinct; auricle narrow and short. Beak prosogyrate. Inner
structure of shell not preserved.
Comparison: Thyasira ( Thyasira ) sp. is similar to the
Cretaceous species, T. (T.) tanabei Kiel, Amano and
Jenkins, 2008 by having a ridged first posterior fold.
However, the wide posterior area of our specimens
enables us to separate T. (T.) sp. from T. (T. ) tanabei.
Page 48
THE NAUTILUS, Vol. 129, No. 2
Figures 8-15. Thyasira (Thyasira) betii new species. All specimens except for one illustrated in Figure 15a, b are from the type
locality (Moonlight North; MN). One specimen of Figure 15a, b is from Bexhaven (BX). 8a-c. Frontal and dorsal views of both
valves; Paratype, UOA L4629. 9a, b. Frontal view of right valve and dorsal view of both valves; Holotype; UOA L4626; white arrow
showing an angulated ventral end of first posterior fold. 10a, b. Inner surface of right valve and its enlargement of the area around
AAS (= anterior adductor scar); Paratype; UOA L4630. 11. Frontal view of left valve; Paratype; UOA L4628. 12. Frontal view of
right valve; UOA L4638. 13. Frontal view of left valve; left valve; UOA L464Q. 14. Frontal view of right valve; UOA L462/. Inner
surface of right valve showing PAS (= posterior adductor scar) and its enlargement; Paratype; UOA L4631.
T. (T. ) xylodia Kiel and Goedert, 2007 comes from
latest Eocene and early Oligocene wood-fall communi¬
ties in Washington State, USA and can be distinguished
from T. (T.) sp. by its larger size (21 mm in length),
deeply concave antero-dorsal margin and narrower pos¬
terior area. T. (71) baca Devjatilova from the Pal eocene
Getldlninskaya Formation of western Kamchatka (Devjatilova
and Volobueva, 1981) differs from T. (71) sp. by having
a triangular shell and narrower posterior area. Thyasira
(71) mironovi Kalishevieh can be distinguished from
T. (T.) sp. by having wider first posterior fold with an
angular ventral end and extending its ventral end to
the ventral margin of main disc.
Distribution: Paleocene, upper part of the Katsuhira
Formation, eastern Hokkaido, Japan.
DISCUSSION
Several Recent species of Thyasira sensu stricto have
been recorded from hydrocarbon seep or hydrothermal
vent sites (Table 4; Clarke, 1989; Dando et ah, 1994;
Oliver and Killeen, 2002; Olu et ah, 2004; Oliver and
Sellanes, 2005; Oliver and Holmes, 2006; Rodrigues
et ah, 2008). Based on recent molecular analysis of nuclear
18S rRNA and 28S rRNA, the subgenus Thyasira sensu
stricto is divided into two clades (Taylor et ah, 2007).
Thyasira (T.)sarsi (Philippi, 1845b) and T. (71) methanophila
Oliver and Sellanes, 2005, from hydrocarbon seeps, form
a monophyletic clade. Thyasira sarsi itself is an oppor¬
tunistic species which is also able to live in sediments
with low organic content, and at relatively low densities
(Keuning et ah, 2011). Another clade includes T. (71)
jlexuosa (Montagu, 1803), T. (71) gouldii (Philippi, 1845a),
and T. (T.) polygonata (Jeffreys, 1864), none of which have
been recorded from seep and vent sites. Morphologi¬
cally, the T. (T.) sarsi-T. (71) methanophila clade differs
from the T. (II) flexuosa-T. (T.) gouldii-T. (71) polygonata
clade by having larger (more than 20 mm in length),
subcircular or a slightly longer shells, without a medial
flattened area. Other Thyasira species found in seep
and vent sites, such as T. (T.) southwardae Oliver and
Holmes, 2006, 71 (71) vulcolutre Rodrigues and Oliver
in Rodrigues et ah, 2008 and T. (T.) oleophila Clarke,
1989, also have similar shell characteristics to the T. (71)
sarsi-T. (T.) methanophila clade.
Payne and Allen (1991) and Dufour (2005) have shown
that in thyasirids demibranch number is related to
body size, because asymbiotic thyasirids with only one
demibranch only have access to a small amount of nutri¬
ents at bathval depths. All the species discussed above
have two demibranchs and chemosynthetic bacteria
(Dufour, 2005; Oliver and Sellanes, 2005; Oliver and
Holmes, 2006; Rodrigues and Oliver, 2008). Almost
certainly because of the abundant supply of hydrogen
sulfide at seep and vent sites, thyasirids living there can
grow to large sizes relative to thyasirids inhabiting other
environments. However, the reason that the thyasirids
living in chemosynthetic environments have subcircular
or longer shells, without a medial flattened area, is
unknown. There are exceptions, as T. (T) striata (Sturany,
1896), found at a Mediterranean seep by Oln et al. (2004),
is characterized by a rather small (ca. 7.5 mm) and higher
shell with a medial flattened area. This morphological
information from Recent seep and vent Thyasira sensu
stricto can be used to infer the paleoecology of fossil
Thyasira sensu stricto species.
As shown in Table 1, Thyasira (71) nakazawai has a
large (length = 28.3 mm) and longer shell (height/
length ratio = 0.85-0.97) without a medial flattened area,
Page 50
THE NAUTILUS, Vol. 129, No. 2
Figures 16-17. Thyasira (' Thyasira ) sp. 16a, b. Frontal view of right valve and dorsal view ofboth valves; JUE no. 15936; Loe. Kl.
17a, b. Frontal and oblique view of right valve; JUE no. 15937; K2.
Table 4. Morphology of recent species of Thyasira sensu strieto. * maximum length (mm); ** Length > Height; + distinctly longer
than high, ± subcircular, — distinctly higher than long; *** Medial flattened area.
compared to Thyasira sensu strieto species from Cenozoic
deposits around the Pacific Rim. Because of this, we
speculate that T. (T. ) nakazawai might have lived in
cold seep areas. In contrast, T. ( T minoensis Itoigawa,
1960 was collected from non-seep sandstones of the
lower Miocene Oidawara Formation; it has a smaller
(length = 14.1 mm) and higher shell (height/length
ratio = 1.08) and with a distinct medial flattened area.
K. Amano et al., 2015
Page 5 1
While the maximum size of T. ( T .) beui new species
is not large (length = 13.8 mm), the species has a
suborbieular shell (height/length ratio = 0.90-1.17)
without a medial flattened area. The carbonates and
associated fauna (see also Campbell et ah, 2008) indi¬
cate this species also thrived at fossil seep sites. Judging
from the lithofacies and the associated fauna of limpets
and provannids, the Paleocene T. (T.) sp. collected from
eastern Hokkaido might have been a member of a fossil
wood-fall community. Despite the small size of T. (7’.) sp.
(length = 9.7-10.1 mm) compared with other seep spe¬
cies, it also is longer than high and has no medial flat¬
tened area. Such small thyasirid species also have been
recognized in Late Cretaceous wood-fall communities
with limpets and provannids by Kiel et al. (2009). In the
northern Pacific area, Thyasira sensu stricto occurred
in hydrocarbon seeps and wood-fall sites during the
Late Cretaceous (Kiel et ah, 2008, 2009). The eastern
Hokkaido Paleocene Thyasira sensu stricto species
might have lived in wood-fall communities. The first large
thvasirid, Thyasira toumsendi (White, 1890) (almost cer¬
tainly a species of Conchocele) appeared in Maastriehtian
seep deposits of Snow Hill Island, Antarctica (Kiel et ah,
2008; Little et al. 2015). The second oldest large thyasirid
species, Conchocele aff. conradi (Rosenkrantz, 1942), is
from die Danian Kangilia Formation of western Greenland
(Rosenkrantz, 3970; Amano, 2014). From Nortii Island,
New Zealand, one specimen of Conchocele sp. has been
recorded from Paleocene deposits at Angora Road, south
of Wimbledon (Beu and Maxwell, 1990; Beu, 2014 per¬
sonal communication). Another Paleocene Conchocele
specimen up to 70 mm in length was collected from
1 km south of Te Kaukau Point, White Rock, South
Wairarapa coast (Beu, 2014 personal communication).
So far, no fossil Conchocele has been recorded from
Paleocene deposits in the northern Pacific area.
Since the Eocene, the genus Conchocele seems to
have replaced the niche of Thyasira sensu stricto in the
northern Pacific. Lots of literature has described the
radiation of Conchocele in Eocene to Recent times in
this region (e.g., Yabe and Nomura, 1925; Grant and
Gale, 1931; Krishtofovich, 1936; Slodkewitsch, 1938;
Weaver, 1942; Hickman, 1984; Moore, 1988; Kamenev
et ah, 2001). Conchocele was also found from Eocene
to Holocene seep sites and in Oligocene to Miocene
whale-fall sites (Goedert et ah, 1995; Majima et ah, 2005;
Amano et ah, 2007; Kiel and Goedert, 2006). Thus the
occurrence of Thyasira ( Thyasira ) nakazatuai from lower
Miocene seep deposits is an exceptional post-Eocene
occurrence of Thyasira ( Thyasira ) species in the north¬
ern Pacific area. In contrast, in New Zealand T. (71 ) beui
occurs in lower to middle Miocene seep sites, in the
absence of Conchocele from the region.
Conchocele might have migrated from western
Greenland to the northern Pacific area (including
Japan) by the Eocene (Amano and Jenkins, 2014), and
once there to have replaced Thyasira sensu stricto
because of its tolerance to lower oxygen environments.
In New Zealand waters, in contrast, Conchocele did not
invade hydrocarbon seep sites and had disappeared
from the region by end of the Paleocene. Thyasirids
(probably Thyasira sensu stricto) from New Zealand
Cretaceous seep deposits (Kiel et ah, 2013) show that
small sized thyasirids have flourished in the area since
that time period.
ACKNOWLEDGMENTS
We thank Alan G. Beu (GNS Science) for supplying
much information, allowing us to refer to the Paleocene
Conchocele from New Zealand and reviewing the manu¬
script; Graham P. Oliver (National Museum Wales), and
John D. Taylor (Natural History Museum) for infor¬
mation on Recent Thyasira ; Bruce Marshall (Te Papa
Museum, Wellington, New Zealand) for showing us
modem Thyasira specimens from New Zealand; Steffen
Kiel (University of Gottingen) for reviewing the manu¬
script; Neville Hudson (University of Auckland, New
Zealand) for his help with fossil curation and access to
material stored in the University of Auckland paleon¬
tological collections; Anton Oleinik (Florida Atlantic
University) for information on Russian literature; Tomoki
Kase (National Museum of Nature and Science. Tokyo),
Tatsuo Oji (Nagoya University Museum), and Hiroshi
Nishi and Jun Nemoto (Tohoku University Museum) for
their help in examining the fossil specimens from Japan.
This study was partly supported by a Grant-in-aid for
Scientific Research from the Japan Society for Promo¬
tion of Science (C, 26400500, 2014-2016) to K.A. and
R.G.J. K.P.S. was financially supported by the National
Science Foundation of China (No. 91114201) and the
Strategic Priority Research Program (B) of the Chinese
Academy of Sciences (XDB03010101). Fieldwork to
the New Zealand seep sites by C.T.S.L. was funded by
a Royal Society International Exchange grant.
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THE NAUTILUS 129(2):54-62, 2015
Page 54
A revision of the fossil taxa assigned to Hijperaulax
(Gastropoda: Odontostomidae), with the description
of a new genus (Gastropoda: Bulimulidae)
Kurt Auffenberg1
John Slapcinsky
Roger W. Portell
Florida Museum of Natural History
University of Florida
P.O. Box 'll 7800
Gainesville, FL 32611 USA
ABSTRACT
Orthalicoid terrestrial snails recorded from the lower Miocene
portion of the upper Oligocene to lower Miocene Tampa
Member of the Arcadia Formation (Hawthorn Group) of south¬
ern Florida and the lower Miocene St. Marks Formation of
northern Florida are reviewed. These taxa, previously allocated
to the genus Hijperaulax Pilsbry, 1897 (Odontostomidae), are
reassigned to Tocobaga new genus (Bulimulidae) on the basis
of a distinctive suite of morphological characters, particularly
those of the peristome and the embryonic whorl sculpture.
Examination of all type material of the fossil taxa histori¬
cally assigned to Hijperaulax reveals that only three species
are separable ( Partula americana Heilprin, 1886; Bulimulus
americanus wakullae Mansfield, 1937; and Bulimus floridanus
Conrad, 1846). The varietal names Bulimulus americanus var.
partulinus and B. americanus var. laxus, both Dali, 1890, are
indistinguishable from the nominate form in any important
morphological character. Bulimulus heilprinianus , Bulimulus
steamsii, both Dali, 1890, and Bulimulus ballistae, Bulimulus
remolina , Bulimulus tampae, and Bulimulus tortilla, all Dali,
1915, are synonyms of B floridanus Conrad, 1846. The status
of B. a. wakullae from northern Florida is problematic.
Although clearly not conspecifie with P. americana , it is ten¬
tatively assigned to Tocobaga new genus and is herein elevated
to species level. The biogeography of Tocobaga new genus is
tentatively discussed. Fossiliferous deposits in North America
and South America have not yielded taxa with the combina¬
tion of shell characters found in the new genus and relationships
with other bulimulid genera are unknown. However, it is proba¬
ble that the new genus, like other non-marine mollusks from
the Tampa Member of the Arcadia Formation, dispersed to
Florida after contact between the Caribbean Plate and the
Bahama Platform (circa 38 Ma).
Additional Keywords: Tocobaga new genus, Miocene, Florida
1 Corresponding author
INTRODUCTION
Hijperaulax Pilsbry, 1897a (type species, Bulimulus
ridleyi E. A. Smith, 1890) was described as a subgenus
of Bulimulus Leach, 1814 (Bulimulidae Tryon, 1867) to
encompass taxa having axial wrinkles on the embry¬
onic whorls and a distinct channel at the posterior inser¬
tion of the outer lip. Pilsbry (1901; 102-103) elevated
Hijperaulax to genus level with the “section” Bonnanius
Jousseaume, 1900 and transferred it to the then subfamily
Odontostominae, but did not alter the composition of
the taxon, which consisted of the extant H. ridleyi and
H. ramagei (both E.A. Smith, 1890) from Fernando
Noronha Island, Brazil and several upper Tampa Member
(Arcadia Formation) species from the Ballast Point site
in Florida. Another fossil species from the western United
States, Bulimulus limnaeiformis Meek and Hayden,
1856 was tentatively transferred to Hijperaulax by Wenz
(1923: 731), but is now assigned to the Viviparidae (see
Henderson, 1935 for references). Bonnanius Jousseaume,
1900 is currently treated as a distinct genus (Simone, 2006;
Breure and Ablett, 2012), type species B ramagei, thus
restricting Hijperaulax to its type species II ridleyi and
the Florida fossil species. The basis for the assignment
of the fossil species to Hijperaulax was the striking simi¬
larity in shell shape, size, and peristome characters
between the fossil taxa and the extant II. ridleyi and, in
particular, the presence in all taxa of a narrow channel at
the junction of the outer lip with the body whorl (Pilsbry,
1897b: 82; 1901: 102-103).
Recently, we examined numerous specimens of Ballast
Point bulimulids housed in the Invertebrate Paleon¬
tology Collection at the Florida Museum of Natural
History (FLMNH) and all type material of Florida fossil
taxa assigned to Hijperaulax. We concluded that several
unnecessary names had been introduced for the Florida
fossils. By comparing these fossil specimens to Hijperaulax
ridleyi and the type species of till other pertinent bulimulid
genera and subgenera, we find that the placement of the
K. Auffenberg et a!., 2015
Page 55
early Miocene fossils is best resolved by formally recog¬
nizing their distinctive suite of characters with a new
generic epithet. We therefore adopt the name:
SYSTEMATIC PALEONTOLOGY
Tocobaga new genus
Diagnosis: Shell ovate to ovate-cylindrical in shape.
Whorls 5.5-6; embryonic whorls approximately 1.2 -1.3,
bluntly rounded and spirally striate; later whorls sculp¬
tured with weak to moderately strong axial riblets, spiral
incised lines often present in interspaces, occasionally
crossing axial riblets; last half of body whorl slightly flat¬
tened behind peristome, ascending for last 0.2 whorl;
sutures shallow to well-impressed. Peristome broadly
expanded and thickened internally, often strongly thick¬
ened basally; inner and outer margins connected by dis¬
tinct parietal callus, interrupted by narrow channel at
posterior insertion of outer lip; marginal palatal tooth
robust, weak or lacking; when prominent, forming distinct
sinus at upper insertion of outer lip, buttressed below.
Aperture ovate to subquadrate in shape. Columella
simple, lacking lamellae. Umbilicus narrow, chink-like.
Type Species: Partula americana Heilprin, 1886.
Ballast Point, Hillsborough Bay, Hillsborough County,
Florida, upper Tampa Member, Arcadia Formation
(early Miocene).
Content: Tocobaga americanus (Heilprin, 1886),
T.jloridanus (Conrad, 1846), and T. wakullae (Mansfield,
1937).
Etymology: The name Tocobaga is derived from the
Tocobaga Tribe of Native Americans that, like these
extinct snails, inhabited the Tampa Bay region, albeit
23 million years later. Tocobaga was first used in Spanish
documents of the 1560s in reference to the male chief
of the group, the chief’s village, as well as the people
themselves. Since no gender was implied when the
name was established, we treat the genus name Tocobaga
as masculine.
Discussion: The primary shell character used to dif¬
ferentiate many genera of Qrthalieoidea Albers, I860 is
the embryonic whorl sculpture (Pilsbry, 1895, 1896). The
efficacy of this character is supported at least in part by
independent genetic evidence (Breure and Romero, 2012).
The embryonic whorl sculpture of Tocobaga americanus
is microscopically spirally striate (Figure 1) while that of
Hyperaulax ridleyi, the type species of that genus, is axi¬
ally wrinkled (Figure 2). This character alone serves to
remove the fossil species from Hyperaulax. In addition,
the first embryonic whorl of the Florida fossils is low,
rounded with a weakly impressed suture (Figure 1 ), while
that of H. ridleyi is greatly elevated, with a deeply
channeled suture (Figure 2). These features of the
embryonic whorl clearly indicate that the fossil taxa
have been erroneously assigned to Hyperaulax.
The early assignment of these species to the orthalicoid
family Buliinulidae was logical given their close resem¬
blance in gross shell moqdiology to several extant bulimulid
taxa and the biogeography of the Caribbean Basin. The
only families outside the orthalicoids that serve as rea¬
sonable alternatives to this arrangement on the basis of
shell morphology are the Partulidae and Enidae. In fact,
Heilprin (1886) described T. americanus as a species of
Partula Ferussac, 1821, and Moellendorff (1901) sug¬
gested that H. ridleyi belonged to the enid genus Napaeus
Albers, 1850. Hence, the evidence for the placement of
the fossil taxa in the Buliinulidae is examined below.
The fossil species are readily distinguished from partulids
by a variety of features. Of greatest importance are
features of the embryonic whorls. Both partulids and
Tocobaga have spirally striate embryonic whorls, but the
striae of the partulids are much coarser. The embryonic
whorls of partulids are conic and flat-sided, with barely
any relief at the suture; the embryonic whorls of Tocobaga
are rounded at the periphery, more globose, and with
more distinct sutures. The later whorls of partulids are
more rapidly expanding than those of Tocobaga and the
Figures 1-2. Embryonic whorl sculpture of Tocobaga new genus and Hyperaulax. 1. Tocobaga americanus (Heilprin, 1886).
UF 66651 (Invertebrate Paleontology Collection). 2. Hyperaulax ridleyi (E.A. Smith, 1890). UF 109915 (Malacology Collection).
Scale bar = 0.5 min.
Page 56
THE NAUTILUS, Vol. 129, No. 2
shells typically have an incomplete peristome lacking
the parietal callus and associated groove at the junction
of the outer lip with the body whorl that characterizes
Tocobaga. From these characters it can readily be seen
that the relationship of the early Miocene fossils is not
with the partulids.
Separating Tocobaga from the Enidae on the basis
of shell morphology is more difficult. Many enid taxa
have shell morphologies very' similar to that of Tocobaga
(i.e., a channel at the posterior insertion of the outer
lip and an expanded peristome). Enid embryonic whorls
may be smooth or spirally striate. The fossil genus
Dendropupa Owen, 1859 (see below), has axial sculpture
on the embryonic whorls (Solem and Yochelson, 1979).
The enid body whorl may ascend behind the peristome,
but does not become flattened near the aperture as in
Tocobaga. The primary reason for excluding Tocobaga
from assignment in the Enidae is biogeographic. The
enids are currently restricted to Europe, northern Africa,
central and southern Asia, and the Pacific from Indonesia
to Melanesia. Known fossils of this family are restricted to
the same locations with the exception of Dendropupa,
which has been collected from Upper Carboniferous
sediments in eastern Canada, France, and Poland (Solem
and Yochelson, 1979). Other enid fossils are known from
the Paleocene of Europe and later records through
the Tertiary of Europe, Africa, and the Middle East
through Central Asia to China (Zilch, 1959). Solem and
Yoehelson’s assignment indicates a possible Laurasian
origin for the Enidae and a distribution of Dendropupa
that pre-dates the separation of North America and
Europe. However, the Enidae is unknown in subsequent
North American land snail faunas and there is no other
evidence that it constituted a component of the American
M iocene fauna. Thus, we agree with Pilsbry (1901) that
the resemblance of Ht/peraulax and the Florida fos¬
sils to certain members of the Enidae is most likely
the result of convergent adaptations to dry habitats.
The thickened peristome allows for an efficient shell/
substrate seal during aestivation, while the canal acts
as a contact to the exterior.
The current geographic distribution of Orthalieoidea
including the Bulimulidae strongly suggests that bulimulids
could be expected in the fossil fauna of the Caribbean
Basin, including early insular environments on the
Florida Platform. Among Orthalieoidea families, Tocobaga
is best assigned to the Bulimulidae. It is readily sepa¬
rated from the Amphibulimidae P. Fischer, 1873 (e.g.,
Amphibulima de Montfort, 1810) on the basis of shell
shape and degree of calcification. The fossils do not
belong with the Orthalieidae Albers, 1860 (e.g., Liguus
de Montfort, 1810) because those taxa have larger,
imperforate shells and a simple lip. Tocobaga cannot
be assigned to the Odontostomidae because of the
elevated embryonic whorls of the latter, lacking in
spiral sculpture as discussed above. In addition, most
odontostomids have apertural barriers. Hence, we agree
with earlier authors that the Florida fossil species are
properly assigned to the Bulimulidae.
The distinctively striate embryonic whorls of Tocobaga
differ from all other North American and Antillean
Bulimulidae which are either axially ribbed as in Bulimulus
Leach, 1814 and Rabdotus Albers, 1850, or both axially
and spirally ribbed as in Dnjmaeu.s Albers, 1850. Dali
(1890) tentatively assigned all Florida taxa known at
that time to the South American group Anctus Martens,
1860, based on similarities such as a laterally com¬
pressed body whorl behind a broadlv reflected peri¬
stome. However, the embryonic whorls of Anctus are
smooth and lack the channeled and calloused parietal
area. Only the South American genera, Lopesianus
Weyrauch, 1958, Leiostracus Albers, 1850, Discoleus
Breure, 1978, and some Bostnjx sensu lato Troschel,
1847 possess a similarly striate embryonic whorl sculp¬
ture. Of these genera, Loj)esianus differs from Tocobaga
in having fewer and stronger spiral striae on the embry¬
onic whorls (Weyrauch, 1958: pi. 6, figs. 8); deeply
channeled, crenulate sutures; no parietal callus; and
an incomplete, simple peristome. Leiostracus differs in
having the spiral striae confined to the lower half of
each whorl; papilliform embryonic whorls; and an
incomplete peristome only slightly expanded. Discoleus
differs in having a larger, more bulbous embryonic
whorl; a subovate aperture; more convex whorls; and a
simple, incomplete peristome.
The polyphyletic genus Bostnjx sensu lato (Breure
and Romero, 2012) is the most difficult genus from
which to distinguish Tocobaga , a reflection of the pre¬
sumably diverse nature of the former. In each of several
characters, one or a few species of Bostnjx sensu lato
can be found that approach the state seen in Tocobaga.
However, in no case does any species of Bostnjx sensu
lato approach the entire suite of characters that serve
to distinguish the fossils from other Bulimulidae.
We examined the type species of all but two ( Elatibostnjx
Weyrauch, 1958 and Kionoptjx Haas, 1966) of the
21 generic synonyms (or subgenera) given by Breure
(1979) as belonging to Bostryx sensu lato to determine
whether any of these names could apply to the early
Miocene fossils. All but Peronaeus Albers, 1850, Plati/hostn/x
Pilsbry, 1896, Phenacotaxus Dali, 1912, Scansicohlea
Pilsbry, 1930, and Pampasinus Weyrauch, 1958 can be
dismissed for having embryonic whorls that are entirely
smooth or having a combination of spiral and axial
sculpture (the type of Bostnjx sensu stricto, B. solutus
(Troschel, 1847) has smooth embryonic whorls). Elatibostnjx
and Kionoptyx are disregarded because of their dis¬
similar embryonic whorl sculpture and/or shell mor¬
phology mentioned in the original descriptions (Haas,
1966: 239; Weyrauch, 1958: 113). Peronaeus , Pampasinus,
and Platybostnjx can be eliminated on the basis of shell
shape. The first is greatly elongated, and the other two
genera are thick and lens-shaped. Phenacotaxus, Scansicohlea,
and Tocobaga have similar shell shapes, but do not agree
in any other of the characters we use to distinguish the
latter. Phenacotaxus and Scansicohlea have simple peri¬
stomes, their body whorls do not ascend near the aper¬
ture and are evenly expanded, not flattened behind the
K. Auff'enberg et a!., 2015
Page 57
Table 1. Shell morphometries of type specimens of nominate forms of Tocobaga. L = shell length, W = shell width, ApL =
aperture length, ApW = aperture width; all measurements in mm, X = no measurement possible. Bold species are considered valid.
peristome. It is clear that the moqdiology of Tocobaga is
unique, that this taxon has no close resemblance to any of
the name-bearing species currently synonomized with
Bo.stryx sensu lato, and that consequently, none of these
names can serve to accommodate the Florida fossils.
Examination of the holotypes of all named fossil
“ Hyperaulax ” indicates that only three species are recog¬
nizable in the Florida fossils; the taxa described by Dali
(1890, 1915) are synonyms of either T. americanus or
T. floridanus. Below we diagnose and discuss each spe¬
cies. Shell length, shell width, aperture length, and aper¬
ture width (Tables 1-2) were measured as described
in Crampton (1916) for similarly shaped Partula. We
use the following institutional abbreviations: ANSP
Table 2. Tocobaga americanus (Heilprin, 1886) shell mor¬
phometries; L = shell length, W = shell width, ApL —
aperture length, ApW = aperture width; number of whorls of all
measured fossils estimated at 6+ due to erosion of embryonic
whorls. All measurements in mm.
(Academy of Natural Sciences of Drexel University),
UF (Florida Museum of Natural History, University of
Florida), USNM (National Museum of Natural History,
Smithsonian Institution), anti WFIS (Wagner Free Insti¬
tute of Science, Philadelphia).
SYSTEMATICS
Tocobaga americanus (Heilprin, 1886) new
combination
(Figures 3-7, Tables 1 and 2)
Partula americana Heilprin, 1886: 115, pi. 16, fig. 60.
Bulimulus (? Anctus) americanus var. partulinus Dali,
1890: 7; 1915: 26, pi. 4, fig. 12.
Bulimulus (? Anctus) americanus var. laxus Dali, 1890: 7;
1915: 26, pi. 4, fig. 14.
Bulimulus ( Hyperaulax ) patulinus Mansfield, 1937: 24-25
(lapsus calami )
Diagnosis: A large species of Tocobaga , 15-17 mm in
height, 6-8 mm in width; shell ovate in shape; whorls
approximately 5. 5-6. 5, slightly convex; shell sculpture of
evenly spaced oblique riblets of width equal to their inter¬
spaces; last half of body whorl slightly flattened, ascending
for last 0.2 whorl; aperture subquadrate; peristome with
a wide, flat expansion, thickened internally throughout,
almost complete, inner and outer margins connected by a
thickened parietal callus; a narrow channel present at pos¬
terior insertion of outer lip, parallel to the long shell axis;
umbilicus narrow; columella simple and straight.
Holotype: WFIS 865 (Figures 3-5) Ballast Point,
Hillsboro [Hillsborough] Bay, [T;unpa, Hillsborough County],
Florida, upper Tampa Member, Arcadia Formation (early
Miocene). J. Willeox and A. Heilprin, 1886.
Remarks: Tocobaga americanus is readily distinguished
from T. floridanus by its larger size and more ovate
shape; more rapidly expanding whorls; peristome that
is expanded along the entire outer margin; channel at
Page 58
THE NAUTILUS, Vol. 129, No. 2
Figures 3-7. Tocobaga americanus (Heilprin, 1886). 3-5. Holotype of Partula americana Heilprin, 1886. WFIS 865. 6. Holotype
of Bulimulus americanus var. laxus Dali, 1890. USNM 111971. 7. Holotype of Bulimulus americanus var. partulinus Dali, 1890.
USNM 119970. Scale bar = 5 mm.
the posterior insertion of the outer lip that is parallel to
the main axis of the shell; and a more open umbilicus.
The holotypes of Dali’s (1890) varieties laxus (Figure 6)
and partulinus (Figure 7) are indistinguishable from
typical T. americanus in any important detail. The shells
of laxus are more slender and have stronger axial sculp¬
ture, while those of partulinus are even more slender.
However, both are also shorter in length than typical
T. americanus and all three have identical length to
width ratios. Similarly, tire length to width ratios of the
aperture is also alike (Table 1). Variation among 16 unbro¬
ken individuals of 59 specimens in UF lots 66651, 66652,
and 66653 (Table 2) include a morphological range in
shell length and width and apertural length and width
that encompasses the two varietal names and we see
no need to recognize this natural variation with formal
epithets. Specimens from Wakulla County, Florida ques¬
tionably referred to these forms (Mansfield, 1937) were
not located.
Material Examined: Partula americana Heilprin, 1886,
Holotype, WFIS 865; Tocobaga americanus (Heilprin,
1886), UF 66651 (1), UF 66653 (57), USNM 111965 (1);
Bulimulus americanus var. partulinus Dali, 1890, Holo¬
type, USNM 111970; Bulimulus americanus var. laxus
Dali, 1890, Holotype, USNM 111971.
K. Auffenberg et al., 2015
Page 59
Tocobaga floridanus (Conrad, 1846) new combination
(Figures 8-16, Table 1)
Bulimus floridanus Conrad, 1846: 399, text-figure.
Bulimus ( Bulimulus ) longaevus Ancey, 1881: 414.
Bulimulus (? Anctus) heilprinianus Dali, 1890: 6-7, pi. 1,
figs. 6b, 10.
Bulimulus (? Anctus) steamsii Dali, 1890: 7-8, pi. 1, fig. 12.
Bulimulus ( Hi/peraulax ) ballistae Dali, 1915: 26-27, pi. 1,
fig. 5.
Bulimulus (Hi/peraulax) remolina Dali, 1915: 27-28,
pi. 1, fig. 18.
Bulimulus ( Hi/peraulax ) tampae Dali, 1915: 26, pi. 1, fig. 3.
Bulimulus (Hi/peraulax) tortilla Dali, 1915: 27, pi. 1, fig. 2.
Diagnosis: A smaller species of Tocobaga, 8.1-14.1 mm
in height, 3. 6-6.0 mm in width; shell ovate to ovate
cylindrical in shape; whorls 5-6.5, slightly convex to
flattened; embryonic whorl blunt, rounded, sculpture
unknown; shell sculpture of evenly spaced oblique riblets
of width equal to their interspaces, weak spiral incised
lines occasionally present in interspaces; body whorl
not evenly expanded, last half whorl slightly flattened,
ascending last 0.2 whorl; aperture oval to subquadrate;
peristome expanded, thickened on inner margin along
the columellar area, basally and along anterior half
of outer lip; inner and outer peristome connected by a
parietal callus; narrow channel present at posterior inser¬
tion of outer lip that is deflected medially at 45° rela¬
tive to shell long axis; umbilicus very narrow, chink-like;
columella simple and straight.
Holotype: ANSP 30607 (Figures 8-10). Nine miles from
Tampa on Hillsboro [Hillsborough] River, [Hillsborough
County], Florida, upper Tampa Member, Arcadia Forma¬
tion (early Miocene). The holotype is a broken speci¬
men. The inadequate original text-figure apparently depicts
a reconstruction.
Remarks: Bulimulus longaevus Ancey, 1881 is an unnec¬
essary replacement name for Bulimus floridanus Conrad,
1846 (see Henderson, 1935), not B floridanus Pfeiffer,
Figures 8-16. Tocobaga floridanus (Conrad, 1846). 8-10. Holotype of Bulimus floridanus Conrad, 1846. ANSP 30607. 11. Holotype
of Bulimulus ballistae Dali, 1915. USNM 165013. 12. Holotype of Bulimulus heilprinianus Dali, 1890. USNM 111962. 13. Holotype
of Bulimulus remolina Dali, 1915. USNM 165014. 14. Holotype of Bulimulus steamsii Dali, 1890. USNM 111964. 15. Holotype of
Bulimulus tampae Dali, 1915. USNM 165012. 16. Holotype of Bulimulus tortilla Dali, 1915. USNM 165015. Scale bar = 5 mm.
Page 60
THE NAUTILUS, Vol. 129, No. 2
1857, as demonstrated by Wood and Gallichan (2008: 60).
Ancey (1881: 414) erroneously stated that B. floridanus
Pfeiffer, 1857 was described prior to Conrad’s taxon
(1846), leading Wood and Gallichan (2008) to mis¬
interpret Ancey’s intent. The holotypes of Dali's spe¬
cies Bulimulus ballistae , B. heilprinianus , B. remolina,
B. steamsii , B. tampae, and B. tortilla while differing
in size are all similar in shape (Figures 8-16) and shell
length to width and aperture length to width ratios
both to each other and to the holotype of Bulimus
floridanus (Table 1). The holotype of Bulimulus ballistae
(Figure 1 1) is nothing more than a rather small specimen
of Tocobaga floridanus differing from that species in no
obvious detail. The holotypes of Bulimulus heilprinianus
(Figure 12) and Bulimulus tortilla (Figure 16) differ from
the typical T. floridanus only in having a more pro¬
nounced thickening on the anterior half of the inner
margin of the outer lip. The holotype of B. floridanus
is not fully mature and so does not prominently exhibit
this character. The holotype of Bulimulus tampae
(Figure 15) is merely a large specimen of T. floridanus
and differs in no characters other than its size. The holo¬
type of Bulimulus remolina (Figure 13) is slightly more
elongate than typical T. floridanus and has a rather flat¬
tened body whorl with the sculpture largely eroded. The
holotype of Bulimulus steamsii (Figure 14) is the most
unusual specimen examined. This specimen is quite elon¬
gate and large; with narrow, flattened whorls; almost
lacking in shell sculpture. We believe the smooth sur¬
face is due to erosion, because traces of the charac¬
teristic ribbing of Tocobaga remain on a few surfaces,
most prominently behind the aperture. The specimen’s
large size and elongate shape appear to represent one
extreme of the variation in T. floridanus. The holotype
of B. remolina and other specimens (USNM 646161)
are intermediate in shape between the holotype of
B. floridanus and that of B. steamsii, or are equally as
flattened as the type of B. steamsii , although not as large.
The extreme elongation in the holotype of B. steamsii
is partly, but not entirely, a reflection of its large size,
though other large specimens of T. floridanus (e.g.,
USNM 646160) are more ovate in shape. Given the
continuum in size and shape that occurs between the
holotypes of B. floridanus and B. steamsii, and the appar¬
ently eroded nature of the latter’s smooth surface, we
believe tire holotype of B. steamsii represents one extreme
of the morphological variation seen within T. floridanus .
Material Examined: Bulimus floridanus Conrad, 1846,
holotype, ANSP 30607; Tocobaga floridanus (Conrad, 1846),
USNM 111960 (1), USNM 111961 (7) USNM 646160
(5), USNM 646161 (5), USNM 646162 (1); Bulimulus
heilprinianus Dali, 1890, holotype, USNM 111962;
Bulimulus steamsii Dali, 1890, holotype, USNM 111964;
Bulimulus tampae Dali, 1915, USNM 165012; Bulimidus
ballistae Dali, 1915, holotype, USNM 165013; Bulimidus
remolina Dali, 1915, holotype, USNM 165014; Bulimulus
tortilla Dali, 1915, holotype, USNM 165015.
Tocobaga wakullae (Mansfield, 1937) new combination
(Figures 17-19, Table 1)
Bulimulus americanus ivakullae Mansfield, 1937: 15:70,
pi. 1, figs. 10, 13.
Diagnosis: A large species of Tocobaga, ca. 26 mm in
height, >10 mm in width; shell ovate in shape; whorls
approximately 6.5, slightly convex; embryonic whorl blunt,
rounded, sculpture unknown; later whorl sculpture of
Figures 17-19. Tocobaga wakullae (Mansfield, 1937). Holotype of Bulimulus americanus wakullae Mansfield, 1937. USNM
495932. Scale bar = 5 mm.
K. Auffenberg et a!., 2015
Page 61
evenly spaced axial riblets more or less parallel to shell
long axis, spiral sculpture not apparent; body whorl
very slightly flattened behind apertural lip, ascending
very slightly for last 0.2 whorl; aperture ovate, slightly
oblique; peristome moderately reflected, but not greatly
expanded or thickened within; parietal area with weak
callus; channel at posterior insertion of outer lip not
apparent; characters of umbilical area unknown; colu¬
mella simple, unadorned.
Holotype: USNM 495932 (Figures 17-19). About
200 yards south of Wakulla Railroad Station, Wakulla
County, Florida. External mold and cast.
Remarks: Mansfield (1937) originally placed this taxon
as a subspecies of Tocobaga americanus , but it differs
from that species in its larger shell size, in having an
ovate instead of subquadrate aperture, in its more inflated
shape, and in having the axial shell sculpture oriented
more nearly parallel to the long axis of the shell. These
differences in shell characters lead us to conclude that
these two taxa are separate species.
Our assignment of this taxon in Tocobaga , however,
is not without reservation. Several of the diagnostic fea¬
tures of that genus, such as the embryonic whorl sculp¬
ture, the channel at the posterior insertion of the outer
lip, and the nearly complete peristome, cannot be exam¬
ined in the holotype of B. wakullae (the only specimen
available to us) because of its poor preservation (an
external mold) and casting material. Collection of addi¬
tional material may allow assessment of these particular
characters, whereby the generic allocation of B. wakullae
may well be revised.
Tocobaga wakullae (Mansfield, 1937) was described
from the “Tampa Limestone” of Wakulla County in
northwestern Florida. Although Mansfield (1937) did
not discuss the stratigraphy of the type locality, the type
specimen was probably collected from what is now con¬
sidered the St. M arks Formation. This formation is early
Miocene in age (Rupert and Spencer, 1988) and approxi¬
mately contemporaneous with the upper Tampa Member
(Arcadia Formation) of the Tampa Bay area.
Material Examined: Bulimulus americanus wakullae
Mansfield, 1937, Holotype, USNM 495932, external mold
and cast.
BIOGEOGRAPHY
Modern orthalicoids are particularly diverse in South
America with a few lineages in Africa, Australia, Melanesia,
and New Zealand. The group appears to have a Gondwanan
distribution (Herbert and Mitchell, 2009). However,
the family is well represented in Mexico, Central America,
and the Antilles and there are representatives of a few
genera in temperate North America including the bulimulids
Drymaeus and Rabdotus. Several other Antillean land
snail groups were widely distributed in North America
during the late Cretaceous and early Tertiary ( Bishop, 1979;
Roth and Hartmann, 1998) and became restricted to the
Antilles as climate cooled in the Tertiary. However, the
oldest North American fossil bulimulids are the Florida
Miocene fossils reviewed here and there is yet no evidence
that bulimulids were present in North America before
then. The oldest bulimulids are from a middle Paleocene
site in southern Brazil where the family accounts for
more than 30% of fossil species diversity (Salvador and
Simone, 2013). The family remains the most diverse
family in South America accounting for approximately
45% of recent South American species (Simone, 2006).
M any of the Florida Tampa Member (Arcadia For¬
mation) land snails are similar in shell morphology to
extant Antillean species (Pilsbry, 1897a; Auffenberg and
Portell, 1990). Fossil species were assigned to the extant
Antillean genera Plagioptycha Pfeiffer, 1855, Cepolis de
Montfort, 1810, and Pleurodonte Fischer de Waldheim,
1807, Gongylostoma Albers, 1850, and Cerion Roding,
1798 (Dali, 1915; Mansfield, 1837). While the placement
of these fossil taxa in modern genera can be questioned,
the fauna surely has a degree of Caribbean affinity. The
Antillean component of the Tampa Member (24-22.5 Ma,
but see Scott, 1988) species apparently dispersed over
water to Florida after contact between the Caribbean
Plate and the Bahama Platform 38 Ma (Duncan and
Hargraves, 1984). Reexamination of the entire fauna
may provide a better understanding of its biogeography.
Relationships between Tocobaga and recent bulimulid
genera remain unknown. Fossiliferous deposits in North
America have not yielded taxa closely similar to Tocobaga
and the extinct South American fossil genera, Paleobulimulus
Parodiz, 1949 and Itaborahia Maury, 1935, have shell
morphologies quite unlike Tocobaga (Parodiz, 1969).
AC K N OWLE D G M E NTS
For donation of fossil specimens to the FLMNH that
were used in this study, we thank S.B. Upchurch (SDII-
Global), T. Estevez, and the Florida Geological Survey.
For specimen loans, we thank J. Thompson and the late
W. Blow (USNM), E. Bolt (WFIS), and G. Rosenberg
(ANSP). Much research during an earlier version of this
study was performed by Fred Kraus (Bishop Museum).
Information on the Tocobaga Tribe was provided by
]. Milanich (FLMNH). Pertinent literature citations and
contemplation was offered by H. G. Lee (FLMNH). For
sharing his vast knowledge of the Orthalicoidea, A. S.H.
Breure (Leiden, The Netherlands) is thanked, as is F.G.
Thompson (FLMNH) for reviewing an earlier draft of
this manuscript. Digital images of the ANSP and WFIS
types were provided by P. Callomon and A. Lawless.
Reviews by A. S.H. Breure and Barry Roth provided
helpful suggestions that improved this manuscript. This is
University of Florida Contribution to Paleobiology 669.
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THE NAUTILUS 129(2):63-70, 2015
Page 63
First report of the Eocene gastropod Mitrella ( Bastropia )
(Neogastropoda: Columbellidae) from the northeast Pacific
and paleobiogeographic implications
Richard L. Squires
Department of Geological] Sciences
California State University
Northridge, CA 91330-8266 USA
and
Invertebrate Paleontology1
Natural History Museum of Los Angeles Countv
Los Angeles, CA 90007 USA
[email protected]
ABSTRACT
The columbellid neogastropod Mitrella ( Bastropia ) llajasensis
new species is described from a lower Eocene siltstone bed in
the shallow-marine part of the Llajas Formation, Ventura
County, southern California. This new species, which lived in
a warm-water environment, is the first record of Bastropia
Palmer, 1937 in the northeast Pacific region. Bastropia differs
from Mitrella Risso, 1826 by having axial riblets on an early
part of the spire and a longer and more constricted anterior
canal. In addition, the majority of species of Bastropia have
ineised-spiral line(s), with or without pits, near the suture on
the posterior part of the lower spire whorls and on the last
whorl. The geologic range of Bastropia is late Paleocene?
(Thanetian?) and early Eocene (Ypresian) to late middle Eocene
(Bartonian). Its previous occurrences are in Alabama (oldest
occurrence), Texas, and South Island, New Zealand (youngest
occurrence). Bastropia most likely had planktotrophic larval
development, and the introduction of Bastropia into California
from the Gulf Coast region was closely associated with the
“Early Eocene Climatic Optimum” (EECO), a time of global
warmth and high sea level.
Additional Keywords: Simi Valley, thermophilic
INTRODUCTION
Neogastropods first became abundant during the Early
Cretaceous (late Albian) (Kiel, 2002) and subsequently
became increasingly diversified. One of the post-Cretaceous
neogastropod groups is family Columbellidae Swainson,
1840. According to Schnetler (2001), the earliest known
Golumbellids are of Paleocene age, belong to Astyris
1 Research Associate
H. and A. Adams, 1853, and are found in northern
Europe and Greenland. The global Paleogene (Paleo¬
cene to Oligoeene) record of columbellids is meager.
In the northeast Pacific, in a region extending from
southwestern Washington to southern California, their
record is extremely rare. The only Paleogene columbellid
reported previously from this particular region is Mitrella
(Mitrella) blackhillsensis (Squires and Goedert, 1994), of
middle early Eocene age from the Crescent Formation
in southwestern Washington. This present study con¬
cerns the columbellid Mitrella (Bastropia) llajasensis
new species of late early Eocene age from the Llajas
Formation in southern California (Figure 1). Bastropia
Palmer, 1937 is known previously from Eocene strata
in Alabama, Texas, and South Island, New Zealand
(Table 1).
Columbellids underwent great diversification during
the Miocene to Holocene. For example, a great array of
columbellids is found in middle Miocene deposits of
the Caribbean region (Woodring, 1964: 246). Today,
columbellids comprise a highly diverse group with about
400 extant species. Their shells are small, normally
between 3 and 20 mm in height. Most columbellids
are active epibenthic carnivores or scavengers, but
some feed on algae. Herbivory is uncommon among
neogastropods, and only the columbellids and nassariine
bueeinids include herbivorous species. Columbellids
today have worldwide distribution, mainly in tropical
and warm-temperate seas, but a few species inhabit
high-latitude and deep-ocean environments (deMaintenon,
1999: 2008). Columbellids have larval-developmental strat¬
egies ranging from embryos hatching into planktotrophic
veliger larvae (indirect development) to embryos hatch¬
ing into small crawling snails (non-pelagic development)
(Bandel, 1974). Columbellids can have determinate
growth, thus, at adulthood there is thickening of the
outer lip, with concomitant development of denticles
Page 64
THE NAUTILUS, Vol. 129, No. 2
Figure I. Index map of LACMIP locality 41604, the type
locality of the new species.
on the interior surface of the outer lip (deMaintenon,
2005: 501; Fernet, 2007).
MATERIALS AND METHODS
The new species is based on a well-preserved single
specimen. Its small size (8 mm height) is most likely
why this species has been previously overlooked. Shells
of this size are commonly firmly cemented in rock matrix
in the Llajas Formation, and, as a result, they are easily
destroyed during the process of collecting. In the case of
the new species, however, it was found in relatively soft
matrix, which was easily removed by picking at the grains
with a very sharp, small needle. The lower and middle
Eocene provincial molluscan “stages” mentioned in the
text are informal (in quotation marks) and, as discussed
by Squires (2003), applv mainly to California. Morpho¬
logic terms are from Cox (1960). Supergeneric classifica¬
tion is based on Bouchet and Rocroi (2005: 254).
STRATIGRAPHY AND DEPOSITIONAL
ENVIRONMENT
Squires (1984, 2001) provided details of the stratigraphy,
depositional environments, and paleontology of the
Llajas Formation. The locality where the new species
was found is in siltstone within the middle lower part
of the “shallow-marine (transgressive) deposits” in the
formation. This part of the formation is of late early
Eocene age. Co-occurring shallow-marine, warm-water
species are the large benthic foraminifer Pseudophragmina
( Proporocyclina ) clarki (Cushman, 1920) and the
goneplacid crab Glyphithyreus iveaveri (Rathbun, 1926).
SYSTEMATIC PALEONTOLOGY
Class Gastropoda Cuvier, 1797
Clade Neogastropoda
Superfamily Buccinoidea Rafinesque, 1815
Family Columbellidae Swainson, 1840
Discussion: DeMaintenon (1999, 2008) reported that
columbellids have retained the basic neogastropod gut
anatomy and that the herbivory observed in columbellids
is probably a derived condition. The classification of
columbellids is presently not well resolved, with many
taxa based on morphologic groups rather than on
monophyietic elades (deMaintenon, 1999). Results of
deMaintenon s (1999: 264) parismony-based phylogenetic
studies support a close relationship between columbellids
and buccinids.
The supraspecific assignment of Paleogene columbellids
has been highly unstable. The historical trend in the
literature has been the usage of names belonging to
extant genera. As noted by deMaintenon (2008: 267),
many extant and fossil species currently are placed in
genus Mitrella Risso, 1826. DeMaintenon (1999, 2008)
reported, however, that Mitrella is not a monophyietic
group, but it is a polyphyletic collective for biconic
columbellids with smooth, unsculptured shells.
Genus Mitrella Risso, 1826
Type Species: Mitrella scripta (Linnaeus, 1758)
[— Mitrella flaminea Risso, 1826], by subsequent desig¬
nation (see Woodring, 1928: 273); extant, Mediterranean
Sea, Portugal, and Morocco (Poppe and Goto, 1991: 152).
Description: Shell small (approximately between 4 and
18 mm height), fusiform. Spire moderately high to high,
apex subrounded to aeieular-Iike. Protoconch low-domal,
paucispiral, smooth. Teleoconch five to eight whorls.
Whorls convex to flat-sided. Suture impressed to indis¬
tinct. Shell generally smooth, except for pillar (neck) area.
Shell can be glossy. Axial sculpture normally absent.
Those in modem taxa having rare exceptions of axial rib-
lets on upper spire (see below) need modern systematic
work and might represent different genera or subgenera.
R.L. Squires, 2015
Page 65
Spiral sculpture prominent on pillar, elsewhere, minor
or absent. Aperture on most species wide, short, and
truncate. Columella generally straight and smooth with
slight callus. Pillar commonly short. Interior of outer lip
can have denticles. Area preceding outer lip can have
“varix”-like swollen area.
Discussion: Keen (1971) reported two species of
Mitrella that have axial riblets on the earliest teleoconch
whorls. Botii species are from the southern tip of Baja
California, Mexico. They are Mitrella haccata (Gaskoin,
1852) (also see Keen, 1971: fig 1231) and Mitrella
caulerpae Keen (1971: fig. 1232, two views). Mitrella
haccata has axial ribs on the first two early teleoconch
whorls and has a very short anterior canal. Mitrella
caulerpae has axial ribs on only the first teleoconch
whorl, a moderately short anterior canal, and either a
thick outer lip or one with a varix.
Most nominal Mitrella , like the extant Mitrella pallaryi
(Dautzenberg, 1927) in the eastern Atlantic, have been
reported as having planktotrophic development, inferred
from the multispiral protoconch of this species (see World
Register of Marine Species at http://www.marinespecies
.org). By similar reasoning, Harzhauser and Kowalke
(2002) inferred that a Miocene species of Mitrella had
planktotrophic development. A few extant species of
Mitrella have been observed as having nonpelagic devel¬
opment (Chaney and deMaintenon, 2009).
Subgenus Bastropia Palmer, 1937
Type Species: Astt/ris hastropensis Harris, 1895, by
original designation, middle Eocene, Texas.
Description: Shell small (up to 20 mm height), slender
fusiform. Spire high. Pleural angle 25° to 40°. Protoconch
three smooth whorls. Teleoconch 5 to 6.5 whorls. First
one to two teleoconch whorls with many and closely
spaced axial riblets. Other spire whorls smooth, but pos¬
terior part of penultimate and last whorls can have
incised spiral line(s), with or without many closely spaced
pits. Columella straight and with slight callus. Aperture
narrow and long. Anterior canal well developed and con¬
stricted. Pillar elongate and covered by many spiral ribs;
those extending onto medial area of columella can create
“plicate” appearance. Aperture narrow and long. Anterior
canal moderately long to long, anterior canal constricted,
siphonal fasciole can be distinct.
Discussion: The type species of Bastropia differs from
the type species of Mitrella by having a narrower last
whorl, axial riblets on the first and/or second teleoconch
whorls, incised spiral line(s) with or without pits com¬
monly present on posterior parts of the lower spire
whorls and on the last whorl, spiral ribs present on base
of the last whorl, absence of thick callus on the inner and
parietal lips, thinner outer lip, and an aperture that is
more distinctly constricted anteriorly. Whether or not
there are denticles on the interior of the outer lip of
Bastropia is unknown because all the specimens of the
all the species of Bastropia have the margin of the outer
lip broken. All these specimens might be juveniles.
Palmer (1937: 282) stated that the height of the spire
of Bastropia is greater than the height of its last whorl,
but, based on measurements of the four known species
of Bastropia listed in Table 1, this statement is incorrect.
The height of the spire is actually less (in most species,
slightly less) than the height of the last whorl. Modern
workers have commonly identified (e.g.. Palmer, 1937)
or compared (e.g., MacNeil and Dockery, 1984) these
four species to Mitrella ( Bastropia ). Maxwell (1992: 130,
fig. 7e) recognized Bastropia as a distinct genus, and
future work might confirm his assertion. Each species of
Bastropia is known only from a single specimen or just a
few specimens, and all specimens are from siliciclastic
sandstone (can be glauconitic) or siltstone.
Bastropia llajasemis new species
(Figures 2-6)
Diagnosis: Bastropia with a relatively low spire and
wide pleural angle (40°), axial riblets on both second
and third teleoconch whorls, incised spiral line(s) with or
without pits commonly present near lower suture of lower
spire whorls and near suture of last whorl, somewhat
inflated last whorl, distinct siphonal fasciole.
Description: Shell small (8 mm height, 3 mm diameter),
fusiform elongate. Entire shell comprised of nine whorls.
Pleural angle 40°. Protoconch three whorls, overall nar¬
rowly conical, smooth; apical whorl low and domal with
other two whorls slighly convex. Teleoconch six whorls,
rather flat-sided except for more convex last whorl.
Suture indented, imparting tabulate look to lower spire
whorls. Axial sculpture confined to first and second
teleoconch whorls (i.e., fourth and fifth whorls of entire
shell) and consisting of approximately 18 narrow, closely
spaced opisthoeyrt riblets extending from suture to
suture and general alignment on both sides of suture.
Spiral sculpture consisting of line(s) and ribs: Incised-
spiral line(s) somewhat irregularly spaced, bearing many
closely spaced pits between slightly raised growth lines,
and confined to posterior part of penultimate and last
whorls (three spiral lines on penultimate whorl and five
on last whorl); spiral ribs confined to pillar (base of last
whorl). Aperture narrow and long, widest posteriorly and
with distinct constriction anteriorly. Outer lip missing.
Inner lip with thin callus. Pillar long and with approxi¬
mately 12 prominent spiral ribs; those on central part of
pillar have “plicate appearance” underneath callus and
onto inner lip but not extending very far into aperture.
Pillar with small but distinct siphonal notch. Siphonal
canal not bent. Growth line on spire whorls prosocline;
growth line on last whorl parasigmoidal near suture but
prosocline on pillar; intersection of growth lines and
spiral ribs on pillar near columella producing weak
eancellate shell-surface structure.
Holotype: LACMIP (Natural History Museum of Los
Angeles County, Invertebrate Paleontology' Section) 14514.
Page 66
THE NAUTILUS, Vol. 129, No. 2
Figures 2-6. Mitrella ( Bastropia ) llajasensis new species.
Holotype LACMIP 14514, LACMIP loc. 41604, height 8 mm.
2. Apertural view. 3. Apertural view turned slightly left.
4. Closeup of tip (height 1 mm) shown in previous figure.
5. Left-lateral view. 6. Abapertural view. Scale bars = 1 mm.
Type Locality: LACMIP 41604, in greenish-brown
siltstone, on north side of paved road leading to a large
water tank on east side of mouth of Runkle Canyon, near
intersection with Sequoia Street, Santa Susan a Quadrangle,
7.5-minute, photorevised 1969, south side of Simi Valley,
Ventura County, southern California (Figure 1). Collector:
R.L. Squires, March 27, 1999.
Geologic Age: Late early Eocene.
Distribution: Upper Ypresian Stage (equivalent to
provincial lower “Domengine Stage”) part of the Llajas
Formation, lower middle part within the “transgressive
shallow marine” deposits, southern Simi Valley, Ventura
County, southern California.
Etymology: Named for the Llajas Formation.
Discussion: A single specimen was found. It shows
excellent preservation, except that it has undergone some
post-depositional crushing. There are cracks in the shell
(see Figure 5), and its outer lip is broken off. It is unfor¬
tunate that the interior moqahology of the outer lip of
the specimen cannot be determined, thus it cannot be
determined if it represents a juvenile or an adult.
The new species differs from Bastropia bastropensis
(Harris, 1895: 74, pi. 8, fig. 2; Palmer, 1937: 283, pi. 37,
figs. 1, 7), from the lower middle Eocene Weehes Forma¬
tion in Bastrop County, Texas and also questionably from
middle Eocene strata in Mississippi (Palmer and Brann,
1966: 774), by having a smaller shell, wader pleural angle,
lower spire, no incised-spiral line anterior to the suture,
incised spiral lines with pits near the suture between the
penultimate and last whorls, last whorl more swollen,
spiral ribs on pillar not as high up on the shell, central
part of pillar with a “plicate appearance,” and the pres¬
ence of a siphonal fascicle.
The new species differs from Bastropia mackaiji
(Suter, 1917: 43^44, pi. 5, fig. 12; Maxwell, 1992: 130,
fig. 7e), from South Island, New Zealand, by having a
smaller shell, wider pleural angle, lower spire, no incised
spiral line on ante-penultimate whorl, additional incised
spiral lines (but with pits) on last whorl and penultimate
whorls, fewer spiral ribs on pillar, shorter posterior part
of aperture, more inflated last whorl, and the presence of a
siphonal fasciole. According to Maxwell (1992), B. mackaiji
occurs in either upper middle Eocene (upper Lutetian)
or upper middle Eocene (Bartonian) strata, thus making
C. (B.) mackaiji the youngest known Bastropia.
The new species differs from Bastropia subfraxa
(Harris, 1899: 58-59, pi. 7, fig. 11), from the lower
Eocene (Ypresian) Bashi Member of the Hatchetigbee
Formation Member in Alabama (Palmer and Brann,
1966: 512 [as Astyris subfraxa]), by having a smaller
shell, wider pleural angle, lower spire, presence of spiral
sculpture on the posterior part of penultimate and last
whorls, and a more inflated last whorl. A questionable
occurrence of B. subfraxa is from the Tuscahoma For¬
mation, Landing Marl Member, in Alabama (Toulmin,
1977: table 2). According to Dockery (1986: fig. 1), this
member is near the late Paleoeene (Thanetian)/early
Eocene (Ypresian) boundary.
The presence of axial riblets on the upper spire of
Mitrella ( Bastropia ) llajasensis new species resembles
those on the incomplete paratype of the early Eocene
columbeilid that Garvie (1996) identified as Mitrella
( Clinurella ) nuttalli (Garvie, 1996: 68-69, pi. 14, fig. 17)
from the lower Eocene Marquez Member of the Reklaw
Formation in Texas. The new species differs from
R.L. Squires, 2015
Page 67
Game’s paratvpe, which consists of only the upper spire,
by not having near the suture one or more relatively wide
spiral ribs with prominent grooves in the interspaces. On
the holotype (an adult specimen) of M. (C.) nuttalli (see
Garvie, 1996: pi. 14, figs. 15-26), whose uppermost spire
whorls have been eroded, these spiral ribs and grooves
become even more prominent on the penultimate and
last whorls. Bastropia llajasensis new species differs also
from the holotype of M. (C. ) nuttalli by having a
narrower shell, incised-spiral lines anterior to the suture,
pits on these spiral lines, absence of prominent spiral
ribs with sunken (flat-bottomed) interspaces on the
Figure 7. Global paleobiogeographic distribution and associated ranges of Bastropia (plotted in order of first-appearance datum);
data derived from Table 1. Geologic time scale, stage ages, and timing of global-climatic events from Gradstein et al. (2012).
Land-mass positions from Smith et al. (1994). Numbers at top of columns refer to geographic regions and primary sources of data:
1, Harris (1899), Palmer (1937), and Toulmin (1977). 2, present study. 3, Harris (1895). 4, Maxwell (1992).
Page 68
THE NAUTILUS, Vol. 129, No. 2
anterior half of the last whorl, a more elongate pillar,
narrower posterior part of aperture, and a more con¬
stricted siphonal canal. Although the outer lip is missing
in the new species, the holotype of M. (C.) nuttalli has
an outer lip interior with denticles. The holotype of
M. ( Clinurella ) nuttalli is much more similar to the type
specimen of the columbellid identified by Cossmann
(1886) as Mitrella ( Columbellopsis ) biarata (Cossmann,
1886: 232-234, pi. 10, fig. 3, two views), from middle
Eocene (Lutetian) strata in France. In addition, as noted
by Garvie (1996: 68), M. ( Clinurella ) nuttalli is similar to
Mitrella ( Clinurella ) buccinifonnis (Heilprin, 1879: 213,
pi. 13, fig. 7) and Mitrella ( Columbellopsis ) mississippiensis
(Meyer and Aldrich, 1886: 43, pi. 38, figs. 16, 17, 21, 22).
The new species differs from the earlier mentioned
Baja California species Mitrella bacata and M. caulerpae,
which have axial riblets on the first one or two teleoconch
whorls, in the following ways: Elongate-fusiform shell
with a long anterior canal that is constricted anteriorly,
indented sutures, subsutural-spiral sculpture consisting
of pitted, incised-spiral lines on the penultimate and
last whorls.
The only other known Paleogene columbellid from
the northeast Pacific region is Mitrella ( Mitrella )
blackhillsensis Squires and Goedert (1994: 261, figs. 21-
23), based on two specimens, from shallow-marine strata
in the middle lower Eocene (“Capay Stage”) Crescent
Formation in the Black Hills, Thurston County, south¬
western Washington. Both specimens, however, are miss¬
ing their upper spire, thus positive assignment of this
species to a subgenus is not possible until more complete
specimens are found. This Washington species is very
similar to Mitrella sensu striata in having an unsculptured
shell (except for spiral ribs on the pillar), a short anterior
canal, denticles on the interior of the outer lip, and a
“varix”-like swelling preceding the outer lip. Mitrella
( Bastropia ) llajasensis n. sp. differs from this Washington
species by having a large size, more slender shell,
indented suture on spire, tabulate appearance to the
spire whorls, much longer siphonal pillar, spiral sculp¬
ture on the posterior part of the penultimate and last
whorls, no posterior channel-like indentation in the
aperture, and no indication of a “varix' -like swelling pre¬
ceding the outer lip.
PALEOBIOGEOGRAPHIC IMPLICATIONS
The details of the geologic range and locale of each of
the four known species of Bastropia are given in Table 1
and are shown chagramically in Figure 7. As discussed
earlier, a late Paleocene occurrence of Ba.stropia in
Alabama is questionable. Tbe earliest known documented
record of this subgenus is early Eocene (middle Ypresian)
in Alabama, and by the middle Eocene, it had spread
westward into nearby Texas.
From the late Paleocene through middle Eocene,
there were pulses of immigration of many shallow-
marine, warm-water (thermophilic) mollusks and other
marine organisms into the northeast Pacific region. Most
arrived from the Old World Tethyan Sea, Gulf Coast,
and Caribbean Sea regions by means of westward¬
flowing currents that passed through a low-latitude sea¬
way (Squires, 1987). The nearness in time between the
earliest confirmed record of Bastropia, in the early
Eocene in Alabama, and when it showed up in southern
California, in the late early Eocene, supports the conclu¬
sion that Bastropia arrived in southern California from
the Gulf Coast. Its dispersal would have been greatly
facilitated if its larvae had planktotrophic development.
This scenario is likely because its protoconch, which
consists of three smooth whorls, is similar to those of
previously mentioned (see “Discussion” of genus Mitrella )
inferred planktotrophic species of Mitrella (e.g., the extant
Mitrella pallanji and a Miocene species). The southern
California arrival of Bastropia was closely associated
with a warming event called the “Early Eocene Climatic
Optimum" (EECO), 51-53 million years ago (Figure 7).
During this event, global temperature reached a long¬
term maximum (Zachos et ah, 2008), and there was a
global-sea-level rise (Miller et ah, 2011), which would
have expedited the dispersal of marine organisms.
With the exception of a species in South Island,
New Zealand, Bastropia is restricted to North America.
This New Zealand occurrence is of late middle Eocene
age (Bartonian) and represents the youngest occurrence
of this subgenus. It also provides limited evidence that
Bastropia had an amphitropic distribution (Figure 7). The
route of dispersal that Bastroj)ia took to reach New Zealand
cannot be determined based on current information.
AC K N O WLEDG M E NTS
The manuscript benefited from careful reviews and
valuable comments by Lindsey T. Groves (Natural History
Museum Los Angeles County, Malacology Section) and
Marta deMaintenon (University of Hawaii, Marine
Science, Hilo).
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THE NAUTILUS 129(2):71-76, 2015
Page 71
Reinstatement of Led a rhytida Dali, 1908 and its reallocation
into Propeleda (Bivalvia: Nuculanidae)
Marina Giiller
Division Invertebrados
Museo Argentine de Ciencias Naturales “Bernardino Rivadavia”
Av. Angel Gallardo 470
(C1405DJR), Buenos Aires, ARGENTINA
[email protected]
Diego G. Zelaya
Departamento de Biodiversidad y Biologia Experimental
Facultad de Ciencias Exactas y Naturales - UBA
Ciudad Universitaria, Pab. 2, Lab. 31
(C1428EHA), Buenos Aires, ARGENTINA
[email protected]. uba. ar
ABSTRACT
In 1908, Dali described “ Leda ” rhytida as occurring off
Acapulco, Mexico. The species was never figured, and has
no subsequent published records. Despite that, it appears
mentioned by several authors in check-lists. The identity of
this species appears as controversial in the literature: some
authors emended the type locality to either eastern South
America or Chile, and the species was regarded either as a
possible synonym of Propeleda longicaudata (Thiele, 1912),
or considered as a nomen duhium. The aims of this study are
to determine the actual provenance of the type material, to
provide a proper redescription of the species, and to revise
its generic placement and current status. The study of the
syntypes and additional material from museum collections
reveals that the species actually comes from southern Chile.
Based on the elongated shell outline and distinctive hinge
morphology, the species is here reallocated into Propeleda.
Comparison of this species with P. longicaudata establishes
that P. rhytida is a distinct and valid species.
Additional Keywords: Protobranchia, Nuculanoidea, Magellan
Region, Chile, Taxonomy
INTRODUCTION
Leda (Leda) rhytida Dali, 1908 is among the numerous
new bivalve species described from the collections made
by the U.S. Albatross expedition along the Americas.
The original description is poor and lacking in detail,
and the species was never figured. According to Dali
(1908) the species was collected “off Acapulco, Mexico,
in 141 fathoms”, at Albatross station 3422, and the
type material was deposited at the United States National
Museum [National Museum of Natural History, Smithsonian
Institution], under number “122,918”. After examining
the type material. Keen (1971) detected an error in
the type locality originally indicated by Dali (1908).
Keen stated that the true provenance of the species
was “Patagonia, eastern South America” (sic). In con¬
trast, Bernard (1983) indicated the type locality to be
“Chile”. Despite that, Zamorano and Hendriekx (2012)
repeated “Acapulco” as the type locality. Apart from
the original collection, no new records for the species
are known. Recently, Coan and Valentich-Scott (2012)
suggested that Leda rhytida could correspond to a
synonym of Propeleda longicaudata (Thiele, 1912), a
species described from the Gauss Expedition in eastern
Antarctica, but currently regarded as widely distributed
in Antarctic and sub- Antarctic waters (Dell, 1964; 1990;
Villarroel and Stuardo, 1998). However, Bouchet (2014)
regarded Leda rhytida as a nomen duhium. The aims of
this contribution are to clarify the identity and precise
provenance of Leda rhytida, to provide a proper rede¬
scription of the species, and revise its generic placement
and its affinity with Propeleda longicaudata.
MATERIALS AND METHODS
We examined photographs of the type material of Leda
rhytida and Leda longicaudata (the latter currently treated
in the genus Propeleda), housed respectively at the
National Museum of Natural History, Smithsonian Insti¬
tution (USNM), Washington, and Zoologisches Museum
Berlin (ZMB). Additional material comes from the
collection of the Museo de Zoologla de la Universidad
de Concepcion (MZUC), Chile, and material sam¬
pled by the authors at the Beagle Channel and the
South Shetland (61°23'45" S, 55°26'33" W) and South
Orkney (60°58'53.4" S 43°26'42.6" W) (sub-Antarctic
and Antarctic waters), currently housed at the Museo
de La Plata (MLP), Argentina.
Shells were measured according to the parameters
and ratios indicated by Kamenev (2014), by using an
ocular micrometer mounted on a stereoscopic micro¬
scope, and a caliper for larger animals. Text abbreviations
are: A, length anterior to beaks; H, maximum height
perpendicular to L; L, maximum shell length; W, shell
width across closed specimens. Details of hinge plate
and protoconch were studied with a Phillips XL-30 scan¬
ning electron microscope (SEM), at the Museo Argentine)
de Ciencias Naturales “Bernardino Rivadavia” (MACN).
Page 72
THE NAUTILUS, Vol. 129, No. 2
SYSTEMATICS
Genus Propeleda Iredale, 1924
Type Species: Leda ensicula Angas, 1877 (OD)
Propeleda rhytida (Dali, 1908) new combination
(Figures 1-16)
Leda (Leda) rhytida Dali, 1908: 219 (listed only), 376
(description).
Propeleda longicaudata Thiele, 1912: Villarroel and
Stuardo, 1998: 142, figs. 113, 114 (in part).
Redescription: Shell club-shaped, delicate, moderately
large for genus (max. observed L = 13.2 mm), elongate
(H/L = 0.44 ± 0.01, n = 8), compressed (W/H = 0.49 ±
0.02, n = 8), slightly gapping posteriorly; equivalve;
markedly inequilateral: anterior end short, widely curved;
posterior end projected into a curved rostrum, three to
four times longer than anterior end (Figures 1-13).
Anterodorsal margin consisting of a very short, almost
horizontal section near beak, followed by a longer,
obliquely sloping distal section, which is widely arched
to slightly straight; both parts continuous or forming a
weak angulation (Figures 1-8). Anterior margin evenly
curved, joining anterodorsal margin in a slight angula¬
tion and continuous to ventral margin. Ventral margin
markedly convex at anterior half and nearly straight at
posterior half (Figures 1-8). Posterior (rostral) margin
somewhat convex, forming well-marked angles at junc¬
tions with posterior part of dorsal and ventral margins.
Posterodorsal margin long, slightly to markedly concave
(Figures 1-8). Umbones opisthogyrous, located on ante¬
rior third (A/L = 0.29 ± 0.02, n = 10), only slightly
raised from dorsal margin. Two strong carinae running
at posterior area of shell, along rostrum: one extending
from umbo to junction of ventral and posterior margins,
another from umbo to junction of postero-dorsal and
posterior margins (Figures 1-8). Shallow sulcus between
these carinae present. Anterior and central areas of shell
sculptured with regularly distributed, rounded, commar-
ginal cords, separated by narrow interspaces. These
cords extend over two posterior carinae and into sulcus.
Escutcheon large, well-defined by prominent dorsal
carina, sculptured with faint lines (Figure 9). Lunule
not differentiated. Prodissoconch small, about 165 pm
long, D-shaped, sculptured with a polygonal, net-like
pattern (Figure 14). Periostracum thin, well-adhered,
greenish-brown in wet specimens, straw-yellow when
dry (Figures 1-8). Inner shell surface white, glossy, with
a strong ridge extending along rostrum from umbo to
approximately half of posterior margin, separating inhal¬
ant and exhalant siphons (Figures 10-13). Pallial sinus
short, extending up to half of posterior adductor muscle
(Figure 12). Anterior adductor muscle scar ovate, per¬
pendicular to hinge; posterior adductor muscle scar nar¬
rower and smaller than anterior one, parallel to hinge
plate. Hinge plate narrow, taxodont, with two series of
teeth interrupted beneath beaks (Figures 12, 13). Anterior
series composed of about 15 teeth, posterior series with
about 27 teeth, regarding larger specimens. Teeth vari¬
able in size and morphology within each series. Ante¬
rior series: first two teeth closest to beak, low, U-shaped;
first evenly narrow, second with stronger ventral part
(Figures 15, 16). One or two following teeth almost
straight anti lamellar. Subsequent teeth V-shaped, with
ventral part gradually longer, increasing in size and height,
shaqily cusped. Most distal tooth greatly reduced in size.
Posterior series: five to eight teeth closest to beaks lamel¬
lar, similar in solidness (Figures 15, 16). Internal liga¬
ment stout, posterior to beaks, slightly larger than resilifer,
which is triangular, oblique, posteriorly directed. External
ligament narrow, elongated, amphidetic, larger anteriorly
and partially sunken below beaks (Figures 15, 16).
Type Locality: [Herein emended] U.S. Albatross station
2783, 51°02'30" S, 74°08/30" W, west coast of Patagonia,
141 fathoms [=256 m].
Type Material: 8 syn types (USNM 96918).
Material Examined: MZUC 10406, 1 specimen, 50° 03' S
74°41/ W, Isla Tobar, 200 m; MZUC 4642, 8 specimens,
and MZUC 4610, 19 valves, 50°09'55" S 74°43'75" W,
Confluence Canales Concepcion and Trinidad, 390—460 m;
MZUC 4627, 1 specimen and 4 valves, same data as
previous lot, but 460 m; MZUC 4646, 4 specimens and
4 valves, 51°00'50" S, 74°14'10" W, Puerto Bueno, Canal
Sarmiento, 21.5-220 m; MLP Mal4041 1 specimen, 54° 52' S,
68°28' W, Bahia Lapataia, Beagle Channel, 120-138 m.
Distribution: Southern Chile and Beagle Channel, living
at 120—460 in depth.
Remarks: Some discrepancies arise when comparing
the original labels of the material currently identified as
type of Leda rhytida (at the USNM) to information in
the original description, in particular in relation to the
repository number and the provenance of the types. The
repository number mentioned by Dali (1908) (“USNM
122,918”) actually corresponds to the holotype of Leda
lohula Dali, 1908, another species described by that
author in the same work. Dali indicated that this latter
species also originated from albatross station 3422 (off
Acapulco, Mexico). In addition, Dali did not provide a
repository number for Leda lohula. The specimen in lot
USNM 122918 agrees with the original description of
Leda lohula , particularly differing from Leda rhytida by
having an “ovate shape”, with both anterior and posterior
ends “rounded” (Dali, 1908: 375). In contrast, the origi¬
nal description of Leda rhytida refers to an “elongate-
ovate” shell, with the anterior end rounded and the
posterior one “produced” in a “strongly recurved ros¬
trum”. This set of characters is in accordance with the
shell morphology/ of specimens in lot USNM 96918, cur¬
rently labeled as syn types of L. rhytida (Figures 1-4),
whose label reads “Station 2783, west coast of Patagonia”.
Thus, it seems clear that there was an error in the pub¬
lished information of repository numbers and provenance
M. Giiller and D.G. Zelaya, 2015
Page 73
Figures 1-18. Propeleda rhytida. 1-4, 10, 11. Four syn types of Leda rhytida (USNM 96918). 1-4. External views. 10, 11. Internal
views. 5—9, 12-16. Specimens from 50°09'55" S, 74°43,75" W, southern Chile (MZUC 4642). 5-8. External views. 9. Dorsal view.
12, 13. Internal views (pallial sinus shown in black line on Figure 12). 14. Detail of prodissoconch. 15, 16. Details ol hinge plate.
15. Left valve. 16. Right valve. Scale bars: 1-11=5 mm; 12, 13 = 5 mm; 14 = 50 pm; 15, 16 = 500 pm. Abbreviations: IL = internal
ligament; EL = external ligament.
Page 74
THE NAUTILUS, Vol 129, No. 2
Figures 17-32. Propeleda longicaudata. 17. Original figure by Thiele (1912). 18-21. Two syn types of Lecla longicaudata (ZMB
63104). 18, 20. Externa] views. 19, 21. Internal views. 22-32. Propeleda el. longicaudata. 22, 23, 27. Specimens from South
Orkney (MLP 7467). 24-26, 28-32. Specimens from South Shetland (MLP 7468). 22-26. External views. 27. Dorsal view.
28, 29. Interna] views. 30. Detail of prodissoconeh. 31, 32. Detail of hinge plate. 31. Left valve. 32. Right valve. Scale bars:
17-27 = 5 mm; 28, 29 = 5 mm; 30 = 50 pm; 31, 32 = 500 pm. Abbreviations: IL = internal ligament; EL = external ligament.
M. Culler and D.G. Zelaya, 2015
Page 75
of the species. Keen (1971) already detected such error,
although she mistakenly wrote “eastern South America”
instead of "western South America” when emending the
type locality. Bernard (1983), on the contrary, did refer
to a locality on the Pacific coast, but was ambiguous in
stating “Chile”.
Based on the original label of the syntypes, the prove¬
nance of “Leda” rhytida is here emended to “51°02'30" S,
74°08'30" W, west coast of Patagonia” (which corre¬
sponds to Albatross station 2783). Study of additional
(conspecific) specimens from southern Chile and the
Beagle Channel allows us to confirm the occurrence of
the species in that area, and the nominal species is no
longer regarded as a nomen dubium (as suggested by
Bouchet, 2014) but as a valid species.
Generic Allocation ok the Species: The genus Leda
Schumacher, 1817 in which L. rhytida was originally
described is an objective junior synonym of Nuculana
Link, 1807. Allen and Hannah (1986) defined this genus
by the presence of robust and “moderately” posteriorly
elongated shells. Allen and Sanders (1996) added as a
diagnostic character the presence of a robust hinge with
V-shaped teeth and a small ligament, and the absence
of an internal posterior ridge running from umbo to
posterior margin. The fact that all teeth are V-shaped
in this genus can be confirmed from the illustrations of
Nuculana pemula (M filler, 1779) in Oliver et al. (2010)
(this species is currently regarded as a senior synonym
of Area rostrata Brugiere, 1789, the type species of
Nuculana) . In addition, these photographs show that the
internal ridge is extremely short in this species, being
present only in the distal part of the rostrum.
This set of characters does not match those present
in “Leda” rhytida, which has delicate, very posteriorly
elongated shells, with a long internal ridge extending
from the umbo to posterior margin, and bearing a large
internal ligament and differentiated lamellar teeth near
the beak. In contrast, the characters present in “Leda”
rhytida are in agreement with those present in Leda
ensicula Angas, 1877, the type species of Propeleda
Iredale, 1924 (Huber, 2010; Kamenev, 2014). Conse¬
quently, “Leda” rhytida is here reallocated into Propeleda.
The distinction of Propeleda from other nuculanid and
siliculid genera was discussed by Huber (2010) and
Kamenev (2014).
Affinities Between Propeleda rhytida and Propeleda
longicaudata: Coan and Valentich-Scott (2012) sug¬
gested that Propeleda rhytida (as Leda rhytida ) could
correspond to a synonym of Propeleda longicaudata
(reported by the authors under Nuculana) . The study of
the syntypes of the latter species (Figures 17-21) reveals
that both species show morphological resemblance.
Nonetheless, P. rhytida may clearly be distinguished by
having shells with shorter and more broadly rounded
anterior end, more projected ventral margin and poste¬
rior rostrum, stronger and more spaced commarginal
sculpture, and the two posterior carinae stronger (wider
and higher) than P. longicaudata . These differences lead
us to consider the two taxa as distinct species.
Propeleda longicaudata is currently regarded as a widely
distributed Antarctic and sub-Antarctic species, occur¬
ring in South Georgia, South Orkneys, South Shetlands,
Weddell Sea, Boss Sea and East Antarctica (Dell,
1964; 1990; Hain, 1990; Aldea and Troncoso, 2010).
The study of two lots of “P. longicaudata” from South
Orkney and South Shetland (MLP 7467, 7468, respec¬
tively) reveals some morphological differences between
these specimens and the syntypes of that species, namely
the presence of a consistently less recurved and longer
rostrum, which consequently generates a more elongated
shell outline (Figures 22-25, 28, 29). At present, the sig¬
nificance of these differences could not be determined,
because the syntypes are represented by only 3 valves,
and no additional specimens from the type locality were
available to evaluate the intraspecific variability of this
taxon. Therefore, we prefer to be conservative and refer
to the South Orkneys and South Shetland specimens
here studied as Propeleda ef. longicaudata. The speci¬
mens here studied of P. c£ longicaudata (Figures 22-32)
also differ from P. rhytida by having less inflated shells
(W/L = 0.16 ± 0.01, n = 10 vs. W/L = 0.22 ± 0.01, n = 8)
and a minor number of posterior teeth, even when con¬
sidering specimens of similar size (23 vs. 27 in specimens
about 13 mm long). Morover, in P. rhytida the lamellar
teeth at the posterior series are shorter and stouter, and
at the anterior series, the two teeth closest to the beaks
are consistently stronger (Figs. 15, 16 vs. 31, 32), and
the V-shaped teeth are taller and more delicate. Fur¬
thermore, judging from the material here available,
P. cf. longicaudata reaches a larger size (23.8 mm vs.
13.2 mm long in P. rhytida).
Propeleda longicaudata was also previously reported
as occurring in southern Chile by Villarroel and Stuardo
(1998). However, the study of the specimens they men¬
tioned (MZUC 4610, 4627, 4642, 4646) indicates that
they were misidentified by those authors. The lots are
here reassigned to Propeleda rhytida.
ACKNOWLEDGMENTS
The authors are grateful to Ellen Strong (USNM) and
Christine Zorn (ZMB) for providing photographs of
the type material. We are also indebted to Jorge Artigas
Coch (MZUC) and Gustavo Darrigran (MLP), for pro¬
viding access to the collections under their charge. Authors
are members of the Consejo Nacional de Investigaciones
Cientiflcas y Tecnieas (CONICET). This study was par¬
tially funded by PICT 2011-2182 to Diego G. Zelaya.
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Bouehet, P. 2014. Lech 7 rhytida Dali. 1908. Accessed
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I lain, S. 1990. Die beschalten benthischen Mollusken (Gastropoda
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profundas del sur del golfo de California. In: Zamorano,
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THE NAUTILUS 129(2):77-82, 2015
Page 77
Shells and fossils collected by the earliest settlers of Jamestown,
Virginia, USA
Juliana M. Harding
Department of Marine Science
Coastal Carolina University
P.O. Box 261954
Conway, SC 29528 USA
[email protected]
Beverly A. Straube"
Jamestown Rediscovery
1365 Colonial Parkway
Jamestown, VA 23081 USA
Brittany L. Grimm
Geerat J. Vermeij
Howard J, Spero
Department of Earth and Planetary Sciences
University of California Davis
Davis, CA 95616 USA
ABSTRACT
The discovery of an intact valve of the fossil bivalve Chesapecten
jeffersoriiiis and shells of three tropical snail species in a c. 1610
James Fort well speaks to the curiosity that European colonists
brought to die New World. While implementing die Virginia
Company of Londons mandate to identify and secure profitable
natural resources, the Jamestown, Virginia, colonists apparendy
also gathered interesting natural objects. The shells may have
been collected either as personal souvenirs, much like modern-
day tourists, or as curios destined for the lucrative European
conchology market. Chesapecten jeffersonius , Virginia’s state
fossil, was collected locally as representatives can still be found
in James River Pliocene deposits near Jamestown. In contrast,
the tropical shells were likely brought to Jamestown in May 1610
by survivors of an English shipwreck on Bermuda. The shells
from both Virginia and Bermuda were discarded in the fort’s
well by June 1610 as the settlers hastily prepared to permanently
abandon Jamestown.
Additional Keywords: Chesapecten jeffersonius , Lohatus gigas,
Strombus pugilis, Cittarium pica, Jamestown
INTRODUCTION
The Virginia Company of London sent colonists to
America to exploit natural resources including timber,
iron, and, particularly, gold. Most of the initial explor¬
atory efforts were focused upon finding resources that
could reward Virginia Company shareholders with imme¬
diate investment returns (Horn, 2005). Gold was a top
priority for the nascent colony, as reflected in Captain
John Smith’s lament that “there was no talke, no hope, no
worke, but dig gold, wash gold, refine gold, load gold”
(Smith, 1986a: 218).
Archaeological investigations of James Fort since 1994
by the Jamestown Rediscovery Project have uncovered
1 Corresponding author
2 Current address: 6 Valentine Court, Newport News, VA
23606, USA
evidence that early endeavors to extract profitable
resources included much more than the search for gold.
Some of the undertakings included specialists from
Germany producing glass; English metallurgists milking
trials of Virginia minerals that might alloy with English
copper to make brass; and a tobacco pipemaker pro¬
ducing tobacco pipes from the local clay (Straube, 2004;
Hudgins, 2005). The 17th-century Englishmen were also
interested in New World flora and fauna that could be
used for medicinal applications. Captain Gabriel Archer,
writing in 1607 of Virginias natural bounty, mentioned
“Apothecary drugges of diverse sortes, some knowne to
be of good estimacion, some strange, of whose vertue the
salvages report wonders” (Archer, 1969: 102).
Botanical and biological specimens were also prized
for being exotic. At the time of Jamestown’s founding,
European curiosity about the natural world resulted in a
market for objects of nature, especially those from newly
explored lands. Naturalia “was a commodity bought, sold,
bartered, and exchanged — the centerpiece of a series
of transactions that connected the world of commerce
to the study of nature” (Findlen, 2002: 298). Nobles,
wealthy gentlemen, and academics who could afford to
do so, assembled these objects in cabinets of curiosities
as eclectic reflections of status, records of travel, or tan¬
gible proof of their quest for knowledge. Indications of
this pursuit for authentic naturalia have been discovered
during archaeological excavations of James Fort’s earliest
contexts and particularly in the settlement’s first well
(Kelso et al. 2012: Structure 185).
In early June 1610, the well that had served as a water
source for the inhabitants of James Fort for two years
was quickly filled with half a million objects (Kelso et al.,
2012, Figure 1). Jamestown was being abandoned, and
these materials represented the remains of meals, the
detritus of everyday life, and objects that were not con¬
sidered valuable enough to transport to England. This
action was precipitated by the preceding winter, dubbed
by John Smith as the “starving time,” which had claimed
the lives of three out of four of the colonists and left
many of the survivors sick and malnourished (Smith,
Page 78
THE NAUTILUS, Vol. 129, No. 2
Jamestown
Current James River
shoreline
0 scale in feet 100'
J = grid areas excavated since 1994
James
River
Figure 1. Archaeological plan of James Fort (Courtesy of J. May, Jamestown Rediscovery Project, Preservation Virginia) on the
current shoreline of the James River, Virginia. Black arrow identifies the Jamestown well (Structure 185) from which the shells
were excavated.
1986b: 340). With few prospects of obtaining food, Gov¬
ernor Thomas Gates decided to sail to Newfoundland
where he hoped that the English fishing fleet could help
transport colonists the rest of the way home. Gates did
not want to attempt a transatlantic voyage from Virginia
as his four small vessels were overloaded with over
150 colonists, the provisions needed for the voyage,
the colony’s weaponry, and “all the best things in the
store,” which hopefully could be sold for profit upon
arrival in England (Strachey, 1973:76).
Four hundred years later, archaeological investigation
of the c. 1608-1610 James Fort well revealed a variety
of marine mollusk shells among the discarded items.
The assemblage included an intact fossil Chesapecten
jeffersonius valve (Say 1824) (Figure 2) as well as
seventeenth-century specimens of tropical marine snails
including the queen conch Lobatus g igas (Linneaus
1758) (Figure 3), the West Indian fighting conch
Strombus pugilis (Linneaus 1758), and the West Indian
top snail Cittarium pica (Linneaus, 1758, Figure 3). The
molluscan fauna recovered from this earliest Jamestown
well offer a glimpse into undocumented pursuits in the
early years of the colony.
DESCRIPTION OF THE SHELLS AND
HYPOTHESES FOR THEIR ORIGINS
Chesapecten jeffersonius ( Rival via) is a large temperate
scallop that occupied the shallow subtropical continental
shelf of North America during the Pliocene, 2-5 million
years B.P. Today, these marine sediments are exposed
across the mid-Atlantic US coastal plain due to the drop
in post-Pliocene sea level after the expansion of the
Antarctic ice sheet (Rovere et ah, 2014). This scallop
species is part of a unique faunal group used by geolo¬
gists to characterize the Sunken Meadow Member or the
oldest strata associated with the Yorktown formation.
J.M. Harding et al., 2015
Page 79
Figure 2. The Chesapecten jeffersonius valve discarded in
the earliest known Jamestown well as James Fort was being
abandoned in June 1610. Courtesy of M. Lavin, Jamestown
Rediscovery Project, Preservation Virginia.
Figure 3. Bermudan shells brought to the New World by Sea
Venture survivors that were thrown away in the Jamestown
well in June 1610 including Lobatus gigas (A) and Cittarium
pica (B). Both courtesy of M. Lavin, Jamestown Rediscovery
Project, Preservation Virginia.
The fossils used as the modem basis for tin's description
were collected from cliffs directly downriver of Sunken
Meadow Pond on the James River in Surry County,
Virginia (Ward and Blackwelder, 1980) (Figure 4).
When Thomas Say (1824) established the formal taxo¬
nomic designation for this scallop (as Pecten jeffersonius),
he relied on illustrations made by Martin Lister in 1687
(Historiae Conehvliorum, Liber III: plate 167; Ward and
Blackwelder, 1975). Thus, these scallops have the distinc¬
tion of being the first American fossils ever described
(Ward and Blackwelder, 1975). Although this species
has been extinct for ~4 million years (Krantz, 1991),
C . jeffersonius has played a role in modern paleoenvi-
ronmental reconstructions (e.g., Krantz 1990), paleon¬
tological stratigraphic characterizations (e.g.. Ward
and Blackwelder, 1975: Zone 1 Yorktown Formation;
Blackwelder, 1981: Sunken Meadow Member, Yorktown
Formation), and now offers a window into Jamestown’s
early years.
Ward and Blackwelder ( 1975) characterized Chesapecten
jeffersonius as having shell heights >12.0 cm where
height is the longest distance from the umbo to the
growth edge. The Jamestown C. jeffersonius valve is
15.6 cm high and 16.8 cm long (distance perpendicu¬
lar to height. Figure 2). This height is greater than
the largest valve height (13.77 cm) reported by Ward
and Blackwelder (1975) and the C . jeffersonius from
Kingsmill, Virginia, used by Krantz (1990, 10.3 cm
shell height). The Sunken Meadow Member paleo-
environment was a temperate, marine habitat in which
“many mollusks attained unusually large sizes” (Ward
and Blackwelder, 1980: D35). Kingsmill is on a northern
shore cliff downriver of Jamestown Island (Figure 4).
At Kingsmill, as well as in other James and York River
sites, fossils are deposited in the modern littoral zone,
as they erode from Yorktown Formation stratigraphic
layers, where they are easily collected.
James Fort was established on the banks of the James
River, which served as the colony’s major thoroughfare
and source of seafood. In the summer of 1609, in an
attempt to take pressure off Jamestown food and water
sources. Captain John Smith dispersed colonists 32 km
downriver to live on the “oyster banks” that flourished in
die saltier water of Hampton Roads and to an Indian
settlement near the mouth of the Nansemond River
where the English traded copper for food (Archer, 1969,
282; Earle, 1978) (Figure 4). An additional 120 colonists
were sent to the fresher water upriver near the ‘fall line’
of the James (Smith, 1998b: 220). It is probable that the
colonists stopped during river trips to investigate food
sources as well as to wait for favorable wind or tidal
conditions to facilitate travel.
As the colonists grew more familiar with the region,
they investigated inland areas as potential sources of
mineral wealth and also to mimic the native Algonquin
seasonal occupancy patterns. It is possible that the fossil
scallop was collected on a trip between Jamestown Island
and the temporary upriver settlement near the “falls”.
Sunken Meadows Pond and the adjacent cliffs, the type
locality for the Sunken Meadow Member of the Yorktown
formation (Ward and Blackwelder, 1980), are approxi¬
mately 16 km upriver of Jamestown Island on the opposite
shoreline (Figure 4). It is likely colonial travelers investi¬
gated these and other James River sites where Yorktown
Formation material was exposed during regular travel as
well as when on exploratory expeditions.
A less likely scenario is that the scallop was discovered
during Smiths initial exploration of the Chesapeake Bay
in summer 1608 and retained as a souvenir. Chesapecten
jeffersonius occurs in Yorktown formation deposits located
along the track of Smith’s second Chesapeake voyage
(Jul. -Sep. 1608, Clark et ah, 2007). The last legs of
this voyage included areas with C. jeffersonius deposits
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THE NAUTILUS, Vol. 129, No. 2
Figure 4. Map of the James River, Virginia, USA showing the likely collection sites for the fossil scallop in relation to Jamestown/
James Fort with the Hampton Roads region (shaded) indicated for reference.
on the western Chesapeake shoreline south from the
Patuxent River into the Rappahannock, Piankatank,
Poquoson, Elizabeth, and Nansemond Rivers (Clark
et ah, 2007).
The possibility that the scallop was a gift to Smith from
one of the Algonquin chiefdoms he visited is more
intriguing but even less likely than the other scenarios.
Although Virginias native Algonquin tribes used small
fossil gastropods (such as the marginellid Prunum
limatulum (Conrad, 1834)) as embroidery ornaments for
ceremonial robes (e.g., “Powhatan’s Mantle”, Rountree
and Turner, 2002: 115-116), Algonquin use of C.
jeffersonius was not documented by the early colonists.
Preservation of a large intact scallop shell through
the rigors of James Fort life during the “starving time” is
impressive. One can only wonder whether the original
collector died and the fossil simply became another item
to be discarded when the order was given to abandon
the fort. While considered of value at one point, it was
deemed of little worth relative to food and colonists
since space on the departing ships was limited. Other
fossils (e.g., sharks teeth) have also been discovered by
archaeologists in other James Fort contexts from the
1607-1610 period. Fossils would not have been recog¬
nized in the early 17th century as representing extinct
ancient life but rather as minerals formed by natural
processes in the earth, which sometimes resembled
living organisms (Cook, 2003; MacGregor, 2007).
Shells of three tropical snail species were also depos¬
ited in the Jamestown well in June 1610. The tropical
snails (Figure 3) are found neither in the Yorktown
formation nor in. modern Chesapeake habitats. In fact,
Cittarium pica does not currently live in Bermuda. Shells
of this species from Bermuda date from the Pleistocene,
but are today (as in the 1600s) commonly inhabited by
the terrestrial hermit crab Coenobita clypeatus (Fabricus,
1787; Olson and Hearty, 2013). These tropical shells were
most likely collected in Bermuda between July 28, 1609
J.M. Harding et al., 2015
Page 81
and May 10, 1610 (Strachey, 1973). These dates encom¬
pass the time span between the Bermuda wreck of
the Sea Venture, carrying prospective Jamestown colo¬
nists, and the survivors’ departure for Jamestown almost
10 months later. The shipwreck survivors, including the
newly appointed governor, Sir Thomas Gates, arrived at
Jamestown shortly before the colony’s abandonment in
June 1610. There was no other recorded contact between
Jamestown and Bermuda prior to this date. Further,
accounts of the voyage from Bermuda to Virginia do
not mention any landfall, but it is possible that either
the queen conch or the West Indian fighting conch
might have been collected during a brief stop for pro¬
visions or water along the Southeast US coast since
both species historically ranged as far north as Georgia
(Abi)ott, 1974).
We hypothesize that the Bermudan shells were col¬
lected by the Sea Venture castaways as attractive curios,
much like modern travelers pick up sea shells. Knowl¬
edge of the shells’ intrinsic worth to collectors in Europe
may have been another motivating factor for the colo¬
nists’ interest in keeping these objects. The study of
mollusean shells, known as conchology, developed as a
scholarly discipline during the 17th century (Huxley,
2003). Shells from around the world could be sold
to private European natural history collectors for a con¬
siderable profit. The discovery of specially gathered
shells in the fort well speaks to the survival mentality of
the remaining colonists when they abandoned Jamestown
in June 1610.
SUMMABY
Mollusean fauna recovered from the earliest Jamestown
well offer a glimpse into undocumented pursuits on the
part of the early English colonists relating to the devel¬
oping commodification of natural objects. Virginia and
its early 17th-century English settlement were part
of the growing network of global trade that placed
value and significance on the exotic, including objects
of nature. A fossil Chesapecten jejfersoniu.s was col¬
lected locally, most likely by a colonist exploring the James
River in search of merchandisable commodities. Tropical
snail shells gathered in Bermuda during the winter of
1609-1610 were sufficiently interesting and perhaps valu¬
able, to be saved and transported by the Sea Venture
shipwreck survivors to James town in May 1610. Despite
the ascribed values and significance that led to them
being picked up initially, all of the shells were ultimately
discarded in June 1610 during the swift abandonment of
James Fort. This departure was only thirty hours in dura¬
tion as the colony was providentially revitalized by the
arrival of the colony’s new governor with copious provi¬
sions and fresh settlers. But, in retrospect, if the settle¬
ment had not experienced this temporary major setback
it is very unlikely that the fossil and tropical mollusks
would have been discarded at Jamestown for archaeolo¬
gists to find and interpret 400 years later.
ACKNOWLEDGMENTS
The authors would like to acknowledge the staff of the
Jamestown Rediscovery Archaeological Project, especially
those involved in the excavation, curation, and inter¬
pretation of the well (Structure 185). These include
Dr. W. Kelso, D. Schmidt, J. May, D. Givens, M. Lavin,
M. Richardson, D. Smith, D. Gamble, and D. Warmke.
BLG acknowledges the support of the University of
California, Davis, Dept, of Earth Sciences, Durrell Fund
during the completion of her M.S. research. We thank
Dr. M.G. Harasewyeh for early discussions on the
identification of the scallop and helpful information on
Bermuda mollusks. We also thank C.T. Sailer for her
comments on an earlier draft of this manuscript.
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Geological Survey Bulletin v. 1482-D, pp. 1-61.
THE NAUTILUS 129(2):83-89, 2015
Page 83
Two new species of Tryonia (Caenogastropoda: Cochliopidae)
from die late Pleistocene of Coahuila, northern Mexico
Alexander Czaja
Jose Luis Estrada-Rodriguez1
Department of Biological Sciences
Juarez University of the State of Durango (UJED)
35010 Gomez Palacio
Durango, MEXICO
[email protected]
ABSTRACT
Two new species of Pleistocene freshwater snails of the genus
Tryonia Stimpson (1865) from Coahuila, northern Mexico, are
described. One of them, Tryonia hershleri new species, belongs
to a group of species that have shells with characteristic promi¬
nent axial ribs. The second species, Tn/onia pseudocircumstriata
new species, is similar to recent and fossil species from the Pecos
River in the southwestern United States. Contrary to earlier prac¬
tice, we propose to use different species names for Pleistocene
and extant Tryonia from the Pecos River.
Additional Keywords: Late Quaternary, Tryonia circumstriata,
Tryonia stocktonensis
INTRODUCTION
Although little investigated, the desert regions of north¬
eastern Mexico contain abundant lacustrine sediments
with rich late Quaternary mollusean faunas (Czaja et ah,
2014a; 2014b). Specimens of Tryonia pseudocircumstriata
new species were collected from eolian sediments (inland
dunes) of an ancient paleolake that occupied a huge drain¬
age basin in the central Laguna Region, Coahuila, Mexico
(Figure 1). Tryonia hershleri was collected in the same
region but from fluvial-lacustrine deposits.
These sites contain one of the most diverse mollusean
faunas of the late Quaternary of northern Mexico, with
countless specimens preserved mainly in dunes sediments
(Czaja et ah, 2014a). The fossil material shows an excellent
preservation that allows for detailed species description.
The genus Tryonia contains 33 recent aquatic species
distributed mainly in western North America, with one
species known from Guatemala and Tryonia porrecta
from Hawaii. More than half of the species of Tryonia
occur in the Chihuahuan Desert, where the genus shows
1 Corresponding author
a high level of local endemism. However, many endemic
species of Trt/onia are endangered or already extinct
because freshwater habitats are disappearing, mainly
as a result of human activities (groundwater mining)
(Hershler et al„ 2011; 2014).
Fossil members of the genus have been described
from the Miocene of Venezuela, Peru, Colombia, Brazilian
Amazonia, and from the Pliocene of Guatemala
(Wesselingh and M acsotay, 2006). Uncertain are Miocene
records of Tn/onia from the southwestern USA, hut some
of the Pliocene shells described as Hydrohia and
Calipyrgula by Pilsbry (1935) from the Kettleman Hills,
California, probably belong to Tryonia (R. Hershler, pers.
comm.). So far, Pleistocene records of the genus had been
known only from the United States (Hershler, 2001).
The aim of the present study is to describe the new
species and to compare them with living and fossil mem¬
bers of the genus Tryonia. The present investigations
form part of a broader study of fossil land and freshwater
mollusks (ecology, systematic^, and biogeography) from
the region that begun in 2013 (Czaja et ah, 2014a; Czaja
et ah, 2014b).
MATERIALS AND METHODS
Shells of Tryonia hershleri new species were collected at
Acatita Valley, Coahuila, Mexico, ca. 8 km north of Charcos
de Risa (26°16'53.77" N, 103°4'25.96" W). These shells
were collected in Pleistocene deposits exposed along the
western side of a road to Laguna del Rey. The shells
derive from a gravel-sand layer of fluvial-lacustrine
deposits and were collected directly from a cut of approx¬
imately 2 m thickness. Similar alluvial sandy gravel beds
known from various sites in the vicinity have been dated
as late Pleistocene (~15,000 BP, l4C dates) by Butzer
et al. (2008).
The shells of Tryonia pseudocircumstriata new species
were collected from dune sediments in the south part of
the Paleo-Lake Irritila, Coahuila, 300 m W of the locality
Gabino Vazquez (25°26' 17.45" N, 102°55'12.93" W).
Page 84
THE NAUTILUS, Vol. 129, No. 2
104°30'0"W
IWOTO
103'30'0‘W
103*0‘0"W
102°30'0‘W
102'crcrw
101*30-0*W
LEGEND
• Dunes
■ Mayor Cities
I Paleo-Lake Irritila Model
77.71 States
Tryonia hershleri
^ Tryonia pseudocircumstriata
World Imagery
Low Resolution 15m Imagery
High Resolution 60cm Imagery
High Resolution 30cm Imagery
Mexico
Paleo-Lake
Figure 1. Map of the Paleolake Irritila in Coahuila. Mexico showing the collecting localities of the new species described in this
paper (modified from Czaja et ah, 2014a).
These eolian dunes are of Holocene age but their fossil
content is older and comes from late Pleistocene lacus¬
trine sediments (Butzer et ah, 2008; Czaja et ah, 2014a).
Unconsolidated sediments of both sites were screened
through 0.5 mm and 0.3 mm sieves. The fossils were
photographed with a Zeiss AxioCam ERc5s microscope-
camera. The collected material is housed at the Faculty
of Biological Science of the Juarez State University of
Durango (UJED), Campus Gomez Palaeio.
SYSTEMATIC PALEONTOLOGY
Class Gastropoda Cuvier, 1795
Subclass Caenogastropoda Cox, 1960
Superfamily Truncatelloidea Criscione and Ponder, 2013
Family Cochliopidae Tryon, 1866
Genus Tryonia Stimpson, 1865
Type Species: Tryonia clathrata Stimpson, 1865 (by
original designation).
Tryonia hershleri new species
(Figures 2-10, 16, Table 1)
Diagnosis: Shell medium- to large-sized, conical to tur-
reted, with robust axial ribs, aperture ovate; distinguished
from similar congeners by the pattern of shell sculpture
(see taxonomic remarks).
Description: Shell medium-sized, conical to turreted,
having 4.75-6.25 weakly shouldered whorls, strongly but
variable sculptured. Height 3.50-5.95 mm, width 2.05-
2.90 mm (shell measurements in Table 1). Protoconch
smooth. Sculpture from low to robust axial ribs or almost
spinose projections, ribs beginning at about second whorl,
reaching their maximum size at whorl 4/5. Approximately
45 ribs present, running almost from suture to suture.
About 16 ribs present on body whorl. Ribs of large shells
connected with up to 5 spiral keels. Spiral lines present
on middle and lower part of whorl height. Axial ribs and
spiral cords forming reticulated pattern (Figure 8). Male (?)
shells only with ribs, without spiral cords (Figure 4). Aper¬
ture ovate pyriform, adnate, angulate above, apertural
margins not thickened on male shells. Umbiculus narrow
or absent. Shells do not show great intraspecific morpho¬
logical variation but sexual dimorphism is pronounced.
Operculum not preserved.
A. Czaja and J. L. Estrada-Rodriguez, 2015
Page 85
Figures 2-9. Tryonia hershleri Czaja and Estrada-Rodriguez new species. 2-3. Holotype (UJMC-130). 4. Paratype (UJMC-131),
possible male shells. 5. Brown colored specimen (UJMC-132a), 6. Small specimen (UJMC-L32b). 7. Apical view of shell showing
spiral sculpture on teleoconch (Holotype). 8. Shell sculpture widi axial ribs and spiral keels. 9. Specimen (UJ’V1C-133). Scale bars = 1mm.
Type Material: Holotype (Figures 2, 3, 7), UJMC-130,
5.45 mm height x 2.63 mm width, 1.95 mm aperture
length, 1.20 aperture width. Paratype 1 (Figure 4),
UJMC-131, 4.05 mm height x 2.05 mm width; 1.25 mm
aperture length, 1.05 aperture width. Paratype 2, UJMC-
132, 5.55 mm height x 2.95 mm width; 2.00 mm aperture
length, 1.25 aperture width. Alexander Czaja and Jose
Luis Estrada-Rodriguez coll., 2013. All from type locality.
Type Locality: Late Pleistocene gravel deposits,
Acatita Valley, west side of the road to Laguna del Rey,
8 km north of Charcos de Risa, Coahuila, Mexico,
26°16'53.77" N, 103°4'25.96" W.
Stratum Typicum: Late Pleistocene.
Material Examined: 320 specimens from the type locality.
Etymology: The new species is named on honor of
Dr. Robert Hershler, Smithsonian Institution, for his
investigations on the genus Tryonia.
Geographic distribution: So far only from the Paleo-
Lake Irritila, Coahuila, Mexico.
Taxonomic Remarks: Tn/onia hershleri new species
shows morphological resemblance with species of the
genus with similar shell sculpture (Figures 1 1-15). We
include in this informal group the following fossil species:
T. spiralistriata Wesselingh
T. gaatemalensis Wesselingh
T. vivas i Wesselingh and Macsotay
T. nuttalli Wesselingh
T. scalarioides tuherculata (de Greve)
T. scalarioides scalarioides (Etheridge)
The only living members of this informal Clathrata-
group are the type species T. clathrata Stimpson from
Nevada, United States, and T. exigua (Morelet) from
Lake Peten Itza, Guatemala.
The species in this group have axial ribs similar to
T. hershleri new species, but the most similar is T. vivasi
from the Cumaea Formation, Middle Miocene of
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THE NAUTILUS, Vol. 129, No. 2
Figure 10. Drawing ol Tryonia hershleri Czaja and Estrada-
Rodrfguez new species (Drawing of Fernando Hernandez).
Venezuela. This South American fossil species have
spiral sculpture very similar to that of T. hershleri new
species (Figure 12). The difference is that, while T. vivasi
has up to 8 fine spiral lines, the new species from Coahuila
develops 2—4 coarse spiral cords. The aperture height in
relation to the height of the shell in T. hershleri
new species is larger than in T. vivasi, but the most
significant difference between these two species is their
size. The smallest specimens of T. vivasi are 6.5 mm
long, the longest shells of T. hershleri new species mea¬
sure 5.6 mm.
Shells of Tryonia scalaroides tuberculata Wesselingh
from the Miocene Pebas Formation of Peruvian Amazonia
also have spiral sculpture but only two cords on each
whorl and three on the body whorl (Figure 13). Further¬
more, this Amazonian species differs from T. hershleri
new species by having a different general shape (weaklv
convex whorls) and one or two more whorls.
Other members of the Clathrata- group such as
T. spiralistriata Wesselingh, T. clathrata Stimpson and
T. scalarioides scalarioides (Etheridge) lack the charac¬
teristic keels (or have them very poorly developed) on
the whorls and therefore the general pattern of the
sculpture is different (Figures 11, 14, 15).
Tryonia pseu do e i rcumstriata new species
(Figures 17-27, Table 2)
Diagnosis: Shell medium- to large-sized, conical, aper¬
ture ovate, distinguished from congeners by different
ornamentation of the whorls (regularly spaced spiral lirae
between sutures).
Description: Shell up to 6.5 mm high and 1.61-
2.33 mm wide (shell measurements in Table 2), elongate
to conic with 5.25-7.00 very weak convex whorls.
Protoconch smooth (Figure 21). Teleoeonch sculptured
with more or less regularly spaced spiral lirae between
sutures, ranging in number from 5-7 on third whorl to
12 on body whorl; last three whorls with weak collabral
ribs that cross spiral sculpture. Aperture ovate -pyriform
and angulate above, lip thin, adnate. Shell imperforate or
narrowly umbilicate. Shells white (typical for Pleistocene
material), original coloring and operculum not preserved.
Type Material: Holotype (Figure 17, 18), UJMC-143,
5.05 mm height x 2.05 mm width; 1.25 mm aperture
length, 1.05 aperture width; Paratvpe 1 (Figures 19, 20),
UJMC-144, 5.15 mm height x 1.95 mm width; 1.30 mm
aperture length, 1.05 aperture width; Paratype 2
(Figures 22, 23), UJMC-145, 4.95 mm height x 1.80 mm
width; 1.25 mm aperture length, 1.05 aperture width.
Alexander Czaja and Jose Luis Estrada-Rodrfguez coll.,
2013. All from type locality.
Table 1. Try onia hershleri new species, shell measurements (in mm). Symbols used are; x = mean; SD = standard deviation;
N = sample size.
Specimen 1 2 3 4 5 6 7 8 9 10 II 12 13 14 15 16 17 18 19 20 x SD N
Length 5.05 4.52 3.94 5.03 4.38 4.01 3.61 3.62 4.01 3.65 4.19 4.28 4.81 4.61 4.32 3.92 5.21 3.61 4.01 4.39 4.26 1.360 20
Width 2.20 2.25 1.99 2.23 2.31 2.12 2.03 2.02 2.10 1.97 2.12 2.29 2.60 2.51 2.33 2.03 2.39 1.87 1.99 2.20 2.18 0.190 20
No. whorls 5.50 5.25 5.25 6.25 5.25 5.00 5.00 4.75 5.25 5.00 5.25 5.00 5.25 5.00 6.00 5.25 6.25 5.00 5.00 5.25 5.29 0.416 20
A. Czaja and }. L. Estrada-Rodrfguez, 2015
Page 87
Figures 11-16. Comparison of sculptured Tnjonia species. 11. Tnjonia spirdUstriata Wesselingh and Maesotay (from Wesselingh
and Maesotay, 2006, fig. 4b, reprinted with author’s permission). 12. Tryonia vivasi Wesselingh and Maesotay (from Wesselingh and
Maesotay, 2006, fig. 3a, reprinted with author’s permission). 13. Tnjonia scalarioides tuherculata (de Greve) Wesselingh (from
Wesselingh, 2006, Fig. 27b, reprinted with author’s permission). 14. Tnjonia scalarioides scalarioides (Etheridge) Wesselingh (from
Wesselingh, 2006, Fig. 24b, reprinted with author’s permission). 15. Tnjonia clathrata Stimpson (from Hershler, 2001, fig. 4E,
reprinted with author’s permission). 16. Tnjonia hershleri new species Scale bars = 1mm.
Figures 17-26. Tnjonia pseudocircunistriata Czaja and Estrada-Rodrfguez new species. 17-18. Holotype (UJMC-143).
19-20. Paratype 1 (UJMC-144). 21. Apical view of shell showing spiral sculpture on teleoconeh (Holotype). 22-23. Paratype 2
(UJMC-145), smooth specimen. 24-26. Width specimen (UJMC 146). Scale bars = 1mm.
Page 88
THE NAUTILUS, Vol. 129, No. 2
Figure 27. Drawing of Tryonia pseudocircumstriata
Czaja and Estrada-Rodrfguez new species (Drawing of
Fernando Hernandez).
Type Locality: Dunas Bilbao near Viesca, Coahuila,
late Pleistocene, 25026'17.45" N, 102o55'12.93" W.
Stratum typicum: Late Pleistocene.
Material examined: 85 specimens from type locality.
Etymology: The species name alludes to its morpho¬
logical resemblance to Tryonia circumstriata (Leonard and
Ho, 1960) from the Pleistocene of Pecos County, Texas,
United States.
Geographic Distribution: Thus far known only from
Bilbao Dunes, southern part of the Paleo-Lake Irritila,
Coahuila, Mexico.
Taxonomic Remarks: The most characteristic feature
of this new species is its teleoconch with regularly spaced
spiral lirae (Figure 27). Tryonia pseudocircumstriata
new species closely resembles (as its name indicates)
T. circumstriata from the Pecos River deposits, Texas,
United States. This species was first described by
Leonard and Ho (I960) based on Pleistocene material.
Subsequently, in the vicinity of the Pleistocene site, a
living species was found by Taylor (1987) and named
Tryonia stocktonensis (Gonzales Spring Snail). However,
Hershler and Thompson (1992) did not find difference
between the fossil and Recent material and place
T. stocktonensis in synonymy with T. circumstriata. How¬
ever, in our opinion, the Pleistocene and recent shells
from Texas are different and should not be considered
as a single species. The general shape of the fossil and
the recent shells is clearly different, as made evident by
the different convexity of the whorls (Figures 28-30).
Sutures are also different, very deeply incised in the
Pleistocene shells and shallow in the recent material
(Figures 28, 29). Finally, there are differences in the size
of the shells that go beyond intraspecific variability.
According to Taylor (1987), the recent and endemic
T. stocktonensis have a length range of 2.34-3.67 mm,
whereas the fossil shells are within the 3.7— 4.9 mm
range, clearly larger.
We think that these mentioned morphological differ¬
ences between the Pleistocene and recent shells from
Texas make it impossible to place both in the same spe¬
cies. We propose to use the name Tryonia circumstriata
only for the Pleistocene material from the Pecos River
and Tryonia stocktonensis Taylor for the Recent endemic
shells of the same region.
Our Pleistocene material from Coahuila shows gener¬
ally more similarities with the Pleistocene shells from
Table 2. Tryonia pseudocircumstriata new species, shell measurements (in mm). Symbols used are: x = mean; SD = standard
deviation; N = sample size.
Specimen 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 x SD N
Length 4.00 5.00 4.00 3.31 4.09 4.72 4.00 3.71 4.00 4.00 4.00 5.01 4.92 4.32 4.12 4.00 3.99 4.00 5.00 4.33 4.23 0.468 20
Width 1.91 1.81 1.89 1.40 1.88 1.95 1.92 1.81 1.59 1.99 1.97 2.09 2.00 1.92 1.61 1.86 1.59 1.75 2.00 1.98 1.85 0.176 20
No. whorls 5.25 6.00 6.00 5.00 6.00 7.00 6.00 6.00 6.00 7.00 6.75 7.00 6.25 6.00 6.00 6.25 6.00 6.25 7.00 6.00 6.19 0.543 20
A. Czaja and J. L. Estrada-Rodrfguez, 2015
Page 89
Figures 28-32. Comparison of Tryonia pseudocircumstriata new species with similar Pleistocene and present-day species.
28. Tryonia stocktonensis Taylor, present-day endemic species from Diamond Y Draw, Pecos County, Texas (from Hershler 2001.
Fig. 4H, USNM 883958, reprinted with authors permission). 29. Tryonia circumstriata (Leonard and Ho) from Late Pleistocene of
Pecos River deposits, Terell County, Texas (holotype Calipyrgula circumstriata Leonard and Ho, 1960, Plate 12, Fig. 1, No. 11301).
30. Tryonia circumstriata (Leonard and Ho) from Late Pleistocene of Pecos River deposits, Terell County, Texas (paratype 1
Calipyrgula circumstriata Leonard and Ho, 1960, Plate 12, Fig. 2, No. 11302). 31. Tryonia pseudocircumstriata new species,
holotype (UJMC-143). 32. Tryonia pseudocircumstriata new species, paratype (UJMC-145). Scale bars = lmm.
Texas (Figures 29-32). The shells from Coahuila differ
from fossil T. circumstriata in having only slightly convex
whorls while the species from Texas have more rounded
whorls. The apertures are also different: in all specimens
of T. pseudocircumstriata new species the inner lip is
always ad n ate to the pariental wall while specimens of
T. circumstriata show a narrow gap between the lip and
pariental wall (Figures 29-32).
ACKNOWLEDGMENTS
The authors are grateful to Dr. Robert Hershler,
Smithsonian Institution, who first recognized that the
Pleistocene specimens from Coahuila are new species of
the genus Tn/onia. Special thank goes to Dr. Heike Reise,
Senekenberg Museum fur Naturkunde Gorlitz, Germany,
for her assistance in the revision of the English text.
LITERATURE CITED
Butzer, K.W., J.T. Abbott, C.D. Frederick, P.H. Lehman, C.E.
Cordova, and J.F. Oswald. 2008. Soil-geomorphology and
“wet” cycles in the Holocene record of North-Central
Mexico. Geomorphology 101: 237-267.
Czaja, A., M.R. Palaeios-Fest, J.L. Estrada- Rodriguez, U.
Romero-Mendez, and J.A. Alba-Avila, A. 2014a. Inland
dunes fauna from the Paleolake I rritila in the Comarca
Lagunera, Coahuila, Northern Mexico. Boletin de la
Sociedad Geologica Mexicana 66 (3): 541-551.
Czaja, A., J.L. Estrada-Rodriguez, and U. Romero-Mendez.
2014b. Freshwater mollusks of the Valley of Sobaco,
Coahuila, Northeastern Mexico — a subfossil ecosystem
similar to Cuatrocienegas. Boletin de la Sociedad Geologica
Mexicana 66 (3): 459—469.
Hershler, R. 2001. Systematic^ of the North and Central
American aquatic snail genus Tryonia (Rissooidea:
Hydrobiidae). Smithsonian Contributions to Zoology 612:
1-53.
Hershler, R., H.-P Liu, and J.J. Landye. 201 1. New species and
records of springsnails (Caenogastropoda: Cochliopidae:
Tn/onia) from the Chihuahuan Desert (Mexico and
United States), an imperiled biodiversity hotspot. Zootaxa
3001: 1-32.
Hershler, R., Landye, J.J., H.-P. Liu, M. de la Maza-Benignos,
P. Ornelas, and E.W. Carson. 2014. New species and records
of Chihuahuan Desert springsnails, with a new combination
for Tn/onia brunei. Western North American Naturalist
74 (1): 47-65.
Hershler, R., and Thompson, F.G. 1992. A review of the
aquatic gastropod subfamily Cochliopinae (Prosob ranchia:
Hydrobiidae). Malacological Review Supplement 5: 1-140.
Leonard, A.B. and T.-Y. Ho. 1960. New Calipyrgula from Pleis¬
tocene of Texas and notes on Cochlio/nna rio grande mis . The
Nautilus 73: 125-129.
Pilsbry, H.A. 1935. Mollusks of the fresh-water Pliocene beds
of the Kettleman Hills and neighboring oil fields, California.
Proceedings of the Academy of Natural Sciences
Philadelphia 86: 541-570.
Taylor, D.W. 1987. Fresh-water molluscs from New Mexico and
vicinity. New Mexico Bureau of Mines and Mineral
Resources Bulletin 116: 1-50.
Wesselingh, F.P. 2006. Molluscs from the Miocene Pebas
Formation of Peruvian and Colombian Amazonia. Scripta
Geologica 133: 19-290.
Wesselingh, F.P and O. Macsotay. 2006. A Miocene molluscan
faunule from Caueagua (Miranda State, Venezuela), with
the description of a new species of Tn/onia (Mollusca,
Gastropoda ) Cainozoic Research 4 (1-2): 61-65.
THE NAUTILUS I29(2):90-93, 2015
Page 90
A new Latiromitra (Gastropoda: Ptychatractidae )
from the Gulf of Mexico
Emilio F. Garcia
115 Oakcrest Dr.
Lafayette, LA 70503 USA
efg21 [email protected]
ABSTRACT
Latiromitra niveobabelis new species is described and com¬
pared with the Philippine species Latiromitra barthelowi
(Bartseh, 1942), and with the Atlantic species Latiromitra
cryptodon (P. Fischer, 1882), L. aratiuncula (Quinn, 1981),
L. meekiana (Dali, 1889), L. costata (Dali. 1890) and L styliola
(Dali, 1927).
INTRODUCTION
For some twenty years the Biology Department at the
University of Louisiana at Lafayette (ULL), has con¬
ducted a series of cruises in the Gulf of Mexico utilizing
the R/V Pelican, a ship managed by the Louisiana Univer¬
sities Marine Consortium (LUMCON). The most recent
cruise, one of five executed under the Gulf of Mexico
Research Initiative (GoMRI), was conducted in Septem¬
ber, 2014. The cruise sampled deep-water areas west of
the Diy Tortugas. It was in this southeastern quadrant
of the Gulf where an empty specimen of an unknown
species of the genus Latiromitra was dredged.
The genus Latiromitra has had a controversial saga in
its systematic history. Species in that genus having been
placed in Buccinidae (later transferred to Costellariidae)
by Thiele (1929), in Fasciolariidae and Mitridae by Dali
(1889 and 1890; and 1927 respectively), in Volutidae by
Weaver and Dupont (1970) and in Volutomitridae by
Cernohorsky (1970). Bayer (1971: 196) noted that the
species, placed in Latiromitra, had “a closer affinity with
the family Turbinellidae”, and latter workers followed his
lead. Although Quinn (1981: 72) thought it necessary to
create the new genus Cyomesus for some members of
this group, Bouchet and Waren (1985: 255) concluded
that Quinns taxon was synonymous with Latiromitra.
Kantor (2014) places Latiromitra in Ptychatractidae,
and I here follow his lead. The familial placement of the
genus is tentative, as this is a “taxonomieally complicated
group” (Kantor, personal communication, Feb. 3, 2015),
very similar to some of the deep water Costellariidae.
Species assigned to the genus Latiromitra are rela¬
tively morphologically conservative; they are represented
in the western Atlantic by five species. The new species
proposed herein is known from only one empty shell;
however, several of its conchological characters are quite
distinct from any other Latiromitra.
SYSTEMATICS
Family Ptychatractidae Stimpson, 1865
Genus Latiromitra Loeard, 1897
Type Species: Latiromitra specialis Loeard, 1897,
by monotypy.
Latiromitra niveobabelis new species
(Figures 1-5)
Diagnosis: A milky-white, almost straight-sided, nar¬
rowly fusiform shell with a constricted subsutural band
bordered posteriorly by strong, adpressed crenulations
and anteriorly by a heavily nodulose spiral cord.
Description. Holotype 46.0 mm in length, strong, nar¬
rowly fusiform (length/width ratio 3. 36); last teleoconch
whorl 21.1 mm in length. Protoconch broadly conical
(Figure 3), light amber in coloration, of approximately
2.5 whorls; first whorl damaged, smooth; second whorl
smooth; last whorl developing weak, prosoeline axial rib-
lets on approximately last half of whorl (Figure 4); riblets
growing stronger at termination of whorl. Transition to
teleoconch abrupt, with a strongly prosoeline transitional
growth mark, a change in coloration from amber to milkv-
white, and a development of a round axial rib at start
of teleoconch (Figures 3-4). Teleoconch of 8.5 whorls;
whorls subsuturally constricted (Figure 5), only slightly
convex anteriorly; last whorl strongly convex peripher¬
ally. Suture deep, heavily crenulate; crenulations strongly
adpressed to earlier whorl, creating a narrow channel
(Figure 5), this feature first appearing between second
and third teleoconch whorls. Axial sculpture of strong,
rounded ribs; ribs as wide as interspaces; 10 ribs on first
E.F. Garcia, 2015
Page 91
Figures 1-10. Latiromitra niveobabelis and other species for comparison. 1-5. Latiromitra niveobabelis new species. Holotype,
USNM 1274447, west of Dry Tortugas, 25°51.104' N, 84°52.278' W to 25°49.867' N, 84°52.400' W, in 1737 m, 46.0 mm.
6. Latiromitra barthelowi (Bartsch, 1942). Holotype, USNM 238444, Cagayan Island, Sulu Sea, Philippines, 905 m, 27.5 mm.
7-8. Latiromitra cryptodon (P. Fischer, 1882). Syntype, off Morocco, 33°29' N, 09° 38' W, 1900 in, 30 mm, photo by M. Caballer
(MNHN). Project : E-Recolnat ANR-ll-INBS-0004. 9. Teramachia chaunax Bayer, 1971. Holotype, USNM 701216 west ol
St. Lucia, 13°45.5' N, 61°05.7' W, 201-589 m, 28 nun, photo by J.F. Quinn, Jr.. 10. Latiromitra aratiuncula (Quinn, 1981).
Holotype, USNM 784594, off Anguilla, 18°26.4' N, 63°63°12.6' W, 430 m, 29 mm (photo credit J. F. Quinn). 11. Latiromitra
meekeana (Dali, 1889). Lectotype, USNM 86970, off Morro Light, Havana, Cuba, 732 m, 15.5 mm, photo by J.F. Quinn, Jr.
Page 92
THE NAUTILUS, Vol. 129, No. 2
whorl, gradually increasing to approximately 21 on pen¬
ultimate whorl and 24 on last whorl; a few, weak second¬
ary axial riblets sporadically appearing on last whorls;
axial sculpture diminishing towards anterior end of last
whorl, completely disappearing at siphonal canal. Spiral
sculpture of two strongly nodulose cords on first whorl;
first cord at suture; second cord just below; sutural
cord later creating a narrow, heavily erenulate channel
(Figure 5); both spiral cords delimiting a constricted
band subsuturally, of approximately 2 mm in width on
last whorl; weak, secondary spiral threads covering the
surface of the shell, crossing over axial ornamentation
creating subtle nodes; approximately 20 stronger spiral
cords developing at anterior end of last whorl. Aperture
narrowly elongate, approximately .19 mm in length; outer
lip thin; parietal wall with three oblique, well-developed
lamelliform plaits; posterior plait strongest. Shell milky-
white, with a faint yellowish coloration showing at, and
restricted to, the subsutural band.
Type Material: Holotype USNM 1274447, length
46.0mm, width 13.7 mm.
Type Locality: West of DrvTortugas, USA (25°51.104/ N,
84°52.278' W to 25°49.867' N, 84o52.400/ W, in 1737 m
(GoMRI-V, station 10).
Distribution: Known only from the type locality.
Habitat: Latiromitra niveobabelis , like other conge¬
ners, is a deep-water species. It was collected in a mud
bottom with rubble. Other notable species dredged in
the same haul were Theta chariessa (Dali, 1889) and
Stellatorna antonia (Dali, 1889).
Etymology: A combination from the Latin adjective
niveo, and the biblical tower of Babel; referring to the
white, tower-like shape of the shell.
Remarks: The tapered profile of the new species is
most similar to the western Pacific Latiromitra barthelowi
(Bartsch, 1942), but the latter has a paueispiral proto¬
conch, weak axial ornamentation after the fourth whorl
and, of the three columellar plaits, the abapical one
is weak to absent (Bouehet and Kantor, 2000: 13). The
new species is also similar to Latiromitra cnjptodon
(P Fischer, 1882) (Figures 6-9), an amphiatlantic species
ranging, in the western Atlantic, from the Bahamas to
Brazil (Rosenberg, 2009). This species differs from
Latiromitra niveobabelis in having a different protoconch
sculpture (see Bouehet and Kantor, 2000: 10, fig. 5K),
in having fewer teleoeoneh whorls, in lacking the well-
developed, constricted subsutural band (compare Figures 5
and 6), in having weaker sutural crenulations, and in having
different coloration. Caribbean specimens of Teratmichia
chaunax F.M. Bayer, 1971 (Figure 9), a junior synonym
of L. cnjptodon , have a more slender profile than tire NE
Atlantic and Brazilian specimens of L. cnjptodon-, how¬
ever, they are otherwise undistinguishahle from the type
material of the latter (Bouehet and Kantor, 2000: 10).
Latiromitra aratiuncula (Quinn, 1981) (Figure 10),
from off Anguilla, lacks the constricted subsutural band,
has coarser shell ornamentation with stronger spiral
cords, and fewer axial ribs. Latiromitra meekeana (Dali,
1889) (Figure 11), the only Latiromitra recorded from
the Gulf of Mexico (Rosenber et al. 2009), has a
paueispiral protoconch of 1.3 whorls, is pale waxen or
brownish in coloration, lacks the constricted subsutrural
band and is almost smooth after the fourth whorl.
Latiromitra costata (Dali, 1890) is much smaller, has
strongly convex whorls, and fewer, stronger axial ribs.
Latiromitra styliola (Dali, 1927), which has tentatively
been placed in Latiromitra by Bouehet and Kantor
(2000) has a protoconch of 1.5 whorls, has 5 teleoeoneh
whorls and reaches 11 mm in length.
ACKNOWLEDGMENTS
My thanks to my colleagues Drs. Darryl Felder and
Suzanne Fredericq, Biology Department at ULL, for
inviting me to join them on the GoMRI project; and
to Ms. Virginie Heros, Museum National d’Histoire
Naturelle, Paris and Dr. Jose II. Leal, editor. The Nautilus ,
for allowing me to reproduce images 6-8 and 9-11
respectively. I would also like to acknowledge Drs. M.G.
Harasewych and Yuri Kantor for reviewing the manuscript
and improving its quality. The GoMRI cruises were con¬
ducted with grants from British Petroleum.
LITERATURE CITED
Bayer, F.M. 1971. New and Unusual Mollusks Collected by
R/V John Elliott Pillsbury and RA7 Gerda in the tropi¬
cal Western Atlantic. Biological results of the University
of Miami Deep-Sea Expeditions. 79. Bulletin of Marine
Science 21: 111-236.
Bouehet, P. and A. Waren. 1985. Revision of the northeast
Atlantic bathyal and abyssal Neogastropoda excluding the
Turridae (Mollusca, Gastropoda). Bollettino Malacologico,
Supplemento 1: 121-296.
Bouehet, P. and Yu. I. Kantor. 2000. The anatomy and system¬
atic^ of Latiromitra a genus of tropical deep-water
Ptychatractinae (Gastropoda: Turbinellidae). The Veliger
43: 1-23.
Cernohorsky, W.O. 1970. Systematics of the families Mitridae
and Volutomitridae. Bulletin of the Auckland Institute and
Museum 8, iv + 190 pp., 18 pis.
Dali, W.H. 1889. Reports on the results of dredgings, under
the supervision of Alexander Agassiz, in the Gulf of
Mexico (1877-78) and in the Caribbean Sea (1879-
80), by the U. S. Coast Survey Steamer “Blake”. Bulle¬
tin of the Museum of Comparative Zoology 18: 1-492,
pis. 10-40.
Dali, W.H. 1890. Scientific results of explorations by the U. S.
Fish Commission Steamer “Albatross”. No. VII. Prelimi¬
nary report on the collection of Mollusca and Brachiopoda
obtained in 1887-88. Proceedings of the United States
National Museum 12(773): 219-362, pis. 5-14.
E.F. Garcia, 2015
Page 93
Dali, W. H. 1927. Small shells from dredgings off the southeast
coast of the United States by the United States Fisheries
Steamer "Albatross” in 1885 and 1886. Proceedings of the
United States National Museum 70(2667): 1-134.
Kantor, Y. 2014. Ptychatractidae Stimpson, 1865. Accessed
through: World Register of Marine Species at http://www
.marinespecies.org/aphia.phppp = taxdetails&id= 41 1810 on
Nov. 1 1 , 2014.
Quinn, J.F., Jr. 1981. A new genus of Turbinellidae (Gastropoda:
Prosobranchia), with the description of a new species from
the Caribbean Sea. The Nautilus 95: 72-77.
Rosenberg, G. 2009. Malacolog 4.1.1: A Database of Western
Atlantic Marine Mollusca. [WWW database (version 4.1.1)]
U RL http://www.malacolog.org/.
Rosenberg, G., F. Moretzsohn, and E.F. Garcia. 2009.
Gastropoda (Mollusca) of the Gulf of Mexico. In: Felder,
D.L. and D. K. Camp (eds.) Gulf of Mexico: Its Origins,
Waters, and Biota. II. Biodiversity. Texas A&M University
Press, College Station, pp. 579-699.
Weaver, C.S. and J.E. duPont. 1970. Living Volutes. Delaware
Museum of Natural History, Greenville, xv + 375 pp.,
79 pis.
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Jacksonville, FL 32210
Dr. Charles Lydeard
Biodiversity and Systematics
Department of Biological Sciences
University of Alabama
Tuscaloosa, AL 35487
Dr. Bruce A. Marshall
Museum of New Zealand
Te Papa Tongarewa
P.O. Box 467
Wellington, NEW ZEALAND
Dr. Paula M. Mikkelsen
Paleontological Research
Institution
1259 Trumansburg Road
Ithaca, NY 14850
Dr. Diarmaid 6 Foighil
Museum of Zoology and Department
of Biology
University of Michigan
Ann Arbor, MI 48109-1079
Dr. Gustav Paulay
Florida Museum of Natural History
University of Florida
Gainesville, FL 32611-2035
Dr. Gary Rosenberg
Department of Mollusks
The Academy of Natural Sciences
1900 Benjamin Franklin Parkway
Philadelphia, PA 19103
Dr. Angel Valdes
Department of Malacology
Natural History Museum
of Los Angeles County
900 Exposition Boulevard
Los Angeles, CA 90007
Dr. Geerat J. Vermeij
Department of Geology
University of California at Davis
Davis, CA 95616
Dr. G. Thomas Watters
Aquatic Ecology Laboratory
1314 Kinnear Road
Columbus, OH 43212-1194
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THE0NAUTILU S
Volume 129, Number 3
September 1, 2015
ISSN 0028-1344
CONTENTS
G. Thomas Watters A revision of the western Atlantic Ocean genus Efflgina with notes
Koen Fraussen on Hesperistemia (Gastropoda: Buccinidae: Pisaniinae) . 95
Angel Valdes On two abyssal species of Scaphandridae G.O. Sars, 1878
James H. McLean (Gastropoda: Cephalaspidea) from the eastern Pacific . 118
Rafael Araujo Conservation of two endangered European freshwater mussels
Carles Feo (Bivalvia: Unionidae): A three-year, semi-natural breeding experiment . 126
Quim Pou
Miquel Campos
Emilio F. Gareia A new species of Glyphostoma (Gastropoda: Clathurellidae)
from the Gulf of Mexico . 136
Research Note
Bruce A. Marshall
Di Tracey
First evidence for deep-sea hot venting or cold seepage
in the Ross Sea (Bivalvia: Vesicomyidae) .
140
THE NAUTILUS 129(3):95-117, 2015
Page 95
A revision of the western Atlantic Ocean genus Engina with notes
on Hesperisternia (Gastropoda: Buccinidae: Pisaniinae)
G. Thomas Watters
Department of Evolution, Ecology, and Organismal Biology
The Ohio State University
Columbus, Ohio 43212 USA
[email protected]
Koen Fraussen
Leuven sestraat 25
B-3200 Aarschot, BELGIUM
[email protected]
ABSTRACT
The western Atlantic Ocean members of the genus Engina Gray,
1839, are reviewed. The following taxa are recognized: E. annae
new species, E. corinnae Crovo, 1974, E. demani de Jong
and Coomans, 1988, E. goncalvesi Coltro, 2005, E. itzamnai
(Watters, 2009), E lignea new species, E. permixta new species,
E turbinelh (Kiener, 1836), and E. williamsae new species.
Distributional notes are also given for Hesperisternia karinae
(Nowell-Usticke, 1959).
INTRODUCTION
The systematics of Engina is confusing as the genus has
a complicated taxonomic history. Early species were
described in the genus Valuta by Linnaeus (1758) and
Wood (1828), in Buccinum by Lamarck (1822), in
Columbella by Sowerby (1832) and Duclos (1840), and
in Purpura by Kiener (1836). Even after the description
of the genus Engina in 1839 species continued to be
described in Buccinum by Kiister (1858), in Pollia by
Dunker (1860), in Tritonidea by E.A. Smith (1884), and
other genera, but especially in Ricinula by Reeve (1846).
In addition, aside from actual buccinid species, members
of other families (Murieidae, Columbellidae, etc.) also
were included in those genera and, for the simple reason
that many of those small shells look similar, it was not
until Pease (1860), Melvill (1893; 1894a; 1984b; 1895)
and Melvill and Standen (1895) that the name Engina
was finally used in publications. Tryon (1883) and Pace
(1902) considered Engina to be a columbellid genus.
Engina was introduced by J.E. Gray in The Zoology of
Captain Beechey’s Voyage. In the Introduction to this
publication, Beech ey (in Gray, 1839: vii) stated that
“I wish I could with sincerity have included with the
above-mentioned [acknowledged] names that of Mr. }. E.
Gray, who undertook to describe the shells, but the pub¬
lication has suffered so much by delay in consequence of
his having been connected with it, that it is a matter of the
greatest regret to me that 1 ever acceded to his offer to
engage in it. . . [This eight year delay] has been occasioned
entirely by Mr. Gray’s failing to furnish his part in spite of
every intercession from myself and odiers.” Beechey con¬
tinued in this vein for another page. Gray had listed two
species in the genus, neither collected during the voyage,
both new but unillustrated, and did not select a type of
die genus. Both species, E. zonata and E. elegan.s , have
been the subject of debate due not only to Gray’s lack of
illustration, but to his apparent gross mis-measurement
of the specimens, giving lengths 10 times greater than
the actual shells, and the vague type localities for
both: “Atlantic Ocean.” Gray subsequendy (1847) picked
E. zonata as the type while misspelling Engina as Enzina
(also misspelled in 1842).
Landau and Vermeij (2012) considered Engina to be a
group of intertidal and shallow subtidal species. How¬
ever, most of the species considered here occur in sig¬
nificantly deeper water; live individuals often occur in
depths > 45 m, some to hundreds of meters. Of the
western Atlantic species only E. turhinella truly occurs
predominately in shallow water. Most species are associ¬
ated with rocky reefs or coral debris, whereas some occur
in caves. As with many other gastropod genera, Engina
harbors a number of species that are common or even
abundant whereas several species are quite rare and
rarely represented in collections.
The type species of Engina , E. turhinella , is a member
of the shallow water fauna and therefore shows the shell
characteristics that are representative of Engina species
from that fauna. Most, or all, shallow water Engina spe¬
cies bear the characteristic radially oriented lirae on the
columella that are seen in the type species. Nevertheless
many Engina live in deeper water and differ slightly or
even radically from the shallow water morphs. In deeper
water (ea. > 45 m) species may lack these radially orien¬
tated lirae on the parietal part. The shallow and deep
water moqihs are not well separated and it remains a
question of whedier this is the result of variability (the
opinion followed in the present paper) or rather that our
current concept of Engina is based on a heterogeneous
group (die result of the presence of unrecognized genera).
Page 96
THE NAUTILUS, Vol. 129, No. 3
In the present paper we regard Engina in a rather conser¬
vative way, grouping together the shallow and deep-water
species. Further study may indicate that additional genera
are required.
The type species of Engina , together with the spe¬
cies discussed in the present paper, are members of the
Atlantic Ocean fauna. A much larger number of Engina
species is known from the Indo-West Pacific and eastern
Pacific oceans. We believe that both the Atlantic and Indo-
West Pacific species belong to the same genus, as so many
other genera do, and we see no reason to split Engina
into distinct groups based on geographical distribution.
Apart from a higher biodiversity in the Indian and
Pacific oceans, the morphological variability within the
genus is also higher there than in the Atlantic Ocean.
Slender, fusiform shells are a frequent shape in the Indo-
West Pacific Ocean whereas none are known to us from
the Atlantic Ocean. Those slender species have a different
appearance than do broad ones and tend to live in deeper
water; nevertheless we regard them as included within
the variability of the genus. “Typical” species have radi¬
ating eolumellar riblets. In some species those are well
expressed and striking but other species show this sculp¬
ture only under magnification.
Many Western Atlantic Ocean species have eastern
Pacific Ocean cognates. Atlantic taxa such as E. turbinella
and E. permixta closely resemble Pacific species such as
E. mantensis Bartsch, 1928, E. niaura (Sowerby, 1832),
and E. tabogaensis Bartsch, 1931. The Pacific £. pulchra
(Reeve, 1846) is particularly close to the Atlantic £. clemani.
But taxa such as the Pacific E. fusifomiis Steams, 1894,
and the Atlantic E. g oncalvesi do not have similar cog¬
nates. Vermeij (2006) recognized the close relationship
between the Pacific Hesperistemia jugosa (Adams, 1852),
previously considered an Engina (Keen, 1971), H. lauta
(Reeve, 1846), previously considered a Cantharus (Keen,
1971), and the Atlantic //. karinae (reviewed here).
A fossil record for Engina was nearly lacking until
Landau and Vermeij (2012) reviewed the western Atlantic
taxa. They identified Engina cantaurana Landau and
Vermeij (2012) from the early Miocene of Venezuela as
the oldest known member of the genus but did not believe
it was closely related to any Recent species. They found
that Engina latior Landau and Vermeij (2012) from the
early Pliocene of the Dominican Republic, £. nwinensis
Landau and Vermeij (2012) from the early Pleistocene of
Panama, and £. floridana Olsson and Harbison, 1953,
from the early Pleistocene of Florida, were more similar
to Recent species. Engina nwinensis , in particular, is
very similar to the two rare, deep-water Yucatan taxa
discussed here: E. itzamnai and E. lignea. The extant
E. turbinella was recognized from the middle Miocene
of Panama (Woodring, 1973) and the late Pleistocene
of the Dominican Republic (Landau and Vermeij, 2012).
Spiral and axial sculpture are features with a remark¬
ably low degree of infraspecific variability within the
genus Engina. Variability in pattern and color, to the
contrary, is remarkably high for some Engina. While
most species are moderately uniform in these character¬
istics, without displaying much variation in pattern or
color, a number of other species are strikingly variable.
This was first discussed concerning a Philippine species
(Fraussen and Chino, 2012); in the present paper we
show a similar level lof variability in at least two species.
As remarked by Cemohorsky (1975) and Faber (2007),
Engina cumingiana Melvill, 1895, from “St. Thomas” appears
to be a mislabeled, but as yet unidentified, species. It is not
similar to any of the species covered here. Engina willemsae
de Jong and Coomans (1988) was transferred to Anna by
Watters (2009) and to Ameranna by Landau and Vermeij
(2012). Engina slootsi de Jong and Coomans (1988) is
Habromorula biconica (Blainville, 1832) (Murieidae) from
the Pacific Ocean ( fide Houart, 1994).
MATERIALS AND METHODS
Length is measured from the tip of the apex to the end of
the siphonal canal. Width is measured as the maximum
dimension in a plane with facing the aperture. Spiral sculp¬
ture is counted from the suture to the end of die siphonal
canal. Descriptions of apertural features are based on
the terminology adopted by Landau and Vermeij (2012).
Lirae counts within the outer lip may include bifurcating
lirations. The number of whorls was determined using
the 1 D method of Van Osselaer (1999). Locality infor¬
mation, aside from type locality designations, may be
amplified from the original label for clarification. Given
the generalized nature of most label information, no
attempt has been made to georeference sites that did not
originally include coordinates. Dimensions in captions
refer to shell length. Numbers in ( ) following catalog
numbers indicate the number of specimens in the lot.
Abbreviations used in die text are: AMNH: American
Museum of Natural History, New York City, New York,
USA; BMSM: Bailey- Matthews National Shell Museum,
Sanibel, Florida, USA; NHMUK: Natural History Museum,
London, UK; EFG: Collection of Emilio F. Garcia,
Lafayette, Louisiana, USA; GTW: Collection of G. Thomas
Watters, Columbus, Ohio, USA; HGL: Collection of Harry
G. Lee, Jacksonville, Florida, USA; KF: Collection of Koen
Fraussen, Aarschot, Belgium; MCZ: Museum of Com¬
parative Zoology, Cambridge, Massachusetts, USA; MW:
Collection of Margaret Williams, Tallevast, Florida, USA;
OSUM: Ohio State University Museum of Biological Diver¬
sity, Columbus, Ohio, USA; PRI: Paleontology Research
Institute, Ithaca, New York, USA; UF: Florida Museum
of Natural History, Gainesville, Florida, USA; ZMA:
Zoologisch Museum, Amsterdam, The Netherlands.
SYSTEMATICS
Family Buceinidae Rafinesque, 1815
Subfamily Pisaniinae Gray, 1857
Genus Engina Gray, 1839
Engina Gray, 1839: 112-113.
G.T. Watters and K. Fraussen, 2015
Page 97
Type Species: Engina zonata Gray, 1839, by subse¬
quent designation of Gray (1847) [= Purpura turhinella
Kiener, 1835, see Orr (1962)].
Discussion: The morphological limits and unique fea¬
tures of Engina have not been easy to define. Cernohorsky
(1975: 176) characterized the genus by “die radially ori¬
ented lirae [which] are found only in species of Engina,”
a feature that Landau and Vermeij (2012) nevertheless
noted was not present in all Engina. Landau and Vermeij
(2012: 121) defined the genus by “a shell with a narrow,
strongly denticulate aperture and the presence of distinct
columellar folds” but admitted that “not all species. . .
conform to diis characterization.” Definitive differences
between Engina and Hesperistemia appear to be par¬
ticularly ambiguous. Vermeij (2009) noted that some
Hesperistemia had shell sculpture and apertural features
that were convergent with Engina. Watters (2009) placed
E. janowskyi and his species itzamnai in Hesperistemia ;
Landau and Vermeij (2012) reallocated them to Engina.
That reallocation is followed here and we confirm the
importance of the presence of internal lirae that extend
far into the aperture as an important feature to distin¬
guish Hesperistemia from Engina.
Adding to the problems of identifying the western
Atlantic species is the great variability seen in concholog-
ical features. Coloration, sculpture, degree of elongation,
and apertural features are highly plastic. Given a small
number of specimens it is easy to conclude that numer¬
ous taxa are involved. However, when larger lots are
studied it is apparent that there are intergrades that
unite such disparate forms.
The species recognized in this review are presented in
alphabetical order, including the type species of Engina,
E. turhinella (Kiener, 1835).
Engina annae new species
(Figures 1-4, 13)
Engina species. — Lee, 2009: 108, fig. 518a.
Description (based on 2 adult specimens): Larger
adult specimen seen, 15.6 mm in length (holotype);
smaller specimen seen, 12.9 mm in length; holotype
15.6 mm length x 8.9 mm width. Shell biconic; spire ca.
50% total length. Protoconch minute, of 1.5 smooth, pale
tan whorls. Teleoeonch of 5.5 whorls, demarcated from
protoconch. Teleoeonch sculpture of 9-10 primary
rounded spiral cords, between the suture and the ante¬
rior limit of the siphonal canal, three grouped at periph¬
ery; concave or flat, wide subsutural slope with a single
thin primary cord. Secondary microscopic spiral threads
between primary spiral sculpture. Primary axial sculp¬
ture of broad, low, rounded ribs, very weak on subsutural
slope; 10 ribs on penultimate whorl, 10 on final whorl.
Extremely fine secondary wrinkles occur between axial
ribs. Terminal varix weak. Aperture oval, elongate. Anal
canal bounded by single parietal denticle on columella
and single anal denticle on outer lip. Outer lip with
5-7 denticles, decreasing in size from posterior to ante¬
rior, the posterior-most two with a tendency to be weakly
fused. The thin rib that projects deep into the aperture
and delimits the siphonal canal found in other species is
weak. Five radial lirae present at posterior end, barely
distinct from columellar folds. Anterior to these are 6-8
columellar folds irregular, some lirate, others pustulose.
One or more denticles extending deep into aperture,
forming an interior ledge to columella. Parietal lip erect
for most of its length. Siphonal canal short, open. Color
gray with a tan subsutural band; axial ribs dark brown,
often with a white adapertural face forming a series of
dark and light spots; there is a continuous, white spiral
band just below the periphery. Aperture tan with white
teeth. Operculum, radula, and anatomy unknown.
Holotype: UF 479323 (ex Mark Johnson coll. 0142).
Type Locality: 33 in., 48 km off Masonboro Inlet,
Wrightsville Beach, New Hanover Co., North Carolina,
USA.
Paratype(s): Charlotte Thorpe coll. (1), Amber jack
Hole, E of Mayport, Duval Co., Florida, USA.
Distribution: Only known from offshore of southern¬
most North Carolina and northeast-most Florida. We
assume it occurs off the intervening South Carolina and
Georgia coasts as well.
Habitat: The holotype was a crabbed specimen from
33 m, collected on sandy rubble near rocky outcrops
with sponges.
Shell Variation: The known specimens are remark¬
ably uniform in sculpture and coloration.
Etymology: Named for Ann Johnson, mother of the
collector of the holotype, Mark Johnson.
Comparison with Other Species: The peculiar col¬
oration of dark brown and white spots on a gray back¬
ground has not been found in any other western Atlantic
species. See Table 1 for a comparison with other species.
Discussion: Tins extremely rare species is the northern¬
most member of Engina in the western Atlantic Ocean.
Like most of the species discussed here, it is an offshore
species seldom encountered. It was first recognized as
distinct by Lee (2009).
Engina corinnae Crovo, 1974
(Figures 14-33, 34)
Engina corinnae Crovo, 1974: 30, figs. 1-3; Abbott, 1974:
217, fig. 2398; Ode, 1983: 60; Kaicher, 1986: card
4405; Landau and Vermeij, 2012: 122, 123, 126;
Petueh, 2013: 202.
Engina corinnae of authors, non Crovo, 1971. — Lee,
2009: 108, fig. 518.
Description (based on 38 adult specimens): Larg¬
est adult specimen, 14.8 mm in length; smallest speci¬
men, 9.2 mm in length. Shell biconic; spire ca. 50% total
Page 98
THE NAUTILUS, Vol. 129, No. 3
Figures 1-12. Engina species. 1—4. Engina annae new species. 1-2. Holotype, UF 47923, 15.7 mm. 3-4. Paratype Charlotte Thorpe
coll., Amberjack Hole, E of Mayport, Duval Co., Florida, 12.9 mm. 5-9. Engina goncalvesi Coltro, 2005. 5-6. GTW 12447a, 40-50 m,
in cave, off Arraial do Cabo, Rio de Janeiro State, Brazil, 11.3 mm. 7. GTW 12447b, 150-160 m, off Cabo Frio, Rio de Janeiro State,
Brazil, 13.9 mm. 8. HGL, 40-44 m, in cave, off Arraial do Cabo, Rio de Janeiro State, Brazil, 9.5 mm. 9. GTW 12479a, 40-50 m, in cave,
off Arraial do Cabo, Rio de Janeiro State, Brazil, 1 1.3 mm. 10-12. Engina lignea new species. 10-11. Holotype, UF 479325, 14.3 mm.
12. Paratype, BMSM 76001, 50-55 m, 146 km WSW of Arrecife Alaeran, Campeche Bank, Campeche State, Mexico, 1 1.5 mm.
length. Protoconch minute, of 1.5 smooth whorls, white
with tan patches or bands. Teleoeoneh of 4.75-5.25
whorls, demarcated from protoconch. Teleoeoneh sculp¬
ture of 9-12 (mode=ll) primary sharp spiral cords
between the suture and the anterior limit of the siphon al
canal, weakest on concave or flat subsutural slope; else¬
where of uniform strength except that spiral cords on
siphonal canal are slightly stronger. Secondary microscopic
G.T. Watters and K. Fraussen, 2015
Page 99
Figure 13. Distribution of Engina annae new species (©),
Engina goncalvesi Coltro, 2005 (•), Engina lignea new species
( ♦ ), and Engina williamsue new species( * ).
spiral threads between primary spiral sculpture. Primary
axial sculpture of broad, low ribs, absent from subsutural
slope; 7-10 (mode=9) ribs on penultimate whorl, 6-13
(mode=10) on last whorl, most pronounced at periph¬
ery. Coarse secondary wrinkles occur between axial ribs
which may be lamellate. Terminal varix weak. Aperture
oval, elongate. Anal canal bounded by single parietal den¬
ticle on columella and single anal denticle on outer lip.
Outer lip with 6—8 (mode=6) denticles. The thin rib that
projects deep into the aperture and delimits the siphonal
canal found in other species is weak but always present.
2-5 (mode=3) radial lirae present at posterior end, weak
and irregular, not distinct from columellar folds. Anterior
to these are 4-7 (mode=6) semi-lirate columellar folds
irregular, often limited to edge of parietal wall, weakest
or absent on middle of columella. First or second ante¬
rior denticle often extending deep into aperture forming
an interior ledge to columella. Parietal lip erect for
most of its length. Siphonal canal short, open. Color
white or pale tan with brown blotches, darkest between
axial ribs, particularly on die spire, with a wade, white sub¬
peripheral band; usually two fine, dark threads poste¬
rior to this band on the final whorl. Aperture white,
faint tan, or mauve, darkest at anterior end. Operculum
rounded, leaf-shaped, yellow with brown central radius.
Radula and anatomy unknown.
Holotype: MCZ 277496.
Type Locality: Off Boynton Beach, Palm Beach
County, Florida, USA, 30° 00' N, -79° 57'30" W.
Depth unknown.
Paratype(s): MCZ 277497 (unstated number of speci¬
mens); PRI 28273 (1); all from the type locality.
Other Material Examined (124 specimens): East
Florida: MW (1), 27 m, off’ eastern Florida; MW (1), .34 m,
off Mayport, Duval Co.; MW (1), HGL (6), 30-46 m, off
St. Augustine, St. Johns Co.; UF 35241 (1), 18 m, off
Boynton Beach, Palm Beach Co.; UF 128074 (66), 27 m,
off Boynton Beach, Palm Beach Co.; UF 128051 (3),
27-38 m, off Boynton Beach, Palm Beach Co.; EFG
9553 (2), 38-43 m, off Boynton Beach, Palm Beach Co.;
PIGL (1), Boynton Beach, Palm Beach Co.; UF 352841
(1), 18 m, off Boynton Beach, Palm Beach Co., Florida;
UF 122778 (7), 107 m, off Boynton Inlet, Palm Beach Co.;
UF 126244 (1), 26 m, off Boynton Inlet, Palm Beach
Co.; UF 228793 (6), 73 m, off Palm Beach, Palm Beach
Co.; UF 127142 (1), off Palm Beach, Palm Beach Co.;
UF 229058 (1), 55-73 in, off Palm Beach, Palm Beach
Co.; UF 262537 (2), 91-152 m, off Palm Beach, Palm
Beach Co.; UF 126241 (7), 37 m, off Palm Beach, Palm
Beach Co.; UF 228990 (2), 37-46 in, off Palm Beach
Inlet, Palm Beach Co.; UF 250037 (2), 37-55 m, Dodge
Estate to inlet. Palm Beach Co.; EFG 13895 (1), 30 m, off
West Palm Beach, Palm Beach Co.; HGL (3), 183-213 m,
off Looe Key, Monroe Co. West Florida: EFG 28347 (1),
61 m, off Egmont Key, Hillsborough Co.; MW (1), HGL
(5), 27 m, off Panama City, Bay Co.; MW (2), 6.4 km off
Table 1. Shell characteristics of Engina and Hesperistemia. Numbers in ( ) are modes except for length, which are averages;
* - no inode.
Page 100
THE NAUTILUS, Vol. 129, No. 3
Figures 14-33. Engina corinnae Crovo, 1974. 14-15. Holotype, MCZ 277496, photo courtesy of Jennifer W. Lenihan, © Museum
of Comparative Zoology, Harvard Univ., 13.0 mm. 16. UF 128074, 27 m, off Boynton Beach, Palm Beach Co., Florida, 12.4 mm.
17. EFG 9553, 38-43 m, off Boynton Beach, Palm Beach Co., Florida, 9.9 mm. 18. UF 228793, 73 m, off Palm Beach, Palm Beach
Co., Florida, 9.5 mm. 19. UF 122778, 107 m, off Boynton Inlet, Palm Beach Co., Florida, 9.3 mm. 20. UF 122778, 107 in, off Boynton
Inlet, Palm Beach Co., Florida, 9.3 mm. 21. UF 126241, 36.6 m, off Palm Beach, Palm Beach Co., Florida, 1 1.0 mm. 22. MW, 30-46 m,
off St. Augustine, St. Johns Co., Florida, 12.7 mm. 23. HGL, 183-213 in, off Looe Key, Monroe Co., Florida, 11.5 mm.
24. HGL, 30-46 in, off St. Augustine, St. Johns Co., Florida, 12.2 mm. 25. MW, 6.4 kin off Panama City, Bay Co., Florida, 13.2 mm.
26. HGL, 30-46 m, off St. Augustine, St. Johns Co., Florida, 10.5 mm. 27. EFG 13895, 30 m, off West Palm Beach, Palm Beach
Co., Florida, 14.2 mm. 28. EFG 27567, 39 m, 54 km SSW of Panama City, Bay Co., Florida, 14.7 mm. 29. MW, 27 m, off Panama
City, Bay Co., Florida, 10.5 mm. 30. MW, 31 m, off NE Yucatan, Mexico, 14.8 mm. 31. EFG 28347, 61 in, off Egmont Key,
Hillsborough Co., Florida, 12.6 mm. 32. MW, 34 m, off Mayport, St. Johns Co., Florida, 12.7 mm. 33. ?Engina corinnae. UF 352841,
18 m, off Boynton Beach, Palm Beach Co., Florida, 12.4 mm.
G.T. Watters and K. Fraussen, 2015
Page 101
Figure 34. Distribution of Engina corinnae Crovo, 1974 (©)
and Engina demani de Jong and Coomans, 1988 (•).
Panama City, Bay Co.; EFG 27567 (1), 39 m, 54 km SSW
of Panama City/ Bay Co., 29° 43.32' N, -85° 54.85' W.
Louisiana: EFG 23426 (1), 57-65 m, Ewing Bank, 105 km
off Louisiana coast, 28° 06.07' N, —91° 02.42' W. Mexico:
MW (3), 31 m, off NE Yucatan.
Lee (2009) also lists Xalvis Island, St. Johns Co., Florida.
Distribution: Western Atlantic Ocean, offshore, east¬
ern Florida from St. Johns County to the Florida Keys.
Gulf of Mexico from Hillsborough County, Florida, west
to Louisiana, and off Yucatan Peninsula. The E. corinnae
recorded from the East Flower Garden banks by Ode
(1983) may be this species but we have not seen the
specimen. This species co-occurs with the equally rare
E. williamsae at Egmont Key off Tampa Bay, west¬
ern Florida.
Habitat: Empty shells have been found from 18-122 m
on rubble bottom. Live specimens have been recorded
from 30-73 m. Specimens are locally common and are
occasionally found on Spondylus. Most specimens are
taken as by-catch by shrimp and scallop trawlers. A single
label reads “rock, sand reef,” the only report of a habitat.
Specimens are frequently covered by calcareous algae.
Variation in Specimens: Specimens vary in degree of
coloration, from nearly all white (holotype) to having prom¬
inent brown patches. The intervarical brown banding
varies from diffuse patches to distinctly pigmented spi¬
ral cords, but is always present. Specimens also vary
in the degree of elongation, ranging from compact to
rather fusiform.
Etymology: Named for Corinne E. Edwards (1905-
1989), an “ardent collector of marine life as well as an
educator in popular subjects on natural history” (Crovo,
1974: 30).
Comparison with Other Species: Engina corinnae
superficially resembles E. williamsae new species. Engina
corinnae has more prominent spiral sculpture over
the entire shell, including the sub-sutural band, which
is nearly smooth in E. williamsae . Engina corinnae
is also less solid, has a shorter spire, and averages a
slightly smaller size. The widespread E. turbinella
differs in its (usually) much darker coloration with
white spots as contrasted to the lighter coloration with
dark spots seen in E. corinnae, as well as its dark
aperture with white teeth. See Table 1 for a compari¬
son with other species.
Discussion: Engina corinnae is an offshore species
that varies greatly in the degree of elongation and color¬
ation. The holotype is not typical of the specimens usu¬
ally seen. It is small, compact, nearly uniformly white,
and has fewer axial ribs than average specimens.
Engina demani de Jong and Coomans, 1988
(Figures 34-49)
Engina demani de Jong and Coomans, 1988: 83, pi. 38,
fig. 451; Faber, 2007: 74, figs. 1-3; Watters, 2009:
271.
Engina janowskyi Coltro, 2005: 1, pi. A, figs. 1-5; Faber,
2007: 74, figs. 6, 7; Dornellas and Simone, 2011: 20;
Landau and Vermeij, 2012: 123, 125, 126.
Hesperistemia janowskyi (Coltro, 2005). — Watters,
2009: 271.
Engina demanorum [.sic] de Jong and Coomans, 1988. —
Landau and Vermeij, 2012: 123, 125, 126 [unjusti¬
fied emendation, see “etymology” below].
Description (based on 20 adult specimens): Larg¬
est adult specimen, 15.5 mm in length; smallest speci¬
men, 10.7 mm in length. Shell biconic to fusiform; spire
ca. 50% total length. Protoconch small, conical, of 1.5-
1.75 smooth, white or brown banded whorls. Teleoeoneh
of 5.5 whorls, weakly demarcated from protoconch.
Teleoeoneh sculpture of 10-12 (mode=12) evenly
spaced, rounded, spiral cords between the suture and
the anterior limit of the siphonal canal. Subsutural slope
wide, flat, with single primary thread, well-developed in
some specimens, very weak in others. Between these
cords are ca. 10 fine, regular threads. Axial sculpture of
widely spaced, wide, primary angular ribs; 7 (rarely 6 or
8) on the final whorl, 7 (rarely 6 or 8) on penultimate
whorl, most prominent on periphery, with numerous
secondary fine, axial threads in between primary ribs; in
well-preserved specimens these threads are developed
into oblique lamella, giving a scalloped appearance to
the sculpture. Terminal varix well-developed, angular.
Aperture oval. Parietal wall erect on anterior hall. Anal
canal bounded by single parietal denticle on columella
and single anal denticle on outer lip. Outer lip with 5
(rarely 6) small denticles. The thin rib that projects deep
Page 102
THE NAUTILUS, Vol. 129, No. 3
Figures 35-49. Engina demani de Jong and Coomans, 1988. 35. Holotype, Engina demani de Jong and Coomans, 1988,
11.0 mm, photo courtesy of Marien Faber. 36. Holotype, Engina janotvskyi Coltro, 2005, 13.2 mm, reproduced from Coltro
(2005). 37. HGL, 183 m, 3.2 km W of Sandy Lane Bay, Barbados, 11.2 mm. 38. GTW 13795e, 4 m, Ilha Escalvada, Espfrito
Santo State, Brazil, 13.8 nun. 39. GTW 13795e, 4 m, Ilha Escalvada, Espfrito Santo State, Brazil, 15.2 nun. 40. HGL, 40^45 m,
off Buzios, Sao Paulo State, Brazil, 10.7 mm. 41. HGL, shallow water, Cartagena, Colombia, 13.6 mm. 42. GTW 13795a, 40-45 m,
Ilha de Cabo Frio, Rio de Janeiro State, Brazil, 13.5 mm. 43. GTW 12026d, 3 m, off Guarapari, Espfrito Santo State, Brazil, 15.0 mm.
44. GTW 15579a, French Guiana, 13.1 mm. 45. GTW 13795b, 4 m, Ilha Escalvada, Espfrito Santo State, Brazil, 15.3 mm. 46. GTW
11645a, 15-25 m, off Guarapari, Espfrito Santo State, Brazil, 12.3 mm. 47. GTW 13795e, 4 m, Ilha Escalvada, Espfrito Santo State,
Brazil, 12.3 mm. 48. GTW 13795b, 4 m, Ilha Escalvada, Espfrito Santo State, Brazil, 11.4 mm. 49. GTW 13795e, off Guarapari,
Espfrito Santo State, Brazil, 14.9 mm.
G.T. Watters and K. Fraussen, 2015
Page 103
into the aperture and delimits the siphonal canal found
in other species is absent or nearly so. Radial lirae 1^1
(mode=3) (may be bi- or trifurcate) at posterior end,
weak and irregular, not very distinct from columellar
folds. Anterior to these are columellar folds (4-8,
mode=6) weak, irregular, often limited to edge of parie¬
tal wall. Swelling on interior ledge of columella not pro¬
nounced. Siphonal canal moderately short, open. Base
color variable: dark brown, tan, or yellow, axial ribs usu¬
ally darker, always with a continuous pale band just ante¬
rior to the periphery. The abapertural side of the axial
ribs is usually dark-colored and the adaperture side is
usually pale. Aperture white to tan, the siphonal canal
rimmed in brown. Operculum oval, yellow-brown, with a
subterminal nucleus. Radula and anatomy unknown.
Holotype: Engina demani de Jong and Coomans,
1988: ZMA 3.87.084; Engina janowskyi Coltro, 2005:
M useu de Zoologia da Universidade de Sao Paulo
37178, but apparently lost ( fide D. Cavallari, Jan. 2015).
Type Locality: Engina demani de Jong and Coomans,
1988, Aruba, harbour [Netherlands Antilles], Engina
janowskyi Coltro, 2005, off Guarapari, Espirito Santo
State, Brazil.
Paratype(s): Engina demani de Jong and Coomans,
1988: Faber (2007) illustrated a “paratype” from the
Frere Frederieus Verberne collection. Although de Jong
and Coomans (1988) mentioned additional examples in
the Verberne and de Man collections, they are not
explicitly referred to as types and it is not certain
whether these specimens are valid paratypes. Engina
janowskyi Coltro, 2005: Museu Oeeanografico Eliezer
Rios da Funfayao Universidade de Rio Grande 43853;
Museu Nacional da Universidade Federal do Rio de
Janeiro, unnumbered; PM. Santos Costa coll.; R.
Janowsky coll, [all single specimens]. The localities of
the paratypes were not given but are presumed to be
from the type locality.
Other Material Examined (28 specimens): Barbados:
HGL (1), 183 m, 3.2 km W of Sandy Lane Bay. Colombia:
HGL (1), shallow water, Cartagena. French Guiana: GTW
15579a (1). BrazilL GTW 11645a (3), HGL (1), 15-25 m,
off Guarapari, Espirito Santo State; GTW 13795c (1),
25 m, off Guarapari, Espirito Santo State; GTW 12026c
(2), 12026d (4), 3 m, off Guarapari, Espirito Santo State;
HGL (1), 18 m, off Anchieta, Espirito Santo State; GTW
13795b (3), 13795e (5), 4 m, Ilha Escalvada, Espirito
Santo State; GTW 13795a (1), 40-45 m, Ilha de Cabo
Frio, Rio de Janeiro State; HGL (1), 34-45 m, off Buzios,
Sao Paulo State; HGL (3), 40-45 m, off Buzios, Sao
Paulo State.
Distribution: Apparently widely distributed across
the southern Caribbean and down the Brazilian coast to
Sao Paulo State, but this is based on few records with
large gaps.
Habitat: Empty shells are found at depths between
3-183 m; some live specimens have been collected in
caves at 40-45 m, but the majority of specimens exam¬
ined were found on rubble or under rocks from 4 m.
Variation in Specimens: Shell base color varies from
nearly pure w'hite to nearly all dark brown. Elongation
varies from compact, solid shells (typically from Brazil,
Figures 38, 43) to relatively high-spired, narrow shells
(French Guiana, Figure 44). Despite this great varia¬
tion in color and elongation of the shell, the sculpture
is remarkably uniform in the number of axial and spi¬
ral elements.
Etymology: Engina demani de Jong and Coomans,
1988, named for Ad and Gon de Man, shell collectors
in Aruba. Thus the correct orthography should be
demanorum. However, a name change reflecting this
would be an unjustified emendation. Engina janowskyi
Coltro, 2005, named for Robert H. Janowsky, owner of
MdM Shell Books in Wellington, Florida, USA.
Comparison with Other Species: Some color forms
of E. permixta are similar to E. demani, with which it
may be sympatric. Engina permixta has more axial ribs
on both the final whorl (7-10) and the penultimate whorl
(8-10) than does E. demani (6-8 for both) and has fewer
spiral cords on the final whorl (7-10 vs. 10-12). Addi¬
tionally, the deep ridge bounding the siphonal canal on
the inside of the outer lip is present in E. permixta but
absent or very weak in E. demani. See Table 1 for a com¬
parison with other species.
Discussion: This is a very variable species in colora¬
tion, in degree of elongation, and in the strength of the
sculpture. Engina demani is based on short, compact
specimens. Engina janowskyi is based on elongate forms.
Without seeing intermediates, the two forms would
clearly seem to represent two taxa. However, there is no
clear cut delineation between the two forms in either
color or elongation and the two taxa are synonymous.
This possible synonymy was first suggested by Faber
(2007), who remarked that E. demani may be the shallow
water form and E. janowskyi the deeper water form.
However, our records indicate that there is great varia¬
tion within each population regardless of depth and both
“forms” and intergrades co-occur together.
Engina goncalvesi Coltro, 2005
(Figures 5-9, 13)
Engina goncalvesi Coltro, 2005: 1-2, pi. B, figs. 1-11;
Faber, 2007: 74 [in synonymy of Bailya milleri
Nowell-Usticke, 1959]; Watters, 2009: 270-271, figs.
190, 191; Dornellas and Simone, 2011: 17; Landau
and Vermeij, 2012: 123.
Description (based on 4 adult specimens): Largest
adult specimen, 14.2 mm in lengtli; smallest specimen,
9.5 mm in length. Shell fusiform; spire ea. 50% total
length. Protoconch small, of 1.5 smooth, brown whorls
Page 104
THE NAUTILUS, Vol. 129, No. 3
with pale peripheral band. Teleoconch of 5 whorls,
abruptly arising from protoconch. Teleoconch sculpture
of 12-15 (mode=12) primary flattened, spiral threads on
final whorl between the suture and the anterior limit of
the siphonal canal, with numerous intercalated second¬
ary and tertiary threads. Spiral cords on siphonal canal
slightly stronger. Axial sculpture of broad, low ribs; ea. 20
primary ribs on penultimate whorl, obsolete on most
specimens by last whorl. Intersections of axial and spiral
sculpture with weak, elongated nodules. Terminal varix
well-developed, flaring, moderately narrow. Aperture
oval. Anal canal bounded by single parietal denticle on
columella and single anal denticle on outer lip. Outer lip
with 6-7 weak, irregular denticles. The thin rib that
projects deep into the aperture and delimits the siphonal
canal found in other species is absent. Single (or bifur¬
cating) radial lira at posterior end, very weak and irregu¬
lar, not very distinct from columellar folds. Anterior to
this lira are columellar folds (3—5), usually indistinct,
irregular, often limited to edge of parietal wall. Swelling
on interior ledge of columella distinct, white. Parietal lip
erect for most of its length. Siphonal canal short, open.
Color brown with wide, pale tan spiral band at sub¬
periphery, primary spiral cords often darker. Aperture
with brownish-purple tinge. Operculum leaf-shaped,
golden-tan, with anterior terminal nucleus. Radula and
anatomy unknown.
Holotype: Stated to be in Museu de Zoologia da
Universidade de Sao Paulo, 37179, and listed as such in
Domellas and Simone (2011), but not found ( fide L. R. L.
Simone, pers. comm., 2008).
Type Locality: Off Cabo Frio, Rio de Janeiro State,
Brazil. Depth unknown.
Paratype(s): Museu Qceanografico Eliezer Rios da
Fundayao Universidade de Rio Grande 43854 (1); Museu
Nacional da Universidade Federal do Rio de Janeiro,
unnumbered (2); PM. Santos Costa coll. (1). The localities
of the paratypes were not given but are presumed to be
from the type locality.
Other Material Examined (4 specimens): Brazil.
GTW 10479a (1), 12477a (1), 40-50 m, in cave, off Arraial
do Cabo, Rio de Janeiro State; HGL (1), 40-44 m, in cave,
off Arraial do Cabo, Rio de Janeiro State; GTW 12477b
(1), 150-160 m, off Cabo Frio, Rio de Janeiro State.
Distribution: Brazil, between Cabo Frio, Rio de Janeiro
State, and Ilhabela, Sao Paulo State.
Habitat: Empty shells are found between depths of
25-160 m; live individuals from 25—45 m under rocks,
sometimes in caves.
Variation in Specimens: The few specimens we have
seen are very uniform in all characteristics.
Etymology: Named for Paulo Cesar Pinto Gongalves,
who first collected the species.
Comparison with Other Species: The near lack of
axial sculpture on the final whorl and reduced colu¬
mellar dentition set this species apart from all other
western Atlantic Engina. See Table 1 for a comparison
with other species.
Discussion: This species is placed in Engina with
some reservation. The parietal shield lirae and columel¬
lar folds are nearly absent in some specimens and only
barely expressed in the remainder.
Engina itzamnai (Watters, 2009)
(Figures 50-57, 68)
Hesperistemia itzamnai Watters, 2009: 271, figs. 192-
195, 264.
Engina itzamnai (Watters, 2009). — Landau and Vermeil,
2012: 123.
Engina dicksoni Petuch, 2013: 72, 192, 202, fig. 5.6E.
Description (based on 4 specimens): Largest adult
specimen, 17.8 mm in length (holotype); s null lest adult
specimen, 17.4 mm in length. Shell fusiform; spire ca.
50% total length. Shell relatively thin. Protoconch small,
conical, of 1.5 smooth, white whorls with tan blotches.
Teleoconch of 5.5 whorls, strongly demarcated from
protoconch. Teleoconch sculpture of 11-13 rounded,
widely separated primary spiral cords between the suture
and the anterior limit of the siphonal canal, with numerous
intercalated secondary threads. Subsutural slope wide,
flat, with single primary thread. Spiral cords on siphonal
canal slightly stronger. Axial sculpture of widely spaced,
rounded primary ribs; 8-10 on penultimate whorl, 8-10
on final whorl, with numerous secondary axial threads.
Intersections of axial and spiral sculpture with strong,
elongated nodules, strongest at periphery. Terminal varix
weakly developed, somewhat constricted, narrow. Aper¬
ture oval, outer lip with 4 medial teeth. Anal canal deeply
indented between two teeth; columellar tooth bifid. Pari¬
etal wall erect with 7 weak, lirate teeth. Siphonal canal
moderately long, open. Color white with orangish-tan
interaxial spaces cut by a white subperipheral narrow
band; the spaces form broken flammulations below this
band. Aperture white. Operculum, radula, and anat¬
omy unknown.
Holotype: Hesperistemia itzamnai Watters, 2009, UF
170226. Engina dicksoni Petuch, 2013, UF 328420.
Type Locality: Hesperistemia itzamnai Watters, 2009:
100 fms. [91 m], NE of Contoy Light, Isla Contoy,
Quintana Roo State, Mexico: Engina dicksoni Petuch,
2013: 35 m depth on Campeche Bank, off Puerto Progresso,
Yucatan State, Mexico.
Paratype(s): Hesperistemia itzamnai Watters, 2009, UF
425817 (1), from type locality. Engina dicksoni Petuch,
2013, Petuch coll. (1), from type locality.
Other Material Examined (2 specimens): Mexico:
EFG 25800 (1), 52-53 m, 64 km NNW of Cayos
G.T. Watters and K. Fraussen, 2015
Page 105
Figures 50-67. Engina species. 50-57. Engina itzamnai (Watters, 2009). 50-51. Holotype, UF 170226, 17.9 mm. 52-53. Paratype,
UF 425817, 91 in, NE of Contoy Light, Isla Contoy, Quintana Roo State, Mexico, 16.2 mm. 54-55. MW, 31 m, off NE Yucatan,
Campeche State, Mexico, 17.4 mm. 56-57, EFG 25800, 52-53 m, 64 km NNW of Cayos Areas, Campeche Bank, Campeche State,
Mexico, 17.4 mm. 58-67. Engina permixta new species. 58-59. Holotype, UF 479326, 16.1 mm. 60. Paratype, UF 479324, from
type locality, 14.5 mm. 61. Paratype , OSUM 39958, 21 m, off Piuma, Espirito Santo State, Brazil, 12.7 mm. 62. GTW 4265f, 20-25 m,
off Guarapari, Espirito Santo State, Brazil, 11.4 mm. 63. GTW 4265b, 21 m, off Piuma, Espirito Santo State, Brazil, 13.0 mm.
64. GTW 13807a, 40-50 m, off Conceif ao da Barra, Espirito Santo State, Brazil, 12.0 mm. 65. GTW 4265r, from type locality,
12.1 mm. 66. GTW 12026a, 40-45 m, off Biizios, Rio de Janeiro State, Brazil, 11.2 mm. 67. GTW 4265ah, 10-15 m, Porto da Barra,
Salvador, Bahia State, Brazil, 12.4 mm.
Page 106
THE NAUTILUS, Vol. 129, No. 3
(©) and Hesperistemia karinae (Nowell-Ustdcke, 1959) (•).
Areas, Campeche Bank, Campeche State, 20° 45.59' N,
-92° 12.63' W; MW (1), 31 m, off NE Yucatan,
Campeche State.
Distribution: Gulf of Mexico, Campeche Bank between
Arrecife Alaeran and Cayos Areas.
Habitat: Dead specimens were dredged in 46-180 m
on rubble bottom. No live collected specimens are known
to us.
Variation in Specimens: The few known specimens
vary slightly in the number of axial (8-10) and spiral
(1 1-13) sculptural elements.
Etymology: Hesperistemia itzamnai Watters, 2009:
Mayan, Itzamnd , the creator deity in Mayan mythology.
This species is only known from off the Yucatan Peninsula,
ancestral home of the Mayans. A masculine name. Engina
dicksoni Petuch, 2013: Named for Andrew Dickson, col¬
lector of the type.
Comparison with Other Species: This is the largest
western Atlantic Engina and the most fusiform. See com¬
parison with E. lignea new species, below. See Table 1 for
a comparison with other species.
Discussion: This is apparently a very rare species in
collections. Petuch (2013) renamed this taxon Engina
dicksoni.
Engina lignea new species
(Figures 10-13)
Description (based on 2 specimens): Only adult
specimen seen, holotype 14.3 mm in length x 7.4 mm in
width. Fusiform; spire ea. 50% total length. Shell thick.
solid. Protoconch small, conical, of 1.5 smooth, tan whorls.
Teleoconch of 5 whorls, strongly demarcated from
protoconch. Teleoconch sculpture of 9-10 rounded,
widely separated primary spiral cords between the suture
and the anterior limit of the siphonal canal, with 3-6
intercalated secondary threads. Subsutural slope wide,
flat, with single primary thread. Axial sculpture of widely
spaced, rounded primary ribs; 7 on penultimate whorl,
7 on final whorl, with numerous secondary very fine,
axial threads. Intersections of axial and spiral sculpture
with strong, elongated nodules, strongest at periphery.
Terminal varix weakly developed, low and wide. Aper¬
ture oval, outer lip with 5 lirate denticles extending seep
into aperture. Anal canal shallowly indented between
two weak teeth. Parietal wall erect anteriorly with 5 lirate
columellar folds. Siphonal canal moderately long, open.
Color uniformly light tan to brown, with a vague darker
band on sub-sutural plane, with or without lighter-
colored peripheral axial nodes. Aperture light tan. Oper¬
culum, radula, and anatomy unknown.
Holotype: UF 479325 (ex EFG 26687).
Type Locality: 46-48 m, 145 km WSW of Arrecife
Alaeran, Campeche Bank, Campeche State, Mexico,
22° 10.80' N, -91° 09.00' W.
Paratype(s): BMSM 76001(1), 50-55 m, 146 km
WSW of Arrecife Alaeran, Campeche Bank, Campeche
State, Mexico, 22° 11.46' N, -91° 08.71' W (ex EFG
25822).
Distribution: Known only from off northern Yucatan
Peninsula.
Habitat: Only empty shells have been found between
46-55 m.
Variation in Specimens: The two known specimens
differ in color from light tan to dark brown.
Etymology: Latin lignea , made of wood, in reference
to the texture and colors of the shells.
Comparison with Other Species: This species is
most similar to the sympatric E. itzamnai. It is a smaller
species that differs in having fewer primary axial ribs on
both the final whorl (8-10 in E. itzamnai , 7 in E. lignea)
and on the penultimate whorl (8-10 in E. itzamnai, 7 in
E. lignea), as well as fewer primary spiral cords (1 1-13 in
E. itzamnai, 9-10 in E. lignea). See Table 1 for a compar¬
ison with other species.
Discussion: This is the second new Engina described
from the Yucatan Peninsula. Both are deeper water
species that are rare in collections. Both have some char¬
acteristics of Hesperistemia but based on the analysis
of Landau and Vermeij (2012) we have placed them
in Engina.
Engina permixta new species
(Figures 58-67, 103)
G.T. Watters and K. Fraussen, 2015
Page 107
non Engina turbinella (Kiener, 1836). — Rios, 1975: 93,
pi. 27, fig. 385; Coltro, 2005: 1, pi. A, figs. 8-10
[ misidentif (nations] .
? Engina turbinella (Kiener, 1836). — Lopes and
Alvarenga, 1955: 173; Kempf and Matthews, 1968:
93; Matthews and Kempf, 1970: 32, 46; Eston et ah,
1986: 43; Leal, 1991: 153.
Description (based on 31 adult specimens): Larg¬
est adult specimen, 16.1 mm in length (holotype);
smallest specimen, 9.4 mm in length; holotype 16.1 in
length x 8.3 mm in width. Shell biconic; spire ea. 50%
total length. Protoconch small, conical, of 1.5 smooth,
white or brown banded whorls; last Va whorl may have
axial folds. Teleoconeh of 5.5 whorls, strongly demar¬
cated from protoconch. Teleoconeh sculpture of 7-10
(mode=9) primary spiral cords between the suture and
the anterior limit of the siphonal canal. Subsutural slope
wide, flat, with single primary cord. Three primary cords
gathered at periphery, remaining cords very widely
spaced, flattened or nodulose. 5-6 uniformly wide sec¬
ondary threads between these remaining cords. Axial
sculpture of widely spaced, angular primary ribs; 7-10
(mode=9, but often 8) on the final whorl, 8-10
(mode=8) on penultimate whorl, axial sculpture most
prominent on periphery, with numerous secondary fine,
axial threads in between primary ribs. Terminal varix
only slightly more prominent than preceding axial ribs,
angular. Aperture oval. Anal canal bounded by single
parietal denticle on columella and single anal denticle
on outer lip. Outer lip with 3-6 (mode=5) weak, irreg¬
ular denticles, posterior-most 2 often fused. The thin
rib that projects deep into the aperture and delimits
the siphonal canal found in other species is weak but
always present. 3-5 (mode=4) radial lira present at
posterior end, weak and irregular, not very distinct from
coluinellar folds. Anterior to these are 3-7 (mode=5)
denticulate coluinellar folds, irregular, often limited
to edge of parietal wall. Second or third anterior den¬
ticle often extending deep into aperture. Anterior-most
denticle may form a ridge extending along the parie¬
tal wall to the anterior margin. Interior ledge of colu¬
mella well-defined, often merging with anterior-most
denticle. Siphonal can til moderately short, open. Base
color variable: dark brown, tan, or yellow, axial ribs usu¬
ally darker, always with a continuous white band just
anterior to the periphery; this band may have yellow
portions to it. Aperture white to tan. Operculum oval,
yellow-brown, with a subterminal nucleus. Radula and
anatomy unknown.
Holotype: UF 479326 (ex GTW 4265r).
Type Locality: 1 in, Cajueiro, Rio Grande do Norte
State, Brazil.
Paratype(s): BMSM 76000 (1), from type locality (ex
GTW 4265r); OSUM 39958 (1), 21 m, off Piuma, Espfrito
Santo State, Brazil (ex GTW 13807b); UF 479324 (1),
from type locality (ex GTW 4265r). Other Material Exam¬
ined (51 specimens): Brazil: GTW 13807c (6), 1 in,
Cajueiro, Rio Grande do Norte State; MW (3), intertidal,
Ilha do Frade, Fernando de Noronha, Pernambuco State;
UF 126233 (2), Ponta Verde, Maceio, Alagoas State; MW
(1) , 1 m, Maragogi, Alagoas State; GTW 4265ah (2), EFG
14667 (3), 10-15 in, Porto da Barra, Salvador, Bahia State;
MW (5), Monte Serrat, Bahia State; MW (1), Barra,
Salvador, Bahia State; GTW 13807a (1), 40-50 m, off
Coneeiyao da Barra, Espfrito Santo State; GTW 13807b
(2) , KF 7343 (1), 21 m, off Piuma, Espfrito Santo State;
HGL (1), 15-25 m, off Guarapari, Espfrito Santo State;
GTW 4265f (1), GTW 4265o (1), GTW 4265s (2), HGL
(2), 20-25 in, off Guarapari, Espfrito Santo State; HGL
(1), 10-25 in, off Guarapari, Espfrito Santo State; GTW
13807d (2), 60-80 m, off Guarapari, Espfrito Santo State;
KF 6123, 4.5-60 m, off Guarapari, Espfrito Santo State
(1); KF 4928, 15-30 m, off Guarapari, Espfrito Santo
State (2); KF 80 (1), Rio de Janeiro, Rio de Janeiro State;
GTW 12026a (6), GTW 12026b (1), HGL (1), 40-45 in,
off Buzios, Rio de Janeiro State; HGL (1), 3.5-45 m, off
Buzios, Rio de Janeiro State; HGL (1), 40-46 m, off
Buzios, Rio de Janeiro State.
Distribution: Brazil, from Rio Grande do Norte State
to Sao Paulo State; Fernando de Noronha. If the speci¬
mens referred to E. turbinella by Leal (1991) are this
species, then it is also present at Abrolhos, Atol das
Rocas, Jaseur, and Davis and Dogaressa seamounts.
Habitat: Empty shells have been found from low tide
to 50 m; live individuals have been found from 1-45 m
under rocks and on rubble.
Variation in Specimens: Shell coloration varies from
nearly all dark brown to nearly all pale yellow to white,
with all intergrades. There is no elinal variation in shell
color and all colors may occur together. The dark brown
morph characterized by the holotype appears to be the
most common.
Etymology: Latin permixta , mixed, confused, in rela¬
tion to the disarray of color forms.
Comparison with Other Species: Although often
misidentified as E. corinnae, it is not similar to that
Florida-Gulf of Mexico species. It is most similar to
some color forms of E. demani , with which it may
co-occur. See the comparison under that species for
details. See Table 1 for a comparison with other species.
Discussion: The somewhat weathered leetotype of
Engina zonata Gray, 1839, is very close to some speci¬
mens of tills species. However, the paraleetotype clearly
shows the presence of sub-peripheral white nodules,
which do not occur in E. permixta.
Engina turbinella (Kiener, 1836)
(Figures 69-100, 102)
Page 108
THE NAUTILUS, Vol. 129, No. 3
Figures 69-101. Engina species. 69-100. Engina turbinella (Kiener, 1836). 69. Purpura turbinella Kiener, 1836: 29, pi. 9, fig. 25.
70-71. UF 281377, Scarborough, Tobago. 12.1 mm. 72. KF 4735, Punto San Juan, Venezuela. 12.3 mm. 73. UF 281378, Friendship
Beach, Tobago. 11.1 mm. 74. EFG, 60 ft., Montexuma Shoals, Mustique Island. 12.7 mm. 75. HGL, drift, Playa de Carenera,
Venezuela. 9.6 mm. 76. EFG 25590, 3 km N of Colon Island, Panama, 8.8 mm. 77. GTW 4265af, 1-2 m, Buraco da Rachel,
Fernando de Noronha, 9.3 mm. 78. UF 70393, Bruja Point, Canal Zone, 15.0 mm. 79. GTW 4265e, 0.3-1. 5 m, Punta Robles,
Ambergris Caye, Belize, 13.2 mm. 80-81. NHMUK 1982122, syntype of Engina elegans Gray, 1839, ca. 7.5 mm, photo courtesy
of Phil Hurst, NHMUK. 82. HGL, Isla de Utila, Honduras, 11.3 mm. 83. GTW 4265ag, 2-5 m, Isla la Tortuga, Venezuela, 13.3.
84. GTW 4265b, 7.3 m, Isla Morro Pelotas, Venezuela, 14.2 mm. 85. UF 146654, Miami, Miami-Dade Co., Florida, 14.0 mm.
86. GTW 4265h, 0.3-1. 5 m, Long Reef, Isla Roatan, Honduras, 8.6 mm. 87. GTW 4265c, 1 in, Sandbur Reef, Ambergris Caye,
Belize, 9.3 mm. 88. EFG 19507, Isla de Providencia, Colombia, 13.9 mm. 89. EFG 25673, Bocas del Toro, Panama, 10.4 mm.
90. MW, 1 .0-1.5 m. Sand Key, off Key West, Monroe Co., Florida, 11.0 mm. 91. KF 2503, 12 m, St. Vincent, S end of
Baliceaux Island, Grenadines, 12.3 mm. 92. MW, Pompano Beach, Broward Co., Florida, 13.5 mm. 93. KF 2551, 8-9 m,
Cannoun Island, Grenadines, 13.7 mm. 94. HGL, Current, Eleuthera, 12.1 mm. 95. GTW 4265w, 0.6-1. 8 m. Coral Cove,
Mt. Hartman Bay, Grenada, 10.5 mm. 96. GTW 4265aa, 0.3-1 m, Enrique, La Parguera, Puerto Rico, 9.7 mm. 97-98. UF 126216,
paratype of Engina turbinella cruzana (Nowell-Usticke, 1959), 9.2 mm. 99-100. NHMUK 1982122, lectotype of Engina zonata Gray,
1839, 11.5 mm. 101. Engina cf. turbinella. KF 5262, 12 m, Savan Island, Grenadines, 12.2 mm.
G.T. Watters and K. Fraussen, 2015
Page 109
Figure 102. Distribution of Engina permixta new species (©)
and Engina turbinella (Kiener, 1836) (•) (not shown, E. turbinella
record from Fernando de Noronha).
Buccinum B.r. parvum, striatum , asperum, ex albo...
Lister, 1685: pi. 953, fig. 3.
Purpura turbinella Kiener, 1836: 29, pi. 9, fig. 25.
Engina zonata Gray, 1839: 113; Tryon, 1883: 196, 247;
Bartsch, 1931: pi. 1, fig. 6; Faber, 2007: 73 [in syn¬
onymy of Purpura turbinella Kiener, 1836]; Fraussen
and Chino, 2011: 64 [in synonymy of Purjmra
turbinella Kiener, 1836]; Landau and Vermeij, 2012:
122 [in synonymy of Puqmra turbinella Kiener ,
1836],
Engina elegans Gray, 1839: 113; Tryon, 1883: 192; 213
[in synonymy of Purpura turbinella Kiener, 1836];
Pace, 1902: 78.
Ricinula turbinella (Kiener, 1836). — Reeve, 1846: spe¬
cies 42, pi. 5, fig. 42.
Enzina [sic] zonata Gray, 1847: 133; Tomlin, 1928: 40.
Turbinella ( Fusus ) elegans (Gray, 1839). — Tryon, 1881: 97.
Engina turbinella (Kiener, 1836). — Tryon, 1883: 192,
213, 244, pi. 62, figs. 38, 39; Dali and Simpson, 1901:
402; Warinke and Abbott, 1961: 116, pi. 21, fig. d;
Orr, 1962: text fig. [radula], pi. 10, fig. B; Woodring,
1973: 477^478, pi. 74, figs. 4, 5; Abbott, 1974: 218,
fig. 2399; Ode, 1983: 60, fig. 7; de Jong and Coomans,
1988: 82-83, pi. 38, fig. 450; Merlano and Hedegus,
1994: 187, fig. 710; Redfern, 2001: 93, pi. 43, fig. 395;
Coltro, 2005: 1, pi. A, figs. 6, 7; Faber, 2007: 74,
figs. 4, 5; Massemin et al, 2009: 148 and text figures;
Tunnell et al., 2010: 214; Fraussen and Chino, 2011:
64; Daccarett and Bossio, 2011: 95, fig. 448; Zhang,
2011: 119, figs. 404(1-3); Landau and Vermeij, 2012:
122-127; Redfern, 2013: 144, figs. 414a, b.
Engina turbinella cruzana Nowell-Ustieke, 1959: vi,
68-69; Warmke and Abbott, 1961: 116 [as a form].
Engina turbinella var. cruzana Nowell-Ustieke, 1959:
68-69; Nowell-Ustieke, 1969: 17, pi. 2, fig. “789”
[790]; Boyko and Cordeiro, 2001: 61.
Description (based on 26 adult specimens): Larg¬
est adult specimen, 14.9 mm in length; smallest speci¬
men, 6.6 mm in length. Spire occupies 30-50% of total
length. Protoconch minute, of 1.5 smooth whorls, tan
with or without a darker band. Protoconch almost always
eroded away, very rarely remaining in adult specimens.
Teleoconch of 5-6 whorls. Spiral sculpture consists of
6-10 (mode=8) primary spiral cords; the wide subsutural
slope has secondary threads and usually a single primary
(rarely two, rarely absent) cord somewhere between the
suture and the mid-point of the subsutural slope; this
cord may abruptly turn posteriorly at the edge of the
varix. The periphery has 2-3 primary cords grouped over
the axial nodes and microscopic threads; anteriorly there
are 3-7 wide, rounded ribs widely separated by 3—5 sec¬
ondary threads. Axial sculpture 8-13 (mode=10) elon¬
gated, undulating peripheral nodes on the final whorl,
9-14 (mode=10) on the penultimate whorl; usually
prominent but less so on small specimens. Terminal varix
slightly more developed than previous axial sculpture.
Aperture small, constricted. Anal canal bounded by
single parietal denticle on columella and single anal den¬
ticle on outer lip. Outer lip with 3-7 (mode=5) denticles;
the denticles are elongate but not lirate and do not
project far into the aperture. The posterior-most two
denticles are usually the most prominent and are fused;
the anterior-most denticle forming a strong, long, thin
rib that projects deep into the aperture and delimits the
siphonal canal. Radial lirae (3-7, mode=4) at posterior
end, usually long and thin but often not well-differentiated
from eolumellar folds. Anterior to these are columellar
folds (4-8, mode=6), irregular, some pustulose, some
lirate, often limited to edge of parietal wall, anterior-
most best-developed and forming a ridge on the inte¬
rior of the columella. In some specimens, particularly
small ones, the posterior half of the columella may be
deeply excavated. Background color usually dark brown
to black, rarely orange in some southeastern popula¬
tions. Axial ribs at periphery always colored white, either
as distinct spots or merged into a single spiral band;
these white spots may be limited to the adapertural side
of the ribs. Elsewhere, intersections of axial and spiral
sculpture may or may not be present as white spots.
1° spiral cord on sub-sutural band may be wliite at ter¬
mination. Aperture dark brown to black, fading to a
lighter shade interiorly; outer lip denticles white, colu¬
mellar ridge white. Operculum rounded, leaf-shaped,
yellow with brown central radius. Radula figured by On
(1962). Anatomy unknown.
Types: Purpura turbinella Kiener, 1836: Types not
located at the Museum d’histoire naturelle de la Ville de
Geneve (Y. Finet, pers. comm., 2011), the Museum
national d’Histoire naturelle, Paris (online database,
2011), nor tire Natural History Museum UK (K. Way,
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THE NAUTILUS, Vol. 129, No. 3
pers. comm., 2011), and are presumed lost. Engina
elegans Gray, 1839: Syn types, NHMUK 1982122 (5).
The specimens are heavily worn and/or immature. Engina
zonata Gray, 1839: Lectotype and paralectotype, NHMUK
1967592; the lectotype is the larger of the two, designated
by Orr (1962). Engina turbinella cruzana Nowell- Usticke,
1959: Lectotype, AMNH 198494, designated by Boyko
and Cordeiro (2001) [although listed as a variety on page
68 of Nowell-Ustieke, 1959, the name was immediately
used as a subspecies on the same page as well as earlier
on page vi], additional specimens implied but their dispo¬
sition is unknown except for UF 126216, Judith’s Fancy,
St. Croix, 7 paralectotypes.
Type Locality: Purpura turbinella Kiener, 1836:
"Unknown.” Engina elegans Gray, 1839, Atlantic Ocean.
Engina zonata Gray, 1839: Atlantic Ocean. Lectotype
label reads “West Indies.” Engina turbinella cruzana
Nowell-Ustieke, 1959: West Coast [place name]. Judiths
Fancy. Christiansted Harbor [all St. Croix]. Restricted by
Boyko and Cordeiro (2001) to West Coast but paralecto¬
types at UF are from Judiths Fancy.
Other Material Examined (951 specimens): Florida:
UF 12718 (1), S of Lake Worth Inlet, Palm Beach Co.;
UF 126238 (1), Boynton Beach, Palm Beach Co.; MW
(3), 18 m, Boynton Beach, Palin Beach Co.; UF 394034
(2) , Hillsborough Light, Broward Co.; MW (2), 18 m,
Pompano Beach, Broward Co.; UF 146654 (1), Miami,
Miami-Dade Co.; UF 185179 (1), Bear Cut, Key
Biseayne, Miami, Miami-Dade Co.; UF 126219 (4),
Miami County Causeway, Miami-Dade Co.; UF 126227
(11), Biseayne Bay, Miami, Miami-Dade Co.; UF 238288
(3) , Miami Beach, Miami-Dade Co.; UF 80978 (1), Key
Largo, Monroe Co.; UF 239825 (1), Marathon, Key
Largo, Monroe Co.; HGL (9), 3—4.5 m, Pickles Reef,
Key Largo, Monroe Co.; UF 126223 (11), Little Molasses
Reef, Key Largo, Monroe Co.; UF 80975 (2), Molasses
Reef, Key Largo, Monroe Co.; UF 126228 (1), 4.0 km
WSW of Carysfort Reef Light, Key Largo, Monroe Co.;
UF 37855 (1), Dry Rocks, Key Largo, Monroe Co.; UF
123130 (51), UF '352066 (7), Looe Key, Big Pine Key,
Monroe Co.; UF 352068 (6), UF 123056 (2), Little Torch
Key, Monroe Co.; UF 61023 (4), UF 70396 (11), UF
192067 (3), Key West, Monroe Co.; UF 126218 (44), UF
126243 (92), UF 153382 (9), Middle Sambo Shoals, Key
West, Monroe Co.; UF 80974 (41), Sambo Shoals, Key
West, Monroe Co.; UF 126236 (3), Sand Key Light,
Key West, Monroe Co.; UF 12714 (2), UF 70395 (3), UF
126231 (20), UF 192158 (2), MW (1), Pelican Shoals,
Key West, Monroe Co.; UF 126222 (3), Washerwomans
Shoals, Key West, Monroe Co.; MW (7), 1.0-1 .5 m, Sand
Key, off Key West, Monroe Co.; UF 12717 (1), UF
126239 (13), Ft. Jefferson, Dry Tortugas, Monroe Co.;
UF 12715 (1), Bush Key Reef, Dry Tortugas, Monroe
Co.; UF 12716 (1); Garden Key, Dry Tortugas, Monroe
Co. Bahamas: UF 126240 (5), North Bimini; GTW 4265a
(1), tide pool. South Bimini; UF 126246 (10), Gun Cay,
off Bimini; UF 80985 (1), Grand Bahama Island; UF
37853 (2), West End, Grand Bahama Island; UF 70391
(2), New Providence; UF 126224 (3), North Cay, Nassau,
New Providence; UF 126226 (9), Delaport Point, New
Providence; UF 126221 (2), Morgans Bluff, Andros; UF
126234 (2), Frazier Hog Cay, Berry Islands; GTW 4265n
(1) , 1. 0-2.5 m., Beachside, Chub Cay, Berry Islands; MW
(2) , Periwinkle Beach, Chub Cay, Berry Islands; HGL (1),
Cat Island; HGL (2), Current, Eleuthera; MW (2), 18.0 m.
Cay Sal. Cuba: UF 80982 (12), UF 80995 (5), Santiago;
UF 80993 (5), Jauco; UF 80983 (7), Varadero; UF 80994
(1) , Baracao; UF 54955 (1), Guantanamo; UF 80992 (4),
Gibara; UF 17997 (6), La Habana. Dominican Republic:
UF 239824 (2), Barahona. Cayman Islands: UF 28935 (4),
Preston Bay, Little Cayman; UF 239823 (1), Cayman
Turtle Farm, Grand Cayman; UF 126230 (1), Red Bay,
Grand Cayman. Jamaica: UF 61025 (4); UF 80987 (10),
Flag Point. Puerto Rico: UF 80988 (1), Fajardo; GTW
4265z (1), 0.3-1 in, Eseambron, San Juan; GTW 4265aa
(3) , 0.3-1 m, Enrique, La Parguera; GTW7 4265ab (3),
0.3-1 m, Collao, La Parguera; GTW 4265ac (3), 0.3-1 m,
Playa Buye, La Parguera; GTW4265ad (7), 0.3-1 m, Media
la Luna, La Parguera; GTW 4265ae (4), 1-3 m, El Corral,
La Parguera; MW (3), Pihones Beach; HGL (7), Playa de
Sardinera. U.S. Virgin Islands: UF 37856 (2), UF 80979
(43), Water Island; MW (3), 1 m, St. Croix; MW (1), 1 m.
Ham Bay, St. Croix; UF 126216, St. Croix [7 paratypes of
Engina turbinella cruzana (Nowell-Ustieke, 1959)]. British
Virgin Islands; UF 163643 (9), Tortola; KF 3929 (1),
HGL (2), beached, Beef Island; HGL (3), 0.6 m. Buck
Island; UF 163644 (4), Guana Island. Antigua: UF 80977
(3), UF 126235 (7); UF 120786 (2), Curtain Bluff GTW
4265ai (5), 0.3-1 m. Long Bay; GTW 4265aj (2), 0.3-1 m,
Deep Bay; GTW 4265ak (8), 0.3-1 m, Morris Bay; GTW
4265al (1), 0.3-1 in. Valley Church Bay; GTW 4265am
(2) , 0,3-1 m. Half Moon Bay; GTW 4265an (2), 0.3-1 m.
Valley Church Bay; GTW 4265ao (1), 0.3-1 m, Cades
Bay. Grenadines: EFG (1), 18 m, Montexuma Shoals,
Mustique Island; KF 2503 (3), 12 in, St. Vincent, S end
of Baliceaux Island; KF 5262 (3), 12 m, Savan Island;
KF 2551 (1), 8-9 m, Cannoun Island. Grenada: GTW
4265t (12), GTW 4265u (14), GTW 4265v (8), GTW
4265w (4), GTW 4265x (7), GTW 4265y (2), 0.6-1.8 m,
Coral Cove, Mt. Hartman Bay. Barbados: UF 80976
(2) ; UF 126220 (23), Hastings Rock. Tobago: UF 70391
(3) ; UF 80980 (25); UF 126225 (6), UF 126237 (6),
Buccoo Reef UF 281378 (6), Friendship Beach; UF
281377 (7), UF 352067 (2), Scarborough; MW (2),
Lambeau; MW (3), Irvine Bay. Mexico: EFG 26687 (1),
46-48 m, Campeche State, 22° 10.80' N, -91° 09.00' W;
UF 383287 (4), Punta Hogna, Quintana Roo State; UF
264023 (1), Cayos Lobos, Quintana Roo State; UF
38338 (3), Cozumel, Quintana Roo State. Belize: GTW7
4265c (4), 1 m, Sandbur Reef, Ambergris Caye;
GTW 4265d (1), 1.5 m, Tres Cocos, Ambergris Caye;
GTW 4265e (2), 0. 3-1,5 m, Punta Robles, Ambergris
Caye. Honduras: EFG 7365 (9), MW (2), Caribe Point,
Isla Roatan; HGL (1), 0,5-1. 3 m, Isla Roatan; GTW 4265g
(1), 0.3-1. 5 m, West Id me Key, SE Isla Roatan; GTW7
4265h (1), GTW7 4265i (2), GTW 4265j (1), 0.3-1. 5 m,
G.T. Watters and K. Fraussen, 2015
Page 111
Long Reef, Isla Roatan; GTW 4265k (6), SE Isla
Roatan; GTW 42651 (1), 1-1.5 m, Old Port Royal, Isla
Roatan; GTW 4265m (2), 1-1.5 m, Horseshoe Reef,
Isla Roatan; HGL (5), Oak Ridge, Isla Roatan; KF 2173
(1), shallow water, Cayos Cochinos; KF 4141 (2), 2 m,
Isla de Utila; MW (3), intertidal, Isla de Utdla; HGL
(19), Isla de Utila; UF 380423 (2), Guanaja; MW (2),
1 m, Guanaja. Costa Rica: UF 163876 (1), Punta Cahuita.
Panama: UF 126217 (10), Colon; UF 80981 (9), UF
80989 (7), UF 160525 (2), Galeta Point; UF 80990 (2),
Bocas del Toro; UF 160524 (3), Careening Cay, Bocas
del Toro; UF 80991 (3), Devils Beach; KF 3593 (2),
Tiger Island; EFG 25590 (2), Isla Colon; UF 70393 (4),
Bruja Point. Colombia: KF 3590 (3), Santa Marta; EFG
19507 (5), EFG 19508 (1), Isla de Providencia; EFG
25673 (1), Bocas del Toro. Venezuela: GTW 4265b (1),
7.3 in, Isla Morro Pelotas; GTW 4265ag (3), 2-5 m, Isla la
Tortuga; HGL (5), Playa de Carenero; MW (1), Piedras
Negras, Paraguana Peninsula; MW (2), Isla de Margarita;
MW (3), El Tunal, Isla de Margarita; EFG 19264 (1),
Isla Borracha; KF 4735 (1), Punta San Juan. Netherlands
Antilles: UF 80984 (2), UF 163642 (7), Bonaire; MW (6),
1.2 m, Kralendijk, Bonaire; UF 80986 (2), Aruba; UF
126245 (2), Palm Beach, Aruba. Brazil: GTW 4265af
(1), 1-2 m, Buraco da Rachel, Fernando de Noronha;
KF 78 (12), low tide, Fernando de Noronha.
Distribution: Widely distributed in southern Florida,
rarely in the northern Gulf of Mexico, throughout the
Bahamas, the Greater and Lesser Antilles, and along the
Central American and northern South American coasts.
Massemin et al. (2009) recorded this species from lies du
Salut, French Guiana. Daccarett and Bossio (2011)
recorded it from most of the Colombian Caribbean coast.
Perhaps the strangest aspect of this species’ distribution
is its occurrence on Fernando de Noronha, 2,400 km
away from the next known site in French Guiana, where it
co-occurs with E. demani. Tryon (1881) recorded E. elegans
from Sierra Leone but it is not listed by Ardovini and
Cossignani (2004) and this record is probably in error.
Habitat: A shallow water species found from the inter¬
tidal zone to ca. 20 m. It seems to occur in deeper water
off Florida and the northern Gulf of Mexico than else¬
where in the Caribbean. It is most commonly found
under slabs of coral rubble, often in pairs, often almost
completely covered in a thick, coralline algal deposit.
Variation in Specimens: Specimens vary greatly in
size and degree of “fusiformity,” but are relatively uni¬
form in sculpture and, with few exceptions, coloration.
All specimens have a peripheral band of white spots on
the axial cords that may merge into a single unbroken
band. In some populations the single spiral cord on the
sub-sutural plane and/or the spiral cords anterior to the
periphery may have white nodes as well. The two poste¬
rior-most outer lip denticles are almost always fused into
a single large tooth. In the southeastern populations
some specimens are an orange color (see discussion
below) with numerous white spots. Specimens from the
Bahamas and the Lesser Antilles tend to be smaller than
elsewhere, but there are numerous exceptions.
Etymology: Engina turbinella (Kiener, 1836): Latin,
diminutive of turbina, a top. Engina elegans Gray, 1839:
Latin, elegant. Engina zonata Gray, 1839: Latin, banded.
Engina turbinella cruzana Nowell-Ustieke, 1959: Spanish,
cniz (cross), from the original name given by Christopher
Columbus to the island of St. Croix, Santa Cruz.
Comparison with Other Species: The combination
of a uniformly dark background color with white spots or
bands, fused outer lip teeth, and dark aperture with
white teeth characterize this species. Some specimens
are similar to the Brazilian E. permixta-, that is a relatively
larger species with fewer axial ribs on both the final
whorl (8-13 in E. turbinella , 7-10 in E. permixta) and
the penultimate whorl (9-14 in E. turbinella, 8-10 in
E. permixta). Engina turbinella never shows the wide range
of colors seen in E. permixta. See Table I for a comparison
with other species.
Discussion: Engina turbinella (Kiener, 1836) is a
widespread and usually shallow water species occurring
throughout much of the tropical western Atlantic Ocean.
It is a baffling species varying greatly in size, shape, and
coloration. Three color forms are apparent, all named,
that appear rather distinct unless a large number of
specimens is studied. These are discussed below.
Purpura turbinella Kiener, 1836. Louis Charles
Kiener curated the vast (150,000 specimens) collection
of Baron Jules Paul Benjamin Delessert, describing and
illustrating many species in his Specie s General (Dance,
1966). This included the “jolie petite espece” Purpura
turbinella from an unknown habitat, originally from the
Prince Massena collection, which Delessert had acquired
in 1840. Subsequently this name has been almost univer-
sallv applied to all western Atlantic Ocean Engina taxa as
“ Engina turbinella.” Although the type is apparently lost,
the illustrations (Figure 69) clearly show an orange shell
with small white nodules above and below the usual
peripheral row of nodules. This is not the widely distrib¬
uted typical form illustrated in most accounts as Engina
turbinella. Kieners illustrations match our specimens
from Scarborough, Tobago (Figures 70, 71). Although the
size of the figured type specimen - “7 ligjnes]” [= 15.8 mm]
is larger than the largest specimen we have seen from
there (12.3 mm), given the great variation in size among
other Caribbean Engina species we do not feel this is
significant. These specimens represent a morph that
appears to have a rather limited distribution from Barbados
and Tobago to adjacent western Venezuela, including
the Netherlands Antilles. It should be noted that orange
shells occasionally are seen throughout the remainder of
the western Atlantic Ocean and that shells may fade to
an orange or tan color as well.
Engina zonata Gray, 1839. Tomlin (1928) mentioned a
tablet at NHMUK labeled “ Enzina [sic] zonata Gray. B.V.
1 13” with two examples. He misidentified them as Engina
leucozona (Philippi, 1844) from the Mediterranean Sea.
Page 112
THE NAUTILUS, Vol. 129, No. 3
Orr (1962) quoted Tomlin as saying the specimens repre¬
sent “a possible type lot” but this phrase does not appear
in his 1928 paper. Bartsch (1931) did not pick a lectotype
of zonata, but simply illustrated one of the two specimens
as the genotype of Engina without further comment. On'
(1962) selected and illustrated a lectotype from the same
lot that Tomlin saw. Gray (1839) greatly mis -represented
the size of the specimen as 1.5 inches long, a fact appar¬
ently overlooked by Bartsch (1931) but noted by Orr
(1962). Tryon (1883: 196) also commented on Engina
zonata: “This is also a lost species.” But types exist and
prove to be E. turbinella. This is the morph most com¬
monly found throughout the western Atlantic, where it
insensibly grades into the other E. turbinella morphs
described here. It is often quite large, particularly in pop¬
ulations off Miami and Panama. Like the E. turbinella
morph described above it usually has small, white nodules
below the periphery (not apparent on the lectotype but
visible on the paralectotype of E. zonata) but lacks the
white sub-sutural nodules.
Engina elegans Gray, 1839 (and Engina turbinella
cruzana Nowell- Usticke, 1959). This is the morph usually
associated with the Greater Antilles and the Bahamas. It is
often quite small in comparison to the E. zonata morph
and usually lacks the sub-peripheral white nodules.
Examination of over 900 specimens indicates that
these three morphs seem to blend from one to the other
without clear-cut distinctions. Thus we prefer to recog¬
nize only the single species, E. turbinella. However,
phylogenetic work may yet separate this “species” into
two or more potentially cryptic species. The southeastern
Caribbean form, the basis of the name Puqmra turbinella
Kiener, 1836, in particular seems the most morphologi¬
cally and zoogeographically distinct of the three morphs.
A peculiar form from the Grenadines (Figure 101) may
represent a distinct species as well.
This is the only extant Engina species from the west¬
ern Atlantic with a fossil record. It was recognized from
the middle Miocene of Panama by Woodring (1973) and
the late Pleistocene of the Dominican Republic by
Landau and Vermeij (2012).
Engina wtlliamsae new species
(Figures 13, 103-107)
Description (based on 11 adult specimens): Larg¬
est specimen, 15.0 mm in length; smallest specimen,
11.5 mm in length; holotvpe 12.6 in length x 7.4 mm in
width. Spire occupies 60% of total length. Protoconch
minute, of 1.5 smooth whorls, white with tan patches or
bands. Teleoconch of ca. 5 whorls. Spiral sculpture con¬
sists of a wide subsutural slope with a narrow primary
thread with microscopic secondary threads; the periph¬
ery has 4 indistinct primary cords over the axial nodes
and microscopic threads; anteriorly there are 5 wide, flat
primary cords separated by deeply incised channels.
Axial sculpture of numerous microscopic threads on
sub-sutural band and 8-10 (mode=9) elongated primary
peripheral nodes on the final whorl, 9-10 (mode=9) on
penultimate whorl. Terminal varix only slightly more
developed than previous axial sculpture. Aperture small,
constricted. Anal canal bounded by single parietal denti¬
cle on columella and single anal denticle on outer lip.
Outer lip with 5-6 (mode=5) semi-lirate denticles, the
anterior-most forming a weak, long, thin rib that projects
deep into the aperture and delimits the siphonal canal,
barely perceptible in some specimens. Radial lirae (2-5,
mode=5) at posterior end, weak and irregular. Anterior
to these are columellar folds (2-6, mode=6), weak or
absent from center of columella, often limited to edge
of parietal wall, anterior-most two best-developed and
forming a ridge on the interior of the columella. Back¬
ground color white, subsutural slope gray, wide brown
blotches on the subsutural slope and between the axial
ribs on the posterior half, leaving a white sub-peripheral
band, anteriorly colored with brown spots, more or less
aligned axially. Aperture pale tan. Operculum rounded,
leaf-shaped, yellow with brown central radius. Radula
and anatomy unknown.
Holotype: UF 478947 (ex MW).
Type Locality: 15 m, off Egmont Key, Hillsborough
Co., Florida, USA.
Paratype(s): UF 478946 (1), 15 m, off Egmont Key,
Hillsborough Co., Florida, USA (ex MW); BMSM
75998 (1), off Egmont Key, Hillsborough Co., Florida,
USA (ex MW).
Other Material Examined (9 specimens): Western
Florida: UF 239809 (1), 220 m, off Egmont Key,
Hillsborough Co.; HGL (2), MW (4), 15 in, off Egmont
Key, Hillsborough Co.; MW (1), 14 m, off Egmont Key,
Hillsborough Co.; MW (1), 14 m, off Anna Maria Island,
Manatee Co.
Distribution: Known only from off Egmont Key and
adjacent Anna Maria Island, Hillsborough and Manatee
counties, Florida, Gulf of Mexico.
Habitat: Live and dead specimens collected on rock
reef in 15 in. Other specimens known to 220 m.
Variation in Specimens: The few known specimens
are remarkably uniform in coloration and sculpture.
Etymology: Named for Margaret “Peggy” Williams of
Tallavast, Florida, who collected most of the known
examples.
Comparison with Other Species: This species is
similar to E. corinnae. That species has more prominent
spiral sculpture over the entire shell, including the sub¬
sutural band, which is nearly smooth in E. willianisae .
See Table 1 for a comparison with other species.
Discussion: Engina ivilliarmae is only known from
Egmont Key and Anna Maria Island off the Gulf coast
of Florida. These sites are at the mouth of Tampa Bay, a
large open-water estuary. It has not yet been found on
G.T. Watters and K. Fraussen, 2015
Page 113
Figures 103-120. Engina species. 103-107. Engina williamsae new species. 103-104. Holotype, UF 478947, 14.5 mm. 105. Paratype,
BMSM 75998, 13.0 mm. 106. Paratype, UF 479946, 15.0 mm. 107. MW, 13.6 mm. All specimens from 15 m, off Egmont Key,
Hillsborough Co., Florida. 108-119. Hesperistemia karinae (Nowell-Usticke, 1959). 108-109. Holotype, AM Nil 193771, 23mm.
110. KF 2505, 5-7 m, Ronde Island, Grenada, 18.5 mm. 111. HCL, 9.1 m, Pickles Reef, Monroe Co., Florida, 19.9 mm. 112. HGL,
201 m, St. Anne, Guadeloupe, 19.8 mm. 113. GTW 6613c, 20-25 m, off Guarapari, Espirito Santo State, Brazil, 21.0 mm. 114. GTW
6613a, 6.1 m, Tambor Cay, Panama, 18.5 mm. 115. GTW 6613e, French Guiana, 17.1 mm. 116. GTW 6613b, 12 m, off Guarapari,
Espirito Santo State, Brazil, 21.9 mm. 117. KF 4535, Isla Los Aves, Venezuela, 19.3 mm. 118. HGL, Playa de Ponce, Puerto Rico,
21.1 mm. 119. KF 2947, Port Louis, Guadeloupe, 20.8 mm. 120. Hesperistemia sp., GTW 6613d, Amuay, Venezuela, 18.8 mm.
Page 1 14
THE NAUTILUS, Vol. 129, No. 3
adjacent Mullet and Sister keys. Given the amount of
recreational shelling that takes place in the area it is
surprising that more examples have not been found.
Most specimens are worn and crabbed.
Hesperistemio Gardner, 1944
Type Species: Hesperistemio. waltonia Gardner, 1944,
by original designation.
Discussion: In Hesperistemio the lirae extend much
further back within the aperture than in Engina. Vermeij
(2006) reviewed the fossil and Recent species. Additional
notes are given here for Hesperistemio korinae (Nowell-
Ustieke, 1959) because it occupies a much larger range
than is usually thought and is often misidentified in col¬
lections. It is redescribed and its distribution is given so
that its proper place in the western Atlantic fauna may
become more clear.
In addition to H. korinae, other western Atlantic
Ocean Recent taxa include Hesperistemio multangulo
(Philippi, 1848) and Hesperistemio harasewychi (Petueh,
1987). Abbott (1986) named a subspecies, //. multangulo
grandana, which Vermeij (2006) considered a minor
variant, hut which Petueh (2013) raised to a full species.
Hesperistemio itzomnoi Watters, 2009, and Engina
janoivskyi, placed in Hesperistemio by Watters (2009),
are reallocated to Engina based on the study of Landau
and Vermeij (2012). An additional undescribed Hesperistemio
occurs at Amuay, Venezuela, but it is only known to us
from a single specimen (Figure 120).
Hesperistemio korinae (Nowell-Usticke, 1959)
(Figures 68, 108-119)
Tritonidea orbignyi Payraudeau, 1826. Dali and Simpson,
1901: 400 [misidentification],
Canthorus korinae Nowell-Usticke, 1959: 69, pi. 4, fig. 4;
Warmke and Abbott, 1961: 118 [in synonymy of
Canthorus lautus Reeve, 1846]; Nowell-Usticke,
1969: 17, pi. 4, fig. 4; Nowell-Usticke, 1971: 13-14,
pi. 2; Leal, 1991: 153, pi. 19, fig. G.
non Canthorus lautus Reeve, 1846. — Warmke and
Abbott, 1961: 118, pi. 21, fig. c [misidentification],
Pisanio korinae (Nowell-Usticke, 1959). — Kaieher,
1986: card 4370; de Jong and Coomans, 1988: 84.
Hesperistemio korinae (Nowell-Usticke, 1969 [sic]). —
Vermeij, 2006: 81.
Hesperistemio korinae (Nowell-Usticke, 1953 [sic]). —
Watters, 2009: 271.
Description (based on 19 adult specimens): Larg¬
est adult specimen, 25.0 mm in length; smallest speci¬
men, 17.0 mm in length. Spire occupies 50% of total
length. Protoconch minute, of 1.5 smooth whorls, tan
with a brown band. Teleoeonch of ca. 6.25 whorls. Spiral
sculpture of 10-13 (mode =12) primary ribs, usually
absent from the sub-sutural ramp, between which are
single secondary threads flanked by a few tertiary
threads. Axial sculpture of large primary ribs, absent or
greatly reduced on the subsutural slope, most developed
at the periphery; 6-8 (mode=7) on both the last whorl
and on the penultimate whorl. Intersections of axial and
spiral sculpture form elongated spiral pustules on the
primary spiral cords. Terminal varix only slightly more
developed than previous axial sculpture. Aperture oval,
constricted. Anal canal bounded by single parietal denti¬
cle on columella and single anal denticle on outer lip.
Outer lip with 8-10 (mode=10) lirate denticles, in all but
the anterior two, which are quite short, the lirae curve
hack deep into the aperture. The long, thin rib that
projects deep into the aperture and delimits the siphonal
canal in some Engina is not present. Radial lirae at pos¬
terior end composed of combinations of 4-10 pustules
and irregular lirae, often mixed together. Anterior to
these are columellar folds (8-10, mode=10) posteriorly
composed of irregular pustules, anterior 2-3 folds of
long lirae. Interior ledge of columella well-developed.
Background color yellow or tan with a prominent, wide,
undulating, white peripheral band. This band is often
bounded on each side by a thin dark brown line and
may have dark brown flecks embedded within it. A dif¬
fuse, broad, dark band occurs above and below the white
peripheral band. Subsutural slope and pustules lighter in
color - yellow or tan. Aperture white, columella showing
through to brown below. Operculum rounded, leaf¬
shaped, yellow with brown central radius. Radula and
anatomy unknown. Leal (1991) figured the protoconch.
Holotype: AMNH 193771.
Type Locality: N of Fredericksted, [St. Croix, US Virgin
Islands],
Paratype(s): AMNH 294356 (1); UF 154704 (1); both
from the type locality.
Other Material Examined (35 specimens): Florida:
UF 352842 (1), UF 352843 (1), dredged off Delray
Beach, Palm Beach Co.; UF 126511 (1), 27 m, off Palm
Beach Inlet, Palm Beach Co.; UF 126510 (1), 37 m, off
Palm Beach, Palm Beach Co.; UF 157523 (4), 18 m,
Pompano Beach, Broward Co.; UF 398301 (1), 365.8 m,
off Hillsboro, Broward Co.; HGL (1), 9 m, off Pickles
Reef, Key Largo, Monroe Co. Bahamas: UF 168507 (1),
Cat Cay, Bimini. Puerto Rico: UF 162628 (1), Coreega
Beach, Rincon; UF 162629 (1), Rincon; HGL (3), Playa
de Ponce. U.S. Virgin Islands: UF 154704 (1), N of
Fredericksted, St. Croix [paratype], Guadeloupe: HGL
(3), 201 m, St. Anne; KF 2947 (4), Port Louis. Grenada:
KF 2505 (1), 5-7 in, Ronde Island. Panama: GTW 6613a
(1), 6 in, Tambor Cay. Venezuela: KF 4535 (1), Isla Los
Aves. French Guiana: GTW 6613e (1). Brazil: HGL (1),
6 m, Salvador, Bahia State; HGL (1), 10-12 m, Salvador,
Bahia State; HGL (1), off Guarapari, Espirito Santo
State; GTW 6613c (2), 20-25 m, off Guarapari, Espirito
Santo State; GTW 6613b (2), 12 m, off Guarapari,
Espirito Santo State.
G.T. Watters and K. Fraussen, 2015
Page 115
Distribution: This is a rare but widely distributed spe¬
cies: southeastern Florida, sporadically throughout the
eastern Caribbean to Espfrito Santo State, Brazil, but
with considerable gaps in this distribution. It has not
been recorded from Central America beyond Panama,
or from Cuba or Hispaniola, although it probably occurs
there. It was not included as part of the North American
fauna by Abbott (1974). Recorded by Dali and Simpson
(1901) from Ponce and Culebra, Puerto Rico, as Tritonidea
orbigmji Payraudeau, 1826. Nowell-Usticke (1971) added
Antigua, Barbados, Grenada, Curasao, and Aruba to his
original description. Leal (1991) recorded it from Trindade
and Vitoria seamounts.
Habitat: Live and dead individuals have been found
from 6-366 m. It appears to live in shallower water in the
southern Caribbean than in the northern part of its
range. It has been found on calcareous rubble.
Variation in Specimens: The degree of development
of the axial sculpture varies considerably. The overall
coloration varies from yellowish to a mahogany color but
the wide, white peripheral band is always present.
Etymology: Named after Nowell-Ustickes wile, Karin.
Comparison with Other Species: This relatively large
and brightly colored species should not be confused with
any other species from the western Atlantic. No species of
Engina from there has a similar color pattern, or the
elongate, lirate outer lip denticles, or the pustulose radial
lirae. See Table 1 for a comparison with other species.
Discussion: This species has been identified in most
collections as Buccinum lautum Reeve, 1846, described
from an unknown locality. That species is the eastern
Pacific taxon Gemophos lautus (Reeve, 1846).
ACKNOWLEDGMENTS
The authors are indebted to the following individuals for
allowing us to examine their valuable private collections
and for donating type material: Mark Chapman, Emilio
Garcia, Harry Lee, Charlotte Thorpe, David Massemin,
and Peggy Williams. We also thank the following for
allowing us access to collections or providing photo¬
graphs and information: John Slapcinsky and Gustav
Paulay (UF), Kathy Way and Phil Hurst (NHMUK),
Adam Bal dinger and Jennifer Lenihan (MCZ), Mark
Siddall and Sarfraz Lodhi (AM Nil), and Marien Faber
(The Netherlands). Comments by G. Vermeij (Univer¬
sity of Califormia, Davis) and M.G. Harasewych
(USNM) substantially improved this paper.
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THE NAUTILUS 129(3): 118-125, 2015
Page 118
On two abyssal species of Scaphandridae G.O. Sars, 1878
(Gastropoda: Cephalaspidea) from the eastern Pacific
Angel Valdes
Department of Biological Sciences
California State Polytechnic University
3801 West Temple Avenue
Pomona, CA 91768 USA
Email: [email protected]
James H. McLean
Natural History Museum of Los Angeles County
900 Exposition Boulevard
Los Angeles, CA 90007 USA
Email: [email protected]
ABSTRACT
Scaphander interruptus Dali, 1890 is a widespread, deep-water
eastern Pacific species whose range is extended to California.
The anatomical features of this species are similar to those of
other members of Scaphander de Monfort, 1810, including
the presence of a single lateral tooth on each half-row of the
radula and the morphology of the penial complex. Another
eastern Pacific species, S. cijlindrellus Dali, 1908 is reassigned
to the genus Cylichnium Dali, 1908 because of the presence
of a multidenticulate radula, a female copulatory organ and its
shell moqrhology.
Additional Keywords: Scaphander, Cylichnium, Panamie, deep
sea, redescriptions.
INTRODUCTION
The family Scaphandridae, recently resurrected by
Malaqnias et al. (2009), is poorly known in the eastern
Pacific, particularly in the deep sea. Two species are
problematic and in need of revision. A review of the
literature and material collected from several localities
has raised doubts about the taxonomic placement of
Scaphader cylind'rellus Dali, 1908. Newly collected
material of Scaphander interruptus Dali, 1890 showed a
previously unrecorded variability in shell morphology and
has allowed for anatomical studies. The present paper
deals with these two species and attempts to place them
taxonomically in light of recent studies on the system-
aties of Scaphandridae (Valdes 2008; Malaqnias et al.
2009; Eilertsen and Malaqnias, 2013a). Additionally, new
information on their morphological variably, range, and
internal anatomy is provided.
MATERIALS AND METHODS
The material examined is deposited at the Natural History
Museum of Los Angeles County (LACM), tire Department
of Invertebrate Zoology and Geology at the California
Academy of Sciences, San Francisco (CASIZ), the Benthic
Invertebrate Collection of the Seripps Institution of Ocean¬
ography (SIO), and the National Museum of Natural
History (USNM). The material is noted here as “shell” for
empty shells and “specimen” for complete specimens
including shell and soft parts. The material was unsuitable
for molecular work.
The specimens were dissected for examination of the
reproductive and digestive system anatomy. The repro¬
ductive anatomy was examined under a Nikon SMZ-100
dissecting microscope and drawn with the aid of a camera
lucid a. The gizzard plates and radulae were isolated
from surrounding tissue by submerging them in NaOH
10%, then rinsed in water, dried and examined under a
Cambridge 360 Scanning Electron Microscope at the Uni¬
versity of Southern California Center for Electron Micros¬
copy and Microanalysis.
SYSTEM ATICS
Genus Scaphander de Monfort, 1810
Type Species: Bulla lignaria Linnaeus, 1758 (Mediter¬
ranean and northeastern Atlantic), by original designation.
Scaphander de Montfort, 1808-10 [1810]: 334.
Diagnosis: Shell external, oblong, tapered posteriorly,
with the spire concealed by callus and covered with a
thin periostracum. Microscupture consisting of spiral
lines or series of pits. Aperture as long as the shell,
narrow above, much wider below. Radula with single,
hamate lateral teeth. Rachidian teeth generally absent,
a vestigial rachidian tooth present in several species.
Reproductive system monaulic. Penis unarmed, connected
to the prostate by a long duct. Gizzard with two large,
paired kidney-shape to sub-triangular plates and an unpaired
thin, elongate plate.
Remarks: Scaphander is a genus of predominandy
deep-sea cephalaspidean sea slugs, with species distributed
A. Valdes and J.H. McLean, 2015
Page 1 19
worldwide from the Arctic to the Antarctic (Eilertsen
and Malaquias, 2013a). The shell morphology and
anatomy of S. lignarius (Linnaeus, 1758), the type species,
have been summarized and illustrated by Thompson
(1976) and by Eilertsen and Malaquias (2013a). Eilertsen
and Malaquias (2013a) provided the first molecular phy-
logeny for this group including Atlantic and Western
Pacific species, and Eilertsen and Malaquias (2013b)
examined the digestive system morphology and diet of
this group.
Scaphander interruptus Dali, 1890
(Figures 1-10)
Scaphander interruptus Dali, 1890: 297, pi. 12, fig. 12.
Dali, 1908: 239.
Shell Morphology: Shell up to 22 mm in length
and 11 mm in width (LACM 73-109), involute, oval
(Figure 1). Aperture as long as the shell, wider anteri¬
orly and constricted posteriorly. Lip rising slightly above
the apex (Figures 1, 6, 7). The sculpture consists of spiral
lines of oval pits, with alternating rows of wider and
narrower pits (Figure 2).
Digestive System: The buccal mass is small and con¬
nects dorsally with the esophagus and the salivary glands
(Figure 10). At its posterior end two strong retractor
muscles attach. The esophagus opens into a large mus¬
cular gizzard, which contains three gizzard plates. Two of
the gizzard plates are large and oval (Figure 3), whereas
the third one is elongate (Figure 4). The radular formula
is 12 x 1.0.1 (n = 1). Lateral teeth are hamate, with a
number of small denticles (Figure 5).
Reproductive System: The reproductive system is
monaulic, but the female parts of the reproductive system
were not seen. The penial complex consists of a bulbous,
muscular penis and an oval prostate connected to the
penis by an elongate duct (Figure 9).
Material Examined: CALIFORNIA (all specimens
collected by K. L. Smith, R./V New Horizon), approxi¬
mately 226 km west of Point Arguello, Santa Barbara
County (34°44' N, 123°07' W), 4100 in depth, 1 fragment
of shell, 22 July 1991 (LACM 91-130.2); (34°43' N,
123°07' W), 4100 m depth, 1 broken shell, 1 August
1991 (LACM 91-131.3); (34°45' N, 123°04' W), 4100 m
depth, 1 specimen, dissected, 21 October 1991 (LACM
91-133.3); (34° 44' N, 123° 12' W), 4100 in depth, 1 frag¬
ment of shell, 4 November 1993 (LACM 93-74.1);
(34° 42' N, 123°08' W), 4100 m depth, 1 fragment of
shell, 7 November 1993 (LACM 93-75.2); (34° 43' N,
123°14' W), 4100 in depth, 1 shell, 12 October 1996
(LACM 1996-85.3). COSTA RICA, 14 miles from Punta
Guiones (9°45'18" N, 85°52'24" W), 12 May 1973,
1866 in depth, 1 shell, leg. R/V Velero IV (stn. 18932)
(LACM 73-109). PERU, West of Isla Lobos de Tierra
(6°26' S, 81°05' W), 1025 m depth, 1 shell, leg. McLean
and Del Solar, 23 January 1974 (LACM 74-18). CHILE:
R/V USS Albatross Expedition, station 2788, west coast
of Chile (45° 35' S, 75°55' W), 1050 fathoms (1920.24 m),
3 shells, 11 February 1888 (SYNTYPES, USNM 97075).
Geographic Range: This species was originally described
from Chile. It is otherwise known from southern California,
Baja California, Costa Rica, and Peru (present paper).
Remarks: Scaphander interruptus Dali, 1890 is the
only species of the genus known from the eastern Pacific.
Examination of the type series (Figure 6) confirms the
identity of the material here studied as S. interruptus; the
anatomical examinations conducted confirm the placement
of this species in Scaphander. The radula, with a single
lateral tooth, the presence of three gizzard plates, two of
them larger, and the moiphology of the penial complex
(with a simple unarmed penis and a single unmodified
prostate), are characteristics of this genus (Gosliner, 1994;
Eilertsen and Malaquias, 2013a).
Keen (1971, fig- 2256) assigned a shell collected off Bahia
Magdalena, Baja California, Mexico to S. interruptus.
Examination of this specimen (Figure 8; CASIZ 156489)
revealed some differences in shell moqdiology and sculp¬
ture with the other specimens of S. interruptus here
studied. In typical specimens of S. intemiptus the lip
rises over die spire and the sculpture is composed of
clearlv visible oval pits, whereas the specimen from Bahia
Magdalena has a shorter lip and die sculpture is not well
marked. The specimen from Bahia Magdalena is here
regarded as indeterminable.
Genus Cylichnium Dali, 1908
Type Species: Utriculus domitus Dali, 1889 (Caribbean),
by original designation.
Cylichnium Dali, 1908: 242.
Volvulopsis Schepman, 1913: 463. Type species, by orig¬
inal designation: Volvulopsis ancillarioides Schepman,
1913 (Indonesia).
Diagnosis: Shell external, elongated-oval, with some¬
what elevated or covered spire, in most cases with
fine spiral lines. Aperture long, narrow above, wider
below. Radula with several simple and hamate lateral
teeth. Rachidian teeth absent. Reproductive system
monaulic, with a complex female copulatory organ.
Penis unarmed.
Remarks: The genus Cylichnium was originally described
as a subgenus of Cylichnella Gabb, 1873, based on three
species, Utriculus domitus Dali, 1889 (the type species
by original designation), from Guadeloupe, Cylichnella
pizarro Dali, 1908 from Panama, and Cylichnella
atahualpa Dali, 1908 also from Panama (Dali, 1908). Sub¬
sequently, several other species were assigned to this
genus: C. matsumotoi Habe, 1955 (from Japan), C. waldae
Bouehet, 1975 (from the Atlantic), and C. olivifonnis
(Watson, 1883). Bouehet (1975) transferred Aceras
africana Fischer in Loeard, 1897 and its synonym Aceras
Page 120
THE NAUTILUS, Vol. 129, No. 3
5
Figures 1-5. Scaphander intemiptiis Dali, 1890. 1. Costa Rica (LACM 73-109), shell length 24 nun. 2. SEM photograph of the shell
microsculpture (LACM 91-133.3). 3, 4. SEM photographs of a gizzard plates (LACM 91-133.3). 5. Radular teeth (LACM 91-133.3).
fischeri Locard, 1897 to Cylichnium, based on anatomical
examination of newly collected specimens. A review of the
original description of Cylichnium shows that the type
species ( U . domitus ) has a short, conical shell, with an
aperture about 4/5 of the shell length and a pointed apex,
so it appears to be a juvenile specimen. On the other
hand, the two Panamanian species C. pizarro and
C. atahualpa are much smaller than other Cylichnium
and are most likely members of the genus Cylichna
Loven, 1846, as already suggested by Keen (1971). Valdes
A. Valdes and J.H. McLean, 2015
Page 121
Figures 6-8. Scaphander interruptus Dali, 1890. 6. Syntype (USNM 97075). 7. Specimen from Isla Lobos de Tiera, Peru (LACM
74-18), shell length 15 mm. 8. Indeterminable specimen assigned to S. interruptus by Keen (1971), collected off Bahia Magdalena,
Baja California, Mexico (CASIZ 156489), shell length 12 mm.
Figures 9-10. Scaphander interruptus Dali, 1890, anatomy
(LACM 91-133.3). 9. Penial complex; scale bar = 1 mm. 10.
Anterior portion of the digestive system; scale bar = 1 mm.
Abbreviations: bb, buccal bulb; enr, circumesophageal nerve
ring; gz, gizzard; pc, penal capsule; pn, penis; pr, prostate; rm,
retractor muscle; sg, salivary gland.
(2008) described two additional species of Cylichnium,
C. mucronatum Valdes, 2008 and C. nanum Valdes, 2008
from deep waters in the tropical Indo-Pacific, and synon-
ymized C. matsumotoi Habe, 1955 and C. surnatrense
Thiele, 1925 with C. anciUarioides.
Schepinan (1913) described the genus Volvulopsis
based on the single species V anciUarioides Schepman,
1913, from Indonesia (959-1301 m depth). Subsequent
authors regarded Volvulopsis as a synonym of Cylichnium
Dali, 1908 (see Bouchet, 1975; Valdes, 2008). The shell
characteristics of V. anciUarioides resemble those of
adult species assigned to Cylichnium by Habe (1955)
and Bouchet (1975). Cylichnium differs from Scaphander
in the presence of a female copulatory organ, wider
radula with several lateral teeth and a narrower shell
(Valdes, 2008).
Following these revisions, Cylichnium now contains
six valid species [C. domitus (Dali, 1889), C. africanum
(Fischer in Locard, 1897), C. anciUarioides (Schepman,
1913), C. waldae Bouchet, 1975, C. mucronatum Valdes,
2008 and C. nanum Valdes, 2008], all of them from deep
waters in the tropical Indo-Pacific, Japan, the Caribbean,
and the eastern Atlantic.
Cylichnium cylindrellum (Dali, 1908)
(Figures 11-18)
Scajdmnder cylindrellus Dali, 1908: 239, pi. 8. fig. 1;
Keen, 1971: 800, fig. 2255.
Shell Morphology: Shell 33 mm in length and 16 mm
in diameter (hoiotype), somewhat cylindrical, with nearly
parallel sides (Figure 11). Aperture as long as the shell.
Page 122
THE NAUTILUS, Vol. 129, No. 3
Figures 11-16. Cylichnium cylindrellum (Dali, 1908). 11. Holotype (USNM 110563), shell length 29 min. 12. Apex of a shell from
Santa Barbara County, California (LACM 95-126.3). 13. Apex of a shell from Bahia Magdalena, Baja California, Mexico (CASIZ 156490).
14. SEM photograph of the shell microsculpture (LACM 94-58.3). 15. Half-row of the radula (LACM 94-58.3). 16. SEM photograph
of a gizzard plate (LACM 94-58.3), showing several fractures produced as the consequence of the drying process for SEM examination.
wider anteriorly and constricted posteriorly. Apex sunken,
convex. Lip raising above the apex in some specimens
(Figures 11, 13), whereas in others the lip is lower than
the apex (Figure 12). Sculpture with fine spiral grooves
crossed by microscopic axial lines only present in the
spiral grooves (Figure 14). All specimens were preserved,
so information on the external coloration and shape of
the body is unavailable.
A. Valdes and J.H. McLean, 2015
Page 123
Figures 17-18. Cylichnium cylindrellum (Dali, 1908), anatomy (LACM 94-58.3). 17. Reproductive system; scale bar = 1 mm.
18. Anterior portion of the digestive system; scale bar = 1 mm. Abbreviations: am, ampulla; ag, albumen gland; bb, buccal bulb;
be, bursa copulatrix; enr, circumesophageal nerve ring; esg, external seminal groove; fco, female copulatory organ; gz, gizzard;
mg, mucous gland; pn, penis; pr, prostate; sg, salivary gland; sr, seminal receptacle.
Digestive System: The buccal mass is large and heavily
muscular (Figure 18). At its posterior end it connects to the
esophagus and two elongate salivary glands. The esopha¬
gus is short and opens into the muscular gizzard, which
contains three subequal gizzard plates. All three gizzard
plates are oval and smooth (Figure 16). The radular for¬
mula is 20 x 12.0.12 (n = 1). All lateral teeth are hamate,
with no denticulations (Figure 15). The innermost tooth
of each row is wider than the outer lateral teeth and has
a lateral prolongation on its outer edge.
Reproductive System: The reproductive system is
monaulic (Figure 17). The ampulla is large and convoluted.
It connects to the narrow and elongate post-ampullary
duct. The seminal receptacle enters die post-ampullary
duct tiiat continues distally to die common general atrium.
Also entering die common atrium are die female glands
and die bursa copulatrix (or gametolytie gland). From the
gonophore an open seminal groove runs anteriorly to die
protusible cephalic penis and die prostate. At the vaginal
opening there is a complex female copulatory organ.
Material Examined: CALIFORNIA (all specimens
collected by K. L. Smidi, R/V New Horizon), approxi¬
mately 226 km west of Point Arguello, Santa Barbara
County (34° 44' N, 123° IF W), 4100 m depdi, 1 speci¬
men, 18 February 1990 (LACM 90-162.1); (34°45' N,
123°07' W), 4100 m depth, 1 specimen, dissected, 24 June
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THE NAUTILUS, Vol. 129, No. 3
1991 (LACM 91-41.1); (34°43' N, 123°07' W), 4100 m
depth, 1 shell with dried soft parts, 2 August 1991 (LACM
91-132.4); (34° 4 7' N, 123°04' W), 4100 m depth, 1 speci¬
men, 26 February 1992 (LACM 92-109.1); (34°4T N,
123°03/ W), 4134 m depth, 1 specimen, 25 June 1992
(LACM 92-111.1); (34°38' N, 120°0T W), 4134 m depth,
1 specimen, 22 July 1992 (LACM 92-112.1); (34°43' N,
123°04' W), 4100 in depth, 1 fragment of shell, 20 Octo¬
ber 1992 (LACM 92-114.2); (34°45' N, 123°02' W),
4100 m depth, 1 fragment of shell, 24 February 1993
(LACM 93-21.10); (34°42' N, 123°Q8' W), 4100 m depth,
1 shell and a fragment of shell, 20 July 1993 (LACM 93-
43.7); (34° 39' N, 122°58' W), 4100 m depth, 2 fragments
of shell, 17 June 1994 (LACM 94-58.10); 4100 in depth,
2 specimens, 17 June 1994 (LACM 94-58.3); (34° 44' N,
123° 13' W), 4100 m depth, 1 shell, 21 August 1994
(LACM 94-59); (34°42' N, 123°09' W), 4100 m depth,
1 shell and two fragments, 14 February 1995 (LACM
95-19.2); (34° 42' N, 123°09' W), 4100 m depth, 1 specimen,
14 February 1995 (LACM 95-19.3); (34°42' N, 123°05'
W), 4100 m depth, 1 specimen, 17 February 1995 (LACM
95-20.3); (34°40' N, 123°03' W), 4100 m depth, 1 frag¬
ment of shell, 1 May 1995 (LACM 95-21.9); (34°40' N,
123°03' W), 4100 in depth, 1 specimen, 1 May 1995
(LACM 95-21.8); (34° 40' N, 123°11' W), 4100 m depth,
1 specimen, 3 June 1995 (LACM 95-126.3); (34°40' N,
123°11' W), 4100 m depth, 2 specimens, 31 May 1996
(LACM 96-38.1); (34° 40' N, 123°lT W), 4100 m depth,
1 fragment, 5 June 1996 (LACM 96-39.8); (34°41.710' N,
123° 12.570' W), 41 10 m depth, 2 August 2004, 1 fragment
of shell (SIO Ml 1388); (34°41.440; N, 123°03.694' W),
4135 in depth, 30 October 2004, 1 fragment of shell (SIO
Ml 1435); (34°4 1.440' N, 123°03.694' W), 4135 m depth,
30 October 2004, 1 shell (SIO Ml 1420); (34°41.015' N,
123°09.367' W), 4129 m depth, 26 February 2005, 1 spec¬
imen (SIO Ml 1462); (34°41.920' N, 123° 12.257' W),
4079 m depth, 21 June 2005, 1 fragment of shell (SIO
M 11488); (34°39.826' N, 123°5.241' W), 4070 m depth,
23 June 2005, 1 shell (SIO M11496). MEXICO, Off
Bahia Magdalena, Baja California (24°23' N, 113° 18.9' W),
3390-3580 m depth, 1 shell, leg. Lowenstam (CASIZ
156490). PERU, R/V USS Albatross Expedition, station
4672, 88 mi (163 km) southwest of Palominos Light, 2845
fathoms (5200 m), shell with no body parts (IIOLOTYPE,
USNM 110563).
Geographic Range: This species, originally described
from Peru, has also been found in Baja California, Mexico
(Keen, 1971) and southern California (present paper).
Remarks: Our material of Cylichnium cylindrellum
from southern California clearly matches the original
description and holotype of this species. The mid-whorl
microsculpture shows the paired spiral grooves and the
general outline of the shell is very similar to that of
the holotype. Some variability has been observed in the
southern California material, in some specimens the lip
raises slightly above the apex (LACM 95-19.2, CASIZ
156490) (Figures 11, 13), whereas in others (LACM
95-126.3) (Figure 12) it does not.
Keen (1971) recorded this species from Bahia
Magdalena, Baja California, Mexico. Keens two illustra¬
tions are a copy of the original figure and the specimen
from Bahia Magdalena (CASIZ 156490) on the right.
Examination of the material confirmed that Keens speci¬
men also belongs to C. cylindrellum.
The reassignment of this species was here conducted
based on anatomical features. The reproductive system of
C. cylindrellum has the characteristic female copulatory
organ found in other species of the genus (Bouehet, 1975;
Valdes, 2008). Also, the radula is composed of a series of
simple and hamate lateral teeth, very similar to those of tire
Atlantic species C. waldae and C. africanum (see Bouehet,
1975) as well as the Pacific species C. ancillarioides,
C. mucronatum , and C. nanum (see Valdes, 2008).
Cylichnium cylindrellum differs from other species of
the genus by its shell morphology and internal anatomy.
The shells of the Pacific species C. ancillarioides and
C. mucronatum have much narrower bases and aper¬
tures, as well as more pointed apices (Valdes, 2008). The
other Pacific species, C. nanum has a distinct columellar
tooth (Valdes, 2008), absent in C. cylindrellum. Anatomi¬
cally, C. ancillarioides and C. mucronatum have 10 and 6
radular lateral teeth, respectively (Valdes, 2008), whereas
C. cylindrellum has 12. The reproductive system of
C. mucronatum has a female copulatory organ with mar¬
ginal papillae only, whereas in C. cylindrellum die entire
organ is covered with papillae, as in C. ancillarioides
(Valdes, 2008). However, the prostate of C. ancillarioides
is much more elongated than that of C. cylindrellum,
whereas the penis is shorter and wider.
ACKNOWLEDGMENTS
We are grateful to Kent Trego for calling our attention to
collected specimens of the eastern Pacific species assigned
to Scaphander. Manuel Malaquias, Paula Mikkelsen, and
Lindsey Groves critically reviewed the manuscript.
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THE NAUTILUS 129(3):126-135, 2015
Page 126
Conservation of two endangered European freshwater
mussels (Bivalvia: Unionidae): A three-year, semi-natural
breeding experiment
Rafael Araujo
Museo National de Ciencias Naturales-CSIC
c/ Jose Gutierrez Abascal 2
28006 Madrid, SPAIN
[email protected]
Carles Feo
Quim Pou
Miquel Campos
Consorci de l’Estany
Plaga dels Estudis 2
17820 Banyoles, Girona, SPAIN
[email protected]
[email protected]
[email protected]
ABSTRACT
Freshwater mussels are among the most imperilled ol all ani¬
mal groups. The populations of the endangered Unio mancus
Lamarck, 1819 and U. ravoisieri Deshayes, 1847 (both desig¬
nated as U. elongatulus C. Pfeiffer, 1825 in the European
Habitat Directive) have declined severely over recent years
in Spain. To conserve these species in Lake Banyoles (Girona,
Spain), a total of 108,875 U. mancus and 27,423 U. ravoisieri
juveniles produced by artificial infection of larvae on host fish
were grown in a number of semi-natural, sequential breeding
systems, which involved the use of water and sediment from
their natural habitat, plus pools, plastic outdoor channels, and/
or cages. Across the tested systems, U. mancus reached a
mean length of 9.7 mm (SD±1.53) in one year and 12.4 mm
(SD±1.55) in two years; for U. ravoisieri these values were
15.8 (SD±0.76) and 21.2 mm (SD±2.45). In a experiment
adding extra food, the growth rates were much lower than
those recorded for the other systems. In October 2013, 278
2+ juveniles of U. mancus and 224 2+ juveniles U. ravoisieri
were released into the lake, increasing their original popula¬
tions by some 40% and 200% respectively. Preliminary obser¬
vations made eight months later showed that several tens of
these mussels were still alive. The large numbers of juveniles
raised in the semi-natural systems will help conserve future
generations for these bivalves in Lake Banyoles. Over the
three years of the project, 3,510 fish infected with a total of
some 500,000 gloehidia of one or the other species were also
released. After 1.5 years, hundreds of juveniles (13-35 mm)
arising from this release were detected. This is the first time
in Europe that thousands of juveniles of any endangered
freshwater mussel species have been bred in captivity without
the addition of extra nutrients, demonstrating the practicality
of low-tech and economical approaches to mussel popula¬
tion restoration.
Additional Keywords: Unio, Unio elongatulus, Unio mancus,
Unio ravoisieri, growth, juveniles, restocking, Lake Banyoles
INTRODUCTION
Freshwater mussels, or naiads (Order Unionida), are
among the most imperilled of all animal groups. Their
numbers have drastically declined due to pollution, habi¬
tat deterioration, and declining numbers of host fish
(Lydeard et al., 2004; Strayer et al., 2004). More than half
of the USAs near 300 species are now either extinct,
endangered, or threatened, and in Europe the extinction
rate for naiad populations is growing (Cuttelod et al.,
2011). In the USA, this scenario has encouraged attempts
to develop naiad captive breeding techniques — work that
has inspired similar attempts in Europe. The first docu¬
mented studies on naiad artificial reproduction and
propagation were performed on commercial species in
the USA (Lefevre and Curtis, 1912; Coker et ah, 1921;
Howard, 1922); nacre for buttons was of great economic
importance in North America at the beginning of the
20th century (Anonymous, 1914; Claassen, 1994). In
addition to providing an excellent compendium of the
natural history of freshwater mussels, these pioneering
papers summarized knowledge on mussel breeding and
cultivation that is still useful today.
The reproductive strategy of freshwater mussels
involves an obligatory parasitic stage, in which the lar¬
vae (gloehidia) attach to the external surface of a suit¬
able host and metamorphose into free-living juveniles
(Lefevre and Curtis, 1912; Kat 1984; Waehtler et ah,
2001; Araujo et ah, 2002; Rogers-Lowery and Dimock
Jr, 2006; Barnhart et ah, 2008). This, of course, is a major
problem in the development of controlled naiad breed¬
ing systems. Controlled breeding, of which the main
objective is to obtain larger numbers of juveniles from
fish infected with gloehidia than would be naturally pro¬
duced, can be carried out in captivity or semi-captivity.
Juveniles then need to be grown before their introduction
R. Araujo et al, 2015
Page 127
into natural habitats where populations have been depleted
or entirely lost.
Gum, Lange, and Geist (2011) recently published a crit¬
ical reflection on some of tire captive breeding techniques
used in Europe and the USA, with emphasis on those for
the freshwater pearl mussel Margaritifera margaritifera
(Linnaeus, 1758). The information in the latter work, and
in other seminal publications in the field (see below) was
used to develop the successful semi-natural systems for
rearing endangered freshwater mussels reported here.
Along with the process offish infection, the provision
of an adequate diet for juveniles is a key problem that
must be solved. The success of Lefevre and Curtis (1912),
Coker et al. (1921), and Howard (1922) in rearing juve¬
niles of several species was dependent on the use of the
water, food, and sediment present in the mussels’ natural
ecosystems. The idea that these elements were necessary
was confirmed many years later in Europe by Hruska
(1999), who hypothesized that the food required by
M. margaritifera juveniles comes from a healthy rhizo-
sphere. Eutrophication, contamination, and silting of the
immediate environment was deemed responsible for
the absence of available habitat, juvenile food and the
recruitment of young mussels. The success of Hruska
(1999), who grew juveniles larger than 5 cm, relied on
river bank restoration and a semi-captive breeding sys¬
tem that provided for natural feeding. In Spain, Comas
and Vails (2007) grew juvenile Unio manciis Lamarck,
1819 to reproductive age in a system involving mini¬
mum management that made use of natural water and
sediment — but not from the river were the mussels nor¬
mally lived — without any extra nutrients. However, this
work was only published as an internal document of
the Catalan Regional Government. Other authors have
developed more controlled systems (with more or less
success) inspired by systems used in marine bivalve
aquaculture, providing extra food in the form of algae
(Hudson and Isom, 1984; Gatenby et al., 1996; Gatenby
et ah, 1997; O’Beirn et al, 1998; Henley et al, 2001;
Araujo et al, 2003; Gatenby et al, 2003; Beck and Neves,
2003; Liberty, 2004; Guyot, 2005; Jones et al, 2005;
Barnhart, 2006; Kovitvadhi et al, 2006; Liberty et al,
2007; Eversole, 2008; Kovitvadhi et al, 2008). These more
controlled systems have inspired the main cultivation
programs for M. margaritifera in Europe (Gum et al,
2011; Eybe, etal, 2013).
Although the use of algae has sometimes been suc¬
cessful in the rearing of presumably healthy juveniles,
Nichols and Garling (2000, 2002) report the main dietary
source of carbon for naiads living in rivers and lakes to be
bacterial. Algae do, however, appear to provide key nutri¬
ents such as vitamins and phytosterols. Much remains to
be learned about the diet of juvenile naiads in natural
environments and in captivity. As part of the LIFE 08
NAT/E/000078 “Estany Project” which is dedicated to
restoring the native aquatic fauna of Lake Banyoles
(Girona, Spain; a Natura 2000 site), semi-natural systems
were developed to rear two endangered European species
of mussel: U. mancus and U. ravoisieri Deshayes, 1847
(both designated as U. elongatulus C. Pfeiffer, 1825 under
the Habitat Directive, the main European law for species
conservation). The first of these species lives in Spanish
and French Mediterranean rivers; the limit of its eastward
range, however, remains unknown (Araujo et al, 2009a;
Pile and Puillandre, 2013). It is considered “near threat¬
ened” by the IUCN (Cuttelod et al, 2011). The second
species, U. ravoisieri, is restricted to just two localities in
Spain (Araujo et al, 2009a; Khalloufi et al, 2011). The
Lake Banyoles populations of both species have been in
severe decline in recent years, a consequence of the pro¬
liferation of invasive predatory fish. The five native fish
species ( Anguilla anguilla, Gasterosteus aculeatus, Barbus
meridionalis, Squalius laietanus and Solaria fluviatilis)
have been partially eliminated and replaced by the exotic
Microjiterus salmoides, Lepomis gibhosus, Cyprinus
carpio, Perea fluviatilis, and Sander luciperca (Moreno-
Amich et al, 2006).
It was hypothesized that cultivation systems connected
to the natural habitat of these naiads would provide the
unknown natural food required by the juveniles. This
paper presents the first large-scale attempt to raise juve¬
niles of endangered naiad species in Europe, using water
and sediment from the mussels’ natural environment.
MATERIALS AND METHODS
The work was performed at the Naiad Breeding Labora¬
tory at Banyoles (Girona, Spain). This field station is
located 500 m from Lake Banyoles (which lies in the Ter
Basin) and receives a constant supply of hike water.
Although in preliminary work several fish species were
tested (including Solaria fluviatilis, Luciobarbus graellsii,
Phoxinus phoxinus, and Tinea tinea ) as hosts for the mus¬
sel larvae, Barbus meridionalis Risso, 1827 and Squalius
laietanus Doadrio, Kottelat and Sostoa, 2007 were chosen
since these species are native to the lake. The two naiad
species raised were U. mancus and U. ravoisieri, both of
which are native to the lake basin. The number of speci¬
mens of fish and naiads involved differed over the three
years of the project (2010-2013) (Table 1).
The fish used as hosts were collected from the Rivers
Ter, Terri, Brugent, Llemena and Osor (all in the Ter
Basin), 1^ weeks before infection with glochidia. Fol¬
lowing capture these fish were maintained in outdoor
pools (1,600 L) that received a flow of lake water.
The gravid naiads used came from die lake (U. ravoisieri)
or its effluents (U. mancus); these were collected over the
spring (the water temperature of the lake was moni¬
tored six times per day using a submerged thermometer
(Thermotronie Geteeh Innova) to determine the water
temperature suitable for the reproductive cycle to begin).
These mussels were maintained in indoor aquaria for the
collection of released glochidia (these remained viable for
48-72 h); they were then returned to their natural habitat.
Mature glochidia were collected with pipettes and placed
in aerated water for 5 min in a plastic Tupperware vessel
(500 ml) containing a single fish. Infected fish were then
Page 128 THE NAUTILUS, Vol. 129, No. 3
Table 1. Fish and naiad specimens used.
kept in aerated conical tanks (180 L) at a temperature
ranging between 15 and 22 °C; only fish infected over
the same 3-day period were placed in the same tank.
These tanks were equipped with biological and mechani¬
cal filters. The fish were provided feed every day until
three days before the release of juveniles. The number of
degree/days needed for metamorphosis to occur was
based on Araujo et al. (2005) and Reis et al. (2013). The
water in these tanks was filtered (filter mesh size 200 pm)
and renewed three times per day.
Two days before juvenile release, a 200 pm mesh col¬
lector was installed in the water circuit to retrieve them.
This collector was checked and juveniles collected once
or twice daily until no more were found.
The juveniles obtained were observed under a binoc¬
ular microscope to check their viability; those deemed
alive were then assigned to either (Figure 1):
System 1) Plastic tray (240 cm long, 60 cm wide, 17 cm high)
containing 5 cm-deep un-sieved lake sediment (water
depth to sediment surface = 9 cm) supplied with a con¬
stant water flow. This system was maintained indoors under
Figure 1. Schematic diagram of the experimental design.
normal photoperiod conditions. Juveniles of both naiad spe¬
cies were placed togedier in this system. Su nival was checked
periodically and a random sample of specimens measured
until the emptving of the system in October 2013, or:
System 2) Three outdoor cubic pools (150 x 150 x 150 cm)
containing 20 cm-deep un-sieved lake sediment, (water
depth to sediment surface = 110 cm) supplied with a
constant water flow. Unio rnancus juveniles were released
into pool 1 and U. ravoisieri into pool 2 in 2011 and 2012;
in 2012 the two species were also released together in
pool 3. Survival was checked periodically and a random
sample of specimens measured until pools 1 and 2 were
emptied and the sediment filtered in February 2013, and
until the same was performed with pool 3 in October 2013.
[Note: new generations of U. mancus and U. ravoisieri are
currently being raised in pools 1 and 2],
The surviving juveniles from System 2 were seeded into
the following subsystems in February, April and October
2013 (Figure 1):
a) plastic tray containing lake sediment (depth 30 cm), placed
at the bottom of the lake (depth 2 m).
b) outdoor plastic channels (6 m long, 50 cm wide, 28 cm high)
containing 10 cm-deep lake sediment, supplied with a 1 L/s
constant water flow (water velocity 50-100 cin/min) from
the lake. Survival was checked periodically and a random
sample of specimens measured each time until the end of
the project.
c) plastic cages (30 x 15 x 15 cm; mesh 1 x 1 cm) placed on
the lake bottom (depth 2 m) (only U. mancus).
d) cages (30 x 15 x 15 cm; mesh lxl cm) placed in the lake
water column (depth 1 m) (only V. mancus).
e) cages (100 x 25 x 25 cm; mesh 1 x 1 cm) placed on the
lake bottom (depth 2 in).
f) directly on the bottom of the lake in areas with no vegetation
(only specimens produced in 2011 that reached a size of at
least 2.5 cm).
Although the aim of the present work was to demonstrate
the effectiveness of the natural diet (food from water and
sediment) in raising the mussels, an experiment involving
an external food source was also designed.
In 2012, 2000 U. mancus juveniles were divided into
5 series of 200 (two replicas) in Tupperware vessels
R. Araujo et al., 2015
Page 129
containing the following: 1) 400 ml lake water; 2) 400 ml
lake water plus 0.8 ml of dehydrated commercial algae
(66.5 x 10J cells of Nannochloropsis, 11.08 x 109 of
Phaeodactylum and 1.25 x 109 of Tetraselmis ); 3) 400 ml
lake water plus a mixture of 2.4 ml of natural algae,
leafs and macrophyte extract; 4) 400 ml lake water plus
0.8 ml of leaf extract and 0.8 ml of dehydrated com¬
mercial algae; and 5) 400 ml kike water plus 0.8 ml of
biofilm extract.
Leaf extract was obtained by washing macrophytes
and the leafs and stems of land plants from the habitat
around the lake; the suspension obtained was filtered
and frozen in doses of 0.8 ml. Biofilm extract was pre¬
pared from 200 g of the biofilm growing on the walls of
the outdoor pools in 2 1 of lake water; this was also
filtered and frozen in doses of 0.8 ml.
All containers were cleaned once per week; dead juve¬
niles were removed, live juveniles were measured, and
the food renewed. The experiment ran between May
2012 and July 2013. The surviving juveniles were trans¬
ferred into the lake in their own cage.
We performed a one-way ANOVA to compare the
growth among some of the different systems.
RESULTS
In 2011-2013, the release of gloehidia by U. nuincus and
U. ravoisieri occurred between April 1 1 and J uly 22, and
April 27 and July 24, respectively (minimum lake temper¬
ature 13°C) (Figure 2). The total number of U. nuincus
and U. ravoisieri juveniles released by the host fish for use
in the different systems was 108,875 and 27,423 respec¬
tively (Table 2). The release of juveniles from the host
fish occurred between days 7 and 33 post infection (PI)
in U. nuincus and 8 and 26 PI in U. ravoisieri, depending
on the water temperature (representing a minimum 145
and maximum 521 degree/days for both species taken
together) (Table 3).
All the systems used in tliis study successfully raised
mussels, but with marked differences in survival and
growth rates. All the juveniles in the indoor plastic tray
(System 1) in 2011 died due to a hardware malfunction,
but in 2012 System I was capable of maintaining live
juveniles (Figure 3). Those that survived one year (12%)
reached a mean length of 4.5 mm (SD±1 .35, n=102)
and a maximum of 8 mm. The total survival rate at day
520 was 3.6%, with 574 live juveniles recovered (mean
size 6.5 mm, minimum 3.5 mm, and maximum 11.5 mm)
for the 16,322 originally seeded.
The best results were obtained with the outdoor
pools (System 2). In 2011, 3,000 U. mancus juveniles
were placed in pool 1 and about 1,800 U. ravoisieri in
pool 2. In 2012 these figures were 9,380 U. mancus
and 5,005 U. ravoisieri in pools 1 and 2 respectively.
In addition, 16,658 U. mancus and 2,412 U. ravoisieri
were mixed in pool 3. [Note: in 2013, once the juveniles
from pools 1 and 2 had been removed and the pools
cleaned, 36,140 U. nuincus and 4,318 U. ravoisieri were
placed in them respectively].
Table 2. Number of juveniles obtained from host fish, and numbers seeded in the different systems. Uma = Unio mancus. Ura =
U. ravoisieri.
2011
2012
2013
Total
Uma Ura Uma
Between summer and autumn 2012 there was a seri¬
ous loss of juveniles in pool 1 probably due to the over¬
growth of benthic algae; this problem was solved by
cleaning the floor of the pool and covering the top. On
day 350, 100 juveniles (>1 cm) from pool 1 were trans¬
ferred to a tray in the lake (System 2 a), and on day 575,
150 (1-1.5 cm) were transferred to tire small cages
described for System 2 e and d. On day 617, the pool was
siphoned and a large number of empty shells detected.
A total 493 remaining live juveniles were placed either in
the outdoor plastic channels (System 2 b) or a large cage
on the lake bottom (System 2e) (Figure 4A). On day 697,
100 juveniles (2-2.5 cm) from pool 2 were transferred to
a large cage in the lake (System 2 e), and the remaining
237 distributed between the outdoor plastic channels
(System 2 b) and a large cage in the lake (System 2 e)
(Figure 4B). On day 325, 129 juveniles were transferred
from pool 3 to a large cage in the lake (System 2 e). On
day 500, the pool was siphoned and the 1,459 remaining
juveniles detected transferred to another large cage in
the lake (System 2 e).
The estimated survival rate at 1 year for the 201 1 gen¬
eration of U. rnancus in pool 1 before any distribution into
any subsystem was 77%. However, this fell to 20% after
two years. The juveniles reached a mean length of 9.7 mm
(SD±1.53, n=220 measured) after one year, and 12.4 mm
-r— 'Mean length
© Max length
® Min length
O Sample size
Figure 3. Growth of the juveniles seeded in 2012 in the indoor plastic tray (System 1 ) (both naiad species mixed). N indicates the
number of measured juveniles.
R. Araujo et al., 2015
Page 131
A Days after sowing
— •—Benthic cage in lake i2c)
—♦““Big cage in lake (2e)
Floating cage in lake (2d)
—♦—Outdoor channel (2b)
■ - Pool 1 _
Figure 4. Juvenile growth in the different systems. A. Unio mancus. B. U. ravoisieri.
(SD±1.55, n=191 measured) after two (Figure 4A). The
generation of juveniles seeded in 2012 were killed by the
algal growth on the bottom.
In pool 2, the survival rate of the U. ravoisieri 2011
generation was 33% for the first year, and 18% for the
second. The mean length reached at the end of the first
and second years was 15.8 mm (SDT0.76, n=22 mea¬
sured) and 21.2 mm (SD±2.45, n=331 measured)
respectively (Figure 4B). The survival rate of the 2012
generation was 0.7%.
On day 500, the survival rate of the mixed population in
pool 3 was 8.2% and the mean length 11 mm (SDT2.95,
n=300 measured).
The survival and growth rates strongly increased when
juveniles reached two years (2+) of age and a size of
1-1.5 cm in U. mancus (Figure 4A) and 2-2.5 cm in
U. ravoisieri (Figure 4B). Of the 493 U. mancus 2+
juveniles in pool 1, 200 were transferred to a cage in the
lake (System 2 e) and the rest to an outdoor plastic chan¬
nel (System 2 b). After 170 days, the survival rates were
100% in the plastic channel and 93% in the cage, and the
corresponding mean lengths were 2.1 (SD±2.33) and
1.8 cm (SD±2.3). In a similar experiment with U. ravoisieri
2+, the same survival rates were recorded but the mean
lengths were much greater: 2.5 cm (SD±3.31) and 2.7 cm
(SD±2.3) respectively. In October 2013 (i.e., at 870 days
of age) a small portion of the juveniles in the outdoor
plastic channels were maintained there; the rest were
placed in the lake, both in cages (System 2 e) and free
(System 2 f). In May 2012, prior to the use of the cages in
the lake, 100 U. mancus juveniles of 1 cm from pool
1 were put in an open tray with sediment and placed in
the lake (System 2 a). After three months the tray was
removed; no living specimens were found but only broken
shells, suggesting that they had fallen prey to fish and
crayfish. However, in October 2012, upon inspection of
the mud underneath where the cage had lain, a mussel
was found measuring 28.1 mm (this mussel was marked
for identification purposes), and in June 2013, another
was found measuring 38 mm. This means that the first
juvenile grew 17 mm in the five months since the cage
was placed in the lake (May 2012) and the second 28 mm
in 13 months. However, this subsystem was no longer
used given the poor results obtained.
The survival rate at 295 days for the 150 U. mancus
juveniles from pool 1 in the small cages in the water
column and at the bottom of the lake (Systems 2 c
and d), was very high at 83% and 86%, respectively.
Growth, however, was greater in the cage on the
bottom (System 2 c) (F=93.7, p< 0.001) (Figures 4A,
B). Indeed, the specimens in the water column cages
were covered in algae and some showed deformities.
The large, bottom-placed cages (System 2 e), though
successful (Figures 4 A, B) were difficult to handle.
All the cages became covered in calcified algae during
spring and summer, blocking the mesh and thus reduc¬
ing oxygen and water flow. In October 2013, all the
juveniles from all these cages were removed, labeled
and placed once more in the kike, either in large cages
(System 2 e) or free (System 2 f).
Page 132
THE NAUTILUS, Vol. 129, No. 3
Over the three years of the project, 3,510 infected fish
carrying an estimated total 500,000 U. mancus and
U. ravoisieri glochidia were released into the lake and
its effluents (Table 1). In May and July 2013 hundreds of
1.5 year-old (13-35 mm) juvenile mussels were observed
in these outflows.
The results suggest that U. mancus reaches 9.7 mm by
the end of its first year and 12.5 mm at the end of the
second (taking all System 2 subsystems together and
excluding System 1). For U. ravoisieri , these values are
15.8 and 21.2 mm. However, the growth rate is not con¬
stant over the year; growth stops between November and
March (Figures 4A, B).
In the experiments involving the provision of extra
food, one replica was followed for 400 days and the other
350. Although growth rates were reduced (maximum
2 mm in one year when provided with commercial algae
and leafs; Figure 5) compared to the above tested sys¬
tems, all survivors were placed in a large cage on the
bottom of the lake (System 2e) in October 2013. At
350 days from the beginning of the experiment, only
two of the five series had juveniles alive, both in the two
replicas, the one with algae and the other with algae and
leaf extract. The growth between the replicas didn’t have
significant differences (p>0.02), but it was different
between the two series. The juveniles fed algae and leaf
extract grew more than the others (F=35.61, p>0.001).
However, the growth rates of the U. mancus juveniles
of the experiment were less than the ones of the pool
1-system 2 (F=1575.6, p>0.001) or the plastic tray-
system 1 (F=271.9, p>0.001).
In summary, the numbers of live juveniles (Figure 6)
raised were: U. mancus: 218 2+ and 43,700 0+;
U. ravoisieri: 100 2+, 64 1+ and 13,400 ()+, plus a mix¬
ture (unknown proportions) of 2,304 1+ U. mancus and
U. ravoisieri raised in pool 3.
A total of 278 and 224 2+ juveniles of U. mancus
and U. ravoisieri , were released free into the lake, repre¬
senting improvements of 40% (estimated population
1,000±500) and 200% (estimated population 110±50)
of their original populations (M. Campos, pers. observ.).
DISCUSSION
Seminatural breeding efforts to rescue endangered pearl
mussel populations can result in adverse effects such as
genetic drift and selection (Geist, 2010), so they should
only be considered as an emergency measure. The goal
of this work was to obtain large numbers of juveniles
of two endangered freshwater mussels, U. mancus and
U. ravoisieri, raising them in a system involving water
and sediment from their natural environment. The sequen¬
tial systems tested maintained juveniles in pools or plastic
channels at a field station, before releasing them into the
wild. The large numbers of 1-3 years old juveniles main¬
tained at the field station and in the lake offers hope for
these endangered species. This is the first time in Europe
that thousands of juveniles of any endangered freshwater
mussel have been bred in captivity for three years without
the addition of extra nutrients.
Although thousands of viable juveniles were raised,
mortality was high, especially during the first year of life.
The mortality recorded in the pools in 2012 was probably
caused by algal overgrowth and subsequent anoxia. This
can be avoided by covering the pool and/or siphoning
and filtering the upper layer of die sediment, and
renewing it after two years. The use of outdoor plastic
channels with a slow water current — but fast enough to
prevent deposition — can also be used. Although no nat¬
ural mortality data are available for these species, juve¬
nile mortality is commonly high among freshwater
mussels (Young and Williams, 1984). The juveniles that
reached two years seemed to show increased viability.
The raising of mussels for one or two years in pools and
then transferring the survivors to outdoor channels or
bottom-lying cages in the wild, would likely provide very
good results (Figures 4A, B).
R. Araujo et al., 2015
Page 133
Figure 6. The juveniles bred. A. Empty shells of Unio martens and B. U. ravoisieri 2+ from System 2e. Scale bar: 1 cm. C. Live
juveniles labeled for release into the lake.
The main problem encountered with the use of the
cages (of both sizes) was the growth of calcified algae
(probably a consequence of the hard water of Lake
Banyoles); this blocked the mesh and isolated the mus¬
sels from the environment. This could be avoided by
regular cage cleaning. This blocking was probably the
reason why the U. ravoisieri juveniles grew better in the
outdoor plastic channels (System 2b) than in the cages
(System 2e) (Figure 4B) (F=93.7, p<0,001). At 2 or
3 years of age, the mussels raised in cages are mature
enough to be released into the wild.
The survival of the juveniles freely seeded in the lake
(System 2f) remains to be fully studied, but preliminary
observations (June 2014) suggest that they are still alive.
The release of infected fish was also shown to be a suc¬
cessful way of seeding the environment, at least in the
lake effluents. This could be an easy and effective means
of repopulating depleted areas but it would also be very
difficult to monitor, and success might vary between one
water system and another.
The results obtained with the indoor plastic trays (Sys¬
tem 1) were not as good as those obtained with the pools
(System 2 and its subsystems); survival and growth were
much slower (Figure 3) (F=66.89, p<0.001). However,
these trays provide an easy way of maintaining juveniles
for study and handling in the laboratory.
In the experiment in which extra food was added
(which was very laborious), the growth rates were
much lower than those recorded for the other systems,
although some juveniles did survive for more than one
year (Figure 5).
The fish populations of Lake Banyoles have com¬
pletely changed over the last century. The five native fish
species ( Anguilla anguilla, Gasterosteus aculeatus, Barbus
meridionalis, Squalius laietanus, and Salaria fluviatilis )
have been partially eliminated and replaced bv the exotic
Micropterus salmoides, Lepomis gibbosus, Cyprinus carpio,
Perea fluviatilis and Sander luciperca (Moreno-Amich
et ah, 2006). This led to the vanishing of the formerly
abundant populations of U. mancus and U. ravoisieri.
Restocking with native fish and naiads using the pres¬
ent systems is vital for the survival of these endangered
molluscs in the lake.
In recent years, our knowledge of the reproductive
biology of several Unio species has greatly increased
(Aldridge and Mcivor, 2003; Araujo et til., 2005; Vincentini,
Araujo et ah, 2009b; Reis et ah, 2013); this information
could be used to better conserve these endangered fresh¬
water mussels. The present results increase our knowl¬
edge of the reproductive strategy of U. mancus and
U. ravoisieri. Several species of fish have been reported
as successful hosts for the production of U. mancus
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THE NAUTILUS, Vol. 129, No. 3
juveniles (Araujo et al, 2005), to which B. meridionalis
and T. tinea (results not shown) can now be added. In
U. ravoisieri, the breeding season of which was unknown,
we now have shown that glochidia are released between
April and July, and B. meridionalis, S. laietanus, L. graeUsi,
S. fluviatilis, T. tinea and P. phoxinus may be valid
hosts, as shown in this work and preliminary testing.
Although much remains to be learned regarding the
diet of freshwater mussels, the present results show that
some species can be cultured in semi-captivity using only
the water and sediment from their natural ecosystem.
The survival rate of these juveniles in the following years
will give a better idea about the viability of these breed¬
ing systems.
ACKNOWLEDGMENTS
This work was funded by the LIFE 08 NAT/E/000078
“Estany Project”. The authors thank Ramon Casadevall
and Arnau Juscafresa for their help in capturing the fish
used in this work.
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THE NAUTILUS 129(3): 136^139, 2015
Page 136
A new species of Glyphostoma (Gastropoda: Clathurellidae)
from the Gulf of Mexico
Emilio F. Garcia
1 15 Oakcrest Dr.
Lafayette, LA 70503 USA
Efg21 [email protected]
ABSTRACT
Glyphostoma coronaseminale new species is described and
compared with the Western Atlantic species G dentiferum
Gabb, 1882, G. epicasta Bartsch, 1934, G. gabbii (Dali, 1889),
G. golfoyaquense Maury, 1917, and G. herminea Bartsch, 1934.
INTRODUCTION
Two decades of research campaigns conducted in the
Gulf of Mexico by the Biology Department at the Uni¬
versity of Louisiana at Lafayette (ULL) have led to many
molluscan discoveries (e.g., Garcia, 2003, 2005, 2006).
The cruises have utilized the R/V Pelican, a ship man¬
aged by the Louisiana Universities Marine Consortium
(LUMCON).
In September, 2014 a cruise that terminated west of
Dry Tortugas, Florida, was conducted by ULL marine
biologists Drs. Darryl Felder and Suzanne Fredericq.
This was the last of five cruises executed under the des¬
ignation Gulf of Mexico Research Initiative (GoMRI).
Two types of dredges were used, the standard box
dredge and the Benthic Skimmer, a large, more efficient
dredge specially designed for soft bottom (see Garcia,
2007a). This dredge was in use when the holotype of the
Glyphostoma species described herein was collected.
The genus Glyphostoma was erected by Gabb for the
fossil taxon G. dentiferum ; it is known from the Pliocene
and Miocene of the Caribbean and the southeastern
United States, but it is also part of the recent fauna.
Powell (1966: 115) considered this taxon to be mostly
American. Although it had been used for recent Indo-
Pacific species, most of these could be assigned to either
Etrema or Lienardia. Nevertheless, many Recent Indo-
Pacific species have been assigned to Glyphostoma
(Worms, 2014).
In the Western Atlantic, the genus Glyphostoma can
only be confused with Lioglyphostoma Woodring, 1928,
which differs by the lack of labral denticles and the
“growth wrinkles on the anal fascicle” (Woodring, 1928:
193). These “wrinkles”, characteristic of the Glyphostoma
species described so far, are lacking in the species
described here; they have been substituted by well-defined
nodes, therefore differentiating the sculpture of the anal
fasciole from till oilier Western Atlantic Glyphostovm.
SYSTEM ATICS
Family Clathurellidae II. Adams & A. Adams, 1858
Genus Glyphostoma Gabb, 1872
Type Species: Glyphostoma dentiferum Gabb, 1872,
by monotypy.
Glyphostoma coronaseminale new species
(Figures 1-8)
Diagnosis: A white shell with a wide peripheral pale-
yellow band on apical whorls and a second, anteriorly
positioned band of the same color on last whorl; with a
constricted anal fasciole ornamented with spirally extended
nodes, and a parietal wall showing a nodose callus at
entrance of anal fasciole, a posterior denticle, two cen¬
trally positioned plicae, and an anterior denticle followed
by increasingly smaller, at times very weak, nodules.
Description: Holotype (Figures 1-6) 23.2 mm in length,
strong, fusiform (length/ widdi ratio 2.64). Protoconch
(Figure 4) conical, of approximately 3.25 smooth, yellow¬
ish whorls; first whorl minute, translucent; following whorls
developing a sub-medial keel of increasing strength; a
second, short keel appearing anteriorly, just above suture,
towards termination of last whorl. Transition to teleoconch
indicated by a shallow, strongly sinuous scar, a change in
coloration from yellowish to white, and the beginnings of
tire adult sculpture, Teleoconch of 7.5 convex, subsuturally
constricted whorls, creating a shoulder. Suture strongly
impressed on early whorls, narrowly channeled on later
whorls, crenulated by a row of strong nodes on anal
fasciole (Figure 6). Axial sculpture of strong opisthocline,
rounded ribs of even size, as wide as interspaces; 10 ribs
on first whorl, progressively increasing to 24 on last whorl,
almost reaching anterior end before evanescing; last two
E.F. Garcia, 2015
Page 137
Figures 1-16. Glyphostoma species. 1-8. Glyphostoma coronas eminale new species. 1-6. Holotype USNM 1274997 west of
Dry Tortugas, SW Florida: 25°31.091/ N, 84°28.391' W to 25°27.939' N, 84°27.145' W; in 352-361 m, 23.2 mm. 7-8. Paratype
USNM 12/4998, soutliwest of Key West, Florida, in 84 in, 17.2 mm. 9-10 Gluyphostoma golfoyaquense Maury, 1917, type figure
(after Maury), Rio Cana, Santo Domingo, 20 mm. 11-13. Glyphostoma gabhii (Dali, 1889). Syntype, USNM 87410, off Barbados,
in 300 in, 17.5 mm. 14-16. Glyphostoma dentiferum Gabb, 1872. Leetotype, ANSP IP2910, fossil, no data, 32.1 mm.
Page 138
THE NAUTILUS, Vol. 129, No. 3
adapertural ribs almost reaching anterior end. Spiral
sculpture of strong cords; cords developing heavy, spirally
extended nodes as they cross axial elements; three cords
on first whorl, progressively increasing to 6 by penultimate
whorl; three peripheral cords strongest; a constricted
hand of strong, spirally extended nodes on anal fasciole
appearing from earliest whorls; nodes not necessarily coin¬
ciding with axial cords, which are undercut by subsutural
constriction; approximately 30 nodes on last whorl. Aper¬
ture (Figure 2) narrowly elongate, 12.2 mm in length, with
short, tapering anterior canal slightly recurved to die right;
outer lip strengthened by an abaperturally concave varix
which terminates posteriorly in a deep, U-shaped sinus;
varix crossed by 18 spiral cords, axially incised by a strong
indentation which creates a secondary, thinner labrum; a
third labral element, terminating in bifurcate denticles
(broken at either side in the holotype), projects out of this
secondary labral element (Figure 5). Inner labrum with
8 strong denticles; posterior denticle slightly stronger,
slightly callused towards sinus. Parietal wall (Figure 2) with
a moderately strong, nodulose callus at sinus entrance;
central parietal wall with a small posterior denticle,
followed by two strong plaits that continue into the inner
aperture, an anterior bifurcate denticle, and a series of
barely discernible nodes. Shell color white, with a pale-
yellow peripheral band the width of 3 spiral cords on apical
whorls and 5 on last whorl; a second sub-peripheral band
on last whorl; coloration somewhat stronger in interstices.
Type Material: Holotype USNM 1274997, length
23.2 mm; width 8.8 mm (Figures 1-6). Paratype USNM
1274998, length 17.2 mm, width 7.1 in (Figures 7-8).
Type Locality: West of Dry Tortugas, SW Florida:
25°31.09T N, 84°28.39T W to 25°27.939' N, 84°27.145' W;
in 352-361 m,
Distribution: Southwest of Key West to west of Dry
Tortugas, southern Florida, in 84-361 m.
Habitat: Glyphostoma coronaseminale new species
inhabits relatively deep water. The holotype was dredged
in a sand bottom with broken shells. Other key species
dredged in the same haul were a live specimen of
Bathytoma viabmnnea Dali, 1889 and empty specimens
of Bartschia frumari Garcia, 2008.
Etymology: A compound word from the Latin corona
and seminale (“crown of seeds”); in reference to the
characteristic spiral row of nodules that is present at the
anal fasciole.
DISCUSSION
The paratype of Glyphostoma coronaseminale (Figures 7-8)
is bleached out, faintly showing the color bands; its central
parietal wall shows the posterior denticle, the two sub¬
sequent plaits, and the anterior denticle, but the following
anterior nodes are stronger than those of the holotype. As
the latter was inhabited by a hermit crab, the weaker
nodes may be a result of erosion The varical structure of
the labrum of the paratype shows tire same three stages of
growth as the holotype, including the thinner, but strong,
bifurcate denticles at the edge.
The specimens identified as Glyphostoma golfoyaquense
Maury, 1917 in Kaieher’s card 3882 (1984), and in
Williams’s (2005) image 5115 are G. coronaseminale.
Glyphostoma golfoyaquense (Figures 9-10) is narrower,
has fewer, wider axial ribs that become narrower and
bifurcate at anterior half of last whorl, has a differently
structured anal fasciole (Figure 10), and different denti¬
tion on parietal wall. The maximum reported size for
G. golfoyaquense is 20 mm (Rosenberg, 2009).
The size, parietal dentition, and numerous, even-size axial
ribs separate this new species from several other Western
Atlantic Glyphostoma. Glyphostoma pilsbryi Schwengel,
1940, which inhabits waters of the Gulf of Mexico (Garcia,
2007b) grows only to 9.5 mm, is relatively wider, and has
fewer, thicker axial ribs. Glyphostoma epicasta Bartsch,
1934 has a more elongated, milky white shell with a longer
siphonal canal, different structure of the anal fasciole, and
different dentition on parietal wall. It grows larger than
the new species, to a maximum reported size of 31 mm
(Rosenberg, 2009). Glyphostoma herminea Bartsch, 1934
grows to a maximum size of 21 mm (Rosenberg, 2009),
is yellowish white in coloration, has fewer, wider axial ribs,
a more sloping, less constricted, differently sculptured anal
fasciole, and different dentition on parietal wall.
The new species is most similar to Glyphostoma gabbii
(Dali, 1889) and Glyphostoma. dentiferum Gabb, 1872.
Glyphostoma gabbii (Figures 1 1-13) has a wider aperture,
fewer axial ribs that are narrower than the interspaces, a
subsutural ornamentation of numerous arched axial riblets
crossed by undulating spiral threads (Figure 13), axial
cords on last whorl that bifurcate anteriorly, and different
denticle structure on parietal wall (Figure 12). It has a
maximum reported size of only 17.5 mm. Glyphostoma
dentiferum (Figures 14-16) has fewer, stronger axial ribs
on apical whorls, an anal fasciole sculptured with numer¬
ous arched axial riblets (Figure 16), differently structured
denticles on parietiil wall (Figure 15), and a longer shipho-
nal canal. It grows larger than G. coronaseminale, with a
maximum reported size of 32 mm (Rosenberg, 2009).
The bifurcated denticles of the projected third ele¬
ment of the labrum of Glyphostoma coronaseminale
(Figure 5), have not been reported for any western
Atlantic Glyphostoma. Notwithstanding the broken ends
of this third element in the holotype, it is not at all
fragile; the damage seems to have been cause by crab
predation. It is possible that this third labral element is
more fragile in other Glyphostoma species and has not
been reported because of damage.
ACKNOWLEDGMENTS
My thanks to my colleagues Drs. Darryl Felder and
Suzanne Frederieq, Biology Department at ULL, for
inviting me to join them on the GoMRI project and to
E.F. Garcia, 2015
Page 139
Linda Ward, National Museum of Natural History,
Smithsonian Institution, and Paul Callomon, Collection
Manager, Academy of Natural Sciences of Drexel
University, Philadelphia, for providing the images of the
type material of Glyphostonia gabhi and G. dentifemm
respectively. The GoMRI cruises were conducted with
grants from British Petroleum.
REFERENCES
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Garcia, E.F. 2006. Six new species of mollusks (Gastropoda:
Cerithioidea, Buccinoidea, Muricoidea) from Bahia de
Campeche, southwestern Gulf of Mexico. Novapex 7(4): 77-89.
Garcia, E.F 2007a. Results of deep-water dredging in the Gulf of
Mexico using die “Bendiie Skimmer”, and report on several
geographic extensions, including two species not previously
reported in die western Adantic. The Festivals 39: 13-18.
Garcia, E.F. 2007b. Report on mollusks collected in a dredging
expedition to Bahia de Campeche, southwestern Gulf of
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Kaicher, S.D. 1984. Card Catalogue of World-wide Shells. Pack
#39 - Turridae Part I. cards [i-ii], 3882-3987. S.D. Kaicher,
St. Petersburg.
Maury, C.J. 1917. Santo Domingo type section and fossils. Part 1.
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Powell, A.W.B. 1966. The molluscan families Speightiidae and
Turridae: an evaluation of the valid taxa, both Recent and
fossil, with lists of characteristics species. Bulletin of the
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Adantic Marine Mollusca. [WWW database (version 4.1.1)]
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THE NAUTILUS 129(3): 140-141, 2015
Page 140
Research Note
First evidence for deep-sea hot venting
or cold seepage in the Ross Sea
(Bivalvia: Vesicomyidae)
Two large dead valves of an undescribed clam of the family
Vesicomyidae were recently collected by a toothfish (family
Nototheniidae) longline vessel and returned to NIWA by a
Ministry of Primary Industries (MPI) observer. Using the
Commission for the Conservation of Antarctic Marine
Living Resources Identification Guide (CCAMLR 2009),
the observer reported that the valves had a distinct sulphur
smell, which, along with the location (Figure 1), suggests
that the shells may have originated from or near an active
seep and that living clams may still be there. The most
complete valve (Figure 2) is very large (280 mm), and both
170°E
180°
150°E 160°E 170°E
170°W
Figure 1 . Location of vesicomyid elain sampled in the Ross Sea.
180
160°W
B.A. Marshall and D. Tracey, 2015
Page 141
Figure 2. Vesicomyid clam from Ross Sea. Shell length = 280 mm.
are clearly old, blackish-stained, and with a sparse cover¬
ing of encrusting bryozoans and calcareous polychaete
tubes. Vesicomyid clams are found globally in chemosyn-
thetic habitats using hydrogen sulfide for metabolism via
symbiotic bacteria (Krylova and Moskalev, 1996; Krylova
et al., 2010). While vent and seep species have been
recorded previously from the Antarctic Region (Domack
et al, 2005; Rogers et al., 2012), no vent or seep mollusks
have been reported to date from the Ross Sea.
Apart from the enormous size, the Antarctic species is
characterized by its exceptionally long ligament (length
about 56% of shell length). In size, strongly posteriorly elon¬
gate shape, and general features of the shell, it most closely
resembles some species of Abyssogena Krylova, Sahling,
and Janssen, 2010, particularly A. pkaseolifonnis (Metivier,
Okutani, and Ohta, 1986) and “ Ectenagena ” extenta
(Krylova and Moskalev, 1996) (lengths up to 180 mm and
235 mm: Krylova et al., 2010). These species have been
found, respectively, in the Japan, Kurile, and Aleutian
trenches at 4550-6329 m, the Gull of Alaska, the Monterey
Canyon, the Kurile Trench, and the Costa Rica subduction
zone at 3000-4445 m. Both species are from cold seeps.
ACKNOWLEDGMENTS
We acknowledge MPI Observer Geoff Dolan for return¬
ing the clam valves, and Ashley Rowden, Steve Parker, and
David Bowden (NIWA) for comments. We thank Kevin
Mackay (NIWA) for preparing Figure 1 and Jean-Claude
Stahl (Te Papa) for preparing the images in Figure 2.
LITERATURE CITED
CCAMLR. 2009. VME Taxa Identification Guide. Commis¬
sion for the Conservation of Antarctic Marine Living
Resources, Hobart, Tasmania, Australia, 4 pp. Available
at http://vvww.cca1nlr.0rg/pu/e/e_pubs/VM E_guide.pdf.
Domack, E., S. Ishman, A. Leventer, S. Sylva, V. Willmott, and
B. Huber. 2005 Chemotrophic ecosystem beneath the
Larsen Ice Shelf. EOS Transactions of the American Geo¬
physical Union 86: 269-276.
Krylova, E. M . , H . Sahling, and R. Janssen. 20 1 0. Abyssogena : a new
genus of tlie family Vesicomyidae (Bivalvia) from deep-water
vents and seeps. Journal of Molluscan Studies 76: 107-132.
Rogers A.D., P.A. Tyler P.A., D.P Connelly, J.T. Copley, R.
James, R.D. Larter, K. Linse, R.A. Mills, A.N. Garabato,
R.D. Pancost, D.A. Pearce, N.V.C. Polunin, C.R. German,
T. Shank, PH. Boersch-Supan, B.J. Alker, A. Aquilina, S.A.
Bennett, A. Clarke, R. J .J . Dinley, A.G.C. Graham, D.R.H.
Green, J.A. Hawkes, L. Hepburn, A. Hilario, V.A.I. Huvenne,
Leigh Marsh, E. Ramirez-Llodra, W.D.K. Reid, C.N.
Roterman, C.J. Sweeting, S. Thatje, and K. Zwirglmaier.
2012. The discovery of new deep-sea hydrothermal vent
communities in the Soudiern Ocean and implications for
biogeography. PLoS Biology 10 (1): el001234. doi:10.1371/
journal. pbio. 1001234.
Bruce A. Marshall
Museum of New Zealand Te Papa Tongarewa
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Bruce [email protected]
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THE0NAUTILUS
Volume 129, Nu mber 4
December 10, 2015
ISSN 0028-1344
CONTENTS
Roland Houart Four new species of Muricidae (Gastropoda) from New Caledonia,
Papua New Guinea, and Indonesia . 143
M.G. Harasewych On the phylogenetic relationships of the genus Mexistrophia and
Amanda M. Windsor of the family Cerionidae (Gastropoda: Eupulmonata) . 156
Estuardo Lopez- Vera
Fred G. Thompson
A. Czaja A new species of the genus Mexipirgus Taylor, 1966 (Caenogastropoda:
Jose Luis Estrada-Rodriguez Truncatelloidea: Cochliopidae) from late Holocene spring deposits in
Ulises Romero-Mendez Viesca, Coahuila, Mexico . 163
Jessica Borquez Intracapsular development in the freshwater gastropod Chilina domheiana
Claudio Valdovinos (Bruguiere, 1789) (Gastropoda: Higroplhla: Chilinidae) . 169
Antonio Brante
Maria Jose Pio Trophon geversianus (Pallas, 1774): the first record of communal
Guido Pastorino egg masses in the muricid subfamily Trophoninae (Gastropoda) . 172
Gregory S. Herbert
Research Note
Arthur E. Bogan Determining the date of publication for Contradens Haas and
, Uniandra Haas (Bivalvia: Unionidae) . 175
Book Review . . . 179
Author Index . 181
$*MTHS0
DEC 1 7 2015
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THE NAUTILUS 129(4): 143-155, 2015
Page 143
Four new species of Muricidae (Gastropoda)
from New Caledonia, Papua New Guinea, and Indonesia
Roland Houart1
Institut royal des Sciences naturelles de Belgique
Rue Vautier, 29
B-1000 Bruxelles, BELGIUM
[email protected]
ABSTRACT
Four new species of Muricidae are described from New
Caledonia, Papua New Guinea and Indonesia and compared
with related species. One Timbellus species was collected in
New Caledonia. Two other species are described from Papua
New Guinea, respectively in Chicopinnatus and Dermomurex.
The fourth species, also belonging in Chicopinnatus, originates
from Indonesia.
Additional Keywords: Muricoidea, Chicoreus, Timbellus,
Dermomurex
INTRODUCTION
Many species have been traditionally classified in
Pterynotus Swainson, 1833 and 16 species are currently
considered as valid for this genus in WoRMS (Bouchet
pt al., 2015). However, according to recent molecular
research (Barco et al., 2010), Pterynotus as traditionally
defined (Vokes, 1964; 1971; Fair, 1976; Radwin and
D’Attilio, 1976; Houart, 1994), appears to be polyphyletic
consisting of at least two independent lineages. One group
includes all the species with a sculptural pattern similar
to the type species of Pterynotus, P. alatus (Roding, 1798)
(= Murex pinnatus Swainson, 1822). A second group,
based on the position in the molecular phylogeny of
P. fulgens Houart, 1988, includes species generally clas¬
sified in Pterynotus sensu stricto, but with a less scabrous
shell sculpture and with three major axial varices
“appearing early during the ontogeny” (Merle et al., 2011).
The genus Timl?ellus was reinstated by Merle et al. (2011)
to include the species that were formerly classified in
Pterynotus but which differ in having a trivaricate smooth
rather than scabrous shell.
Eight species of Timbellus species are known from the
New Caledonian area, six of them originally described in
Pterynotus (Houart, 1987; 1988; 1991; 2001). Two addi¬
1 Research Associate
tional species, formerly confused with Timbellus richeri
(Houart, 1986), were recently separated and described
as new Timbellus species (Houart, 2012). A ninth species
is described here.
Species of Chicopinnatus Houart, 1992, a subgenus of
Chicoreus, have been variously classified in Pterynotus
sensu stricto and other genera. Chicopinnatus originally
included three species: Chicoreus ( Chicopinnatus )
orchidijlorus (Shikama, 1973) (type species), C. (C.)
laqueatus (Sowerby, 1841) and C. (C. ) g uillei (Houart,
1985). Different shell characters separate them from
other genera and from Chicoreus sensu stricto. In this
article, three additional species formerly included in
Pterynotus or in Timbellus are added to Chicopinnatus
and two new species are described from Papua New Guinea
and Indonesia.
Dermomurex Monterosato, 1890 was organized by Vokes
(1976) into five subgenera: Dermomurex, Gracilimurex
Thiele, 1929, Takia Kuroda, 1953, Trialatella Berry,
1964, and Viator Vokes, 1974. All these but Gracilimurex
occur in the Indo-West Pacific. A new species from
Papua New Guinea is described here but its classification
in Dermomurex sensu stricto or D. (Trialatella) is doubt¬
ful, as it shares shell characters of both subgenera. The
new species is described without allocation to a subgenus.
MATERIALS AND METHODS
The material in this paper originates from four sources.
(1) The PAPUA NIUGINI expedition (Principal Investi¬
gator: Philippe Bouchet) conducted by MNHN and Pro-
Natura International (PNI) as part of the Our Planet
Reviewed program; its sponsors include the Total Foun¬
dation, Prince Albert II of Monaco Foundation, Stavros
Niarchos Foundation, Fondation EDF, and Ent repose
Contracting. The project operated under a Memorandum
of Understanding between MNHN and the University of
Papua New Guinea, with permits from the PNG Depart¬
ment of Environment and Conservation. The expedition
took place along the coast of New Guinea Island in the
Bismarck Sea, from the Vitiaz Strait to the border between
Papua New Guinea and Irian Jaya; (2) The EXBODI
Page 144
THE NAUTILUS, Vol. 129, No. 4
cruise (Principal Investigator: Sarah Samadi) as part of
the Tropical Deep-Sea Benthos program conducted by
MNI IN and Institut de Recherche pour le Developpement
(IRD); (3) The MUSORSTOM 4 cruise (Principal Inves¬
tigator: Bertrand Richer de Forges), in northern and
southern New Caledonia. Leg one of the latter expedition
took place between 12 September and 5 October 1985 off
northern and southern New Caledonia. Leg two took
place between 13 September and 01 October 2011 and
explored seamounts of the Loyalty Ridge; (4) One addi¬
tional new species originates from the personal collection
of Bunjamin Dharma, Indonesia.
The new species descriptions are based on all the
examined specimens. The characters used to describe
the shell morphology are the general aspect of the shell,
its shape and size, color, shape of the spire and number
of protoconch and teleoconch whorls, features of the
protoconch, shape of the teleoconch whorls and features
or form of the suture and of the subsutural band, of axial
and spiral sculptnre, the aperture and siphonal canal.
When known, the characters of the operculum are also used.
All width measurements are taken with the spines
included. Abbreviations are: DW: Waren dredge; IRD:
Institut de Recherche pour le Developpement (formerly
ORSTOM); ORSTOM: Office de la Recherche Seientifique
et Technique d’Outre Mer (now IRD); Collection abbre¬
viations are: MNHN: Museum national d’Histoire
naturelle, Paris, France; MZB: Museum Zoologieum
Bogoriense, Bogor, Java, Indonesia; RH: collection of the
author; ZSM: Zoologischen Staatssammlung, Miinchen,
Germany; Specimen status: dd: empty shell(s); juv: juve¬
nile; lv: live collected.
Terminology used to describe the spiral cords and
apertural denticles, listed according to type of struc¬
ture (after Merle, 2001 and 2005) (Terminology in
parentheses: variable feature) (Figures 1-5): P: pri¬
mary cord; s: secondary cord; t: tertiary' cord; ad: adapieal;
ab: abapieal; IP: infrasutural primary cord (primary cord
on subsutural ramp); adis: adapieal infrasutural secondary
cord (on subsutural ramp); abis: abapieal infrasutural
secondary cord (on subsutural ramp); PI: shoulder cord;
P2-P6: primary cords of the convex part of the teleoconch
whorl; sl-s6: secondary cords of the convex part of the
teleoconch whorl; example: si = secondary cord between
Pi and P2; s2 = secondary cord between P2 and P3, etc.;
ADP: adapertural primary cord on the siphonal canal;
MP: median primary cord on the siphonal canal;
ABP: abapertural primary cord on the siphonal
canal; abs: abapertural secondary cord on the siphonal
canal; Aperture, D1 to D6: abapieal denticles; ID:
Infrasutural denticle.
SYSTEMATICS
Family Muricidae Rafinesque, 1815
Subfamily Muricinae Rafinesque, 1815
Genus Timbellus de Gregorio, 1885
Type Species: Murex latifolius Bellardi, 1872, Middle
Miocene, Italy (subsequent designation by Vokes,
1964: 14)
Timbellus corbariae new species
(Figures 1-2, 6-11)
Description: Shell medium sized for the genus, up
to 39.6 mm in length at maturity ( ho! o type). LengtSi/
width ratio of the holotvpe 1.56. Last whorl slender, trian¬
gular with variceal wings, whorls narrow, almost smooth,
lightly built. Subsutural ramp broad, strongly sloping,
weakly concave.
Light creamy white with scattered brown blotches on
whorls and varices, more particularly visible in paratype.
Aperture white. Spire high, acute, with 7 narrowly convex,
weakly shouldered, almost smooth, teleoconch whorls.
Suture impressed. Protoconch broken in the two speci¬
mens. Axial sculpture of teleoconch whorls consisting of
3 narrow, strongly webbed varices. Each varix bearing
very thin, almost smooth, wing-like expansion from first to
last whorl. Other axial sculpture of 2 or 3 narrow, nodose,
intervariceal ribs, from suture to suture. Last whorl with
3 very low, narrow ribs, middle one weakly higher. Spiral
sculpture of very weak, narrow, smooth, barely visible pri¬
mary and secondary cords, most obvious on abapertural
side of variceal wings. Paratype with P1-P3 visible from
second whorl. Last whorl with adis, IP, abis, P1-P6, s6. No
visible cords on siphonal canal.
Aperture small, ovate. Columellar lip moderately broad,
smooth, adherent at adapieal extremity. Anal notch deep,
broad. Outer lip weakly erect, smooth with 7 strong,
weakly elongate denticles within: ID, D1-D6, decreas¬
ing abapieally in height and strength. Siphonal canal
long, broad, straight, strongly dorsally bent at tip, with
variceal wing over whole length, smooth. Operculum and
radula unknown.
Type Material: Holotype MNHN I M -2000-30342
and 1 paratype MNHN (as listed below).
Other Material Examined: New Caledonia:
EXBODI, stn DW3857, South Durand Bank, 22° 18' S,
168°42' E, 342 m, 14 September 2011 (holotype) (dd);
MUSORSTOM 4, stn DW205, 22°38' S, 167°07 E,
140-160 m, 27 September, 1985, 1 paratype MNHN
I M -2000-30343 (dd).
Type Locality: New Caledonia, South Durand Bank,
22° 18' S, 168°42' E, 342 m.
Distribution: South of New Caledonia, empty shells
in 160-342 m.
Remarks. Eight species of Timbellus occur off
New Caledonia and in the Coral Sea, living in deep
water, mostly between 200 to 400 m. Only two of these
can be reasonably compared with the new species.
Timbellus corbariae new species differs from T. rubidus
Houart, 2001 (Figures 12-13) in having a larger shell.
Timbellus corbariae has two additional teleoconch whorls,
R. Houart, 2015
Page 145
Figures 1-5. New species of Timbellus and Chico reus. Morphology of spiral cords and apertural denticles. 1-2. Timbellus
corbariae new species. 1. Paratype MNHN IM-2000-30343. 2. Holotype MNHN I M -2000-30342. 3. Chicoreus ( Chicopinnatus )
arbaguil new species. Holotype MNHN-IM-2013- 14388. 4. Chicoreus ( Chicopinnatus ) dharmai new species. 5. Dermomurex
fitialeatai new species. Holotype MNHN IM-2013-14300.
Page 146
THE NAUTILUS, Vol 129, No. 4
Figures 6—16. Species of Timbellus . 6—11. Timkellua corbariae new species. 6—8. New Caledonia. EXBODI, stn D W385 < ,
Son tli Durand Bank, 22° 18' S, 168°42' E, 342 m, holotype MNHN IM-20GG-30342, 39.6 mm. 9-1 1. New Caledonia, MUSORSTOM 4,
stn DW205, 22°38' S, 167°07' E, 140-160 m, paratype MNHN I M -2000-30343, 36.5 mm. 12-13. Timbellus rubidus (Houart, 2001).
New Caledonia, Norfolk Ridge, 23° 44' S, 168° 16' E, 394-401 m, holotype MNHN-IM-2000-0346, 13.2 mm (photos MNHN).
14—15. Timbellus fulgens (Houart, 1988). New Caledonia, 390-420 m, 22°52' S - 16i°12' E, holotype MNHN-IM-2Q00-0082,
25.5 mm (photos MNHN). 16. Timbellus flemingi (Beu, 1967). Norfolk Ridge, north of Norfolk Island, 750-774 m, RH, 28.7 mm.
R. H ouart, 2015
Page 147
which could explain its larger size, but the whorls are
also larger and broader and have 2 or 3 low, narrow and
elongate intervarieeal ridges instead of a single, obvious,
small node in T. nibidus, from first to last teleoconch
whorl. The aperture in T. corbariae is also comparatively
larger and broader and the spiral cords are obviously
narrower and shallower on the varieeal wings.
Timbellus corbariae differs from T. fulgens (Houart,
1988) (Figures 14—15) in having a larger shell and a
comparatively larger aperture with more obvious, high,
apertural denticles and in having 2 or 3 low intervarieeal
elongate ridges instead of an almost smooth shell in
T. fulgens.
A third similar species is T. flemingi (Beu, 1967)
(Figure 16) from New Zealand. Timbellus corbariae dif¬
fers in having a more elongate shell instead of strongly
bieonieal in T. flemingi and in having a higher spire.
Timbellus corbariae differs further in having a broader
aperture that is denticulate instead of almost or entirely
smooth, and in having two or three intervarieeal elongate
nodes instead of a single low node or none.
Etymology: Named for Laure Corbari (MNHN), chief
scientist during the second leg of the EXBODI campaign
(13/9 to 01/10/2011) when the holotype was collected.
Genus Chicoreus Monfort, 1810
Subenus Chicopinnatus Houart, 1992
Type Species: Pterynotus orchidiflorus Shikama, 1972,
Indo-West Pacific (original designation).
Remarks: Three species are currently included in
this subgenus: Chicoreus ( Chicopinnatus ) orchidiflorus
(Shikama, 1972) (Figures 49-50), C. (C.) guillei (Houart,
1985), and C. (C.) laqueatus (Sowerby, 1841).
Five additional species are added here: C. (C. )
brianbaileyi (Miihlhiiusser, 1984), C. (C.) loebbeckei
(Kobelt, 1879), C. (C.) miyokoae (Kosuge, 1979), C. (C. )
arbaguil new species and C. (C. ) dharmai new species.
Chicoreus ( Chicopinnatus ) miyokoae was tentatively
included in Timbellus (Merle et ah, 2011: 133) and
C. (C.) brianbaileyi in Pterynotus (Merle et ah, 2011:
121). Both species are similar in shell morphology and,
together with C. loebbeckei and the two new species
described here, share similar shell characters with
Chicopinnatus , such as the rounded or roundly ovate
aperture, the moderately long, spined siphonal canal,
the more or less webbed varieeal spines and the trivaricate
morphology starting from first teleoconch whorl.
Chicopinnatus species differ from Pterynotus alatus
(Roding, 1798) (= Murex pinnatus Swainson, 1822),
the type species of the genus Pterynotus and from
P. albobrunneus Bertsch and D’Attilio, 1980, P. elongatus
(Lightfoot, 1786), P. laurae Houart, 1997, P. patagiatus
(Hedley, 1912) and P. pellucidus (Reeve, 1845) in having
a broader shell with broader, round or roundly ovate
aperture, a comparatively narrower siphonal canal, a
lower, broader spire and a trivaricate shell starting from
first teleoconch whorl rather then from second or third
whorl in Pterynotus.
Chicopinnatus species differ further from Timbellus
species in having a broader shell with broader, more
rounded aperture but mainly in having a squamous
shell compared to the smooth, or almost smooth shell
in Timbellus.
A molecular phylogenetic study of the whole group is
necessary to test this new classification.
Chicoreus (Chicopinnatus) arbaguil new species
(Figures 3, 17-25, 34)
Description: Shell large for the subgenus, up to
85.9 mm in length at maturity (paratype MNHN
IM-2013-14446). Length/width ratio 1.21-1.51. Bieonieal,
broadly ovate, heavy, strongly spinose, nodose. Subsutural
ramp narrow, weakly sloping, convex. Light brown or tan,
occasionally with light purplish spinose fronds (holotype).
Primary and secondary cords lightly darker colored.
Apertural side of siphonal canal lighter colored. Aper¬
ture white. Spire high with 2 protoconch whorls and
teleoconch up to 8 broad, weakly shouldered, spinose
and nodose whorls. Suture impressed. Protoconch small,
weakly flattened, with rounded whorls. Last whorl
minutely punctate, with a narrow, strong keel abapically.
Width 1300 pm, height 1100 pm (paratype MNHN
IM-2000-30344). Terminal lip almost straight, partly
eroded. Axial sculpture of teleoconch whorls consisting
of high, narrow, rounded, nodose varices. Each varix
with long, acute, open, webbed, primary, secondary and
tertiary spines. Shoulder spine longest. P1-P3 and P4-s6
spines joined by thin webbing. Other axial sculpture of
high, strong, rounded intervarieeal ribs. First teleoconch
whorl already starting 3 small, rounded varices with 2
or 3 intervarieeal narrow ribs; second whorl with
intervarieeal ribs becoming broader. Three varices and
2 or 3 broad, high, intervarieeal ribs from fourth to last
whorl. Spiral sculpture of high, narrow, squamous and
nodose primary, secondary and tertiary cords and few,
nodose, narrow threads. First whorl with visible P1-P4;
second whorl starting IP; third and fourth with adis, IP,
PI, P2, P3, s3, P4; fifth with adis, IP, abis, PI, P2, P3, s3,
P4, starting additional threads; seventh whorl of subadult
paratype MNHN IM-2000-30344 with adis, IP, abis and
2 additional threads on subsutural ramp, followed by PI,
P2, P3, s3, P4, s4, P5, s5, P6, s6, tertiary cords and few
threads. Siphonal canal with strongly backward curved
ADP, straight MP and ABP. PI and P2 spines joined by
thin webbing from third to last whorl, including P3 spine
from fifth to last whorl. S3 not joined to other spines. P4
to s6 spines webbed. Adult shell with 8 teleoconch
whorls having similar shell sculpture morphology. Aper¬
ture large, broadly ovate. Columellar lip narrow, smooth,
with low parietal tooth at adapieal extremity. Rim adher¬
ent at adapieal extremity, weakly erect abapically. Anal
notch shallow, broad. Outer lip erect, crenulated, with
strong, low, narrow lirae within: IP split, Dl, D2-D7
occasionally split. Siphonal canal long, narrow, lightly
Page 148
THE NAUTILUS, Vol. 129, No. 4
Figures 17—25. Chicoretis ( Chicopinnatus ) arbaguil new species. 17—19. Papua New Guinea, PAPUA NIUGINI, stn PP06,
Rempi area, east of Tadwai Island, 145°48' E, 04°59' S, 180 m, holotype MNHN IM-2013-14388, 83.3 mm. 20-21. stn PP07, Rempi
area, east of Tadwai Island, 145°48' E, 04°59' S, 150 m, paratype MNHN IM-2013-14446, 85.9 mm. 22-23. Madang Lagoon,
no other data, paratype MNHN I M -2000-30344, 44.1 mm. 24. Protoconch (paratype MNHN I M -2000-30344), scale bar 500 pm.
25. Operculum (holotype IM-2013-14388), scale bar 500 pm.
R. Houart, 2015
Page 149
abaxially bent, weakly dorsally recurved at tip, narrowly
open, with 3 or 4 acute spines: ADP, MP, ABP, (abs),
decreasing in length abapically. Operculum dark brown,
broadly ovate, with subapical nucleus and 13 concentric
ridges. Attached surface with many growth lines and
broad, callused rim. Radula unknown.
Type Material: Holotype MNHN IM-2013-14388 (Iv)
and 3 paratypes MNHN, 1 coll. RH (as listed below).
Material Examined: Papua New Guinea: PAPUA
NIUGINI, stn PP06, Rempi area, east of Tadwai Island,
145°48' E, 04° 59' S, 180 m, 19 November 2012, 1 lv
(holotype IM-2013-14388); stn PP07, Rempi area,
east of Tadwai Island, 145°48' E, 04°59' S, 150 m,
20 November 2012, 1 lv (paratype MNHN IM-2013-
14446); Madang Lagoon, November-Deeember 2012,
no other data, 2 paratypes MNHN I M -2000-30344 (1 lv,
1 dd, juv), 1 paratype RH (dd).
T)pe Locality: Papua New Guinea, Rempi area, east of
Tadwai Island, living at 180 m.
Distribution: Papua New Guinea: Rempi area, east
of Tadwai Island, living at 150-180 m and Madang
Lagoon, no other data.
Remarks: Chicoreus ( Chicopinnatus ) arbaguil differs
from C. brianbaileyi (Ligures 36—13) in having a larger,
more elongate shell with a same number of teleoconch
whorls, a comparatively larger and broader aperture, and
less expanded variceal wings with longer spines. It also
has a comparatively higher spire and a longer siphonal
canal with more broadly spaced spiral cords. The distance
between P4 and ADP is more reduced with less webbed
wings and narrower spiral cords. The protoconch
(Ligure 24) is smaller with broader first whorl and a
strongly keeled last whorl compared to the broader,
smooth protoconch of C. brianbaileyi (Ligure 43).
Chicoreus ( Chicopinnatus ) arbaguil differs from
C. miyokoae (Ligures 44-46) in having a more elongate
and larger shell with a same number of teleoconch
whorls, more broadly spaced spiral cords, and broader
primary cords. The spire is higher, the siphonal canal
longer and the ADP, MP and ABP cords are not webbed
like in C. miyokoae. The aperture is comparatively larger.
The variceal spines are longer, straight and not short,
strongly webbed and adaperturally recurved as in
C. miyokoae. The protoconch is almost twice as wide
with a strongly keeled last whorl while it is small and
smooth in C. miyokoae (Ligure 46).
Chicoreus ( Chicopinnatus ) arbaguil differs from
C. loebbeckei (Ligures 47—48) by the same characters
separating it from C. miyokoae. Chicoreus ( Chicopinnatus )
arbaguil differs further in having lecithotrophic larval
development, as attested by the paucispiral, rounded
protoconch, rather than planktotrophic development as
in C. loebbeckei, which has a conical protoconch of more
than three whorls with a narrow keel abapically and a
terminal lip of sinusigera type (Ligure 48).
Etymology: This new Chicoreus is named after Jo
Arbasto and Noel Saguil who, during the Madang expedi¬
tion, operated the tangle nets that led to its discovery. Jo
Arbasto is a professional tangle net fisherman on the island
of Panglao, in the Philippines, and Noel Saguil has been a
project officer working lor biodiversity projects, also in the
Philippines. Both contributed immensely to the success of
MNHN expeditions, notably the Panglao and Aurora
expeditions in the Philippines, Santo 2006 in Vanuatu,
and now' the Papua New Guinea Madang 2012 expedition.
The word arbaguil is used as a noun in apposition.
Chicoreus (Chicopinnatus) dharmai new species
(Ligures 4, 26-33, 35)
Pterynotus brianbaileyi. — Dharma, 2005: 164, pi. 57,
fig. 14 (not Pterynotus brinbaileyi Miihlhausser, 1984).
Description: Shell large for the subgenus, up to
74.6 mm in length at maturity (holotype). Length/width
ratio 1.2.5-1.26. Bieonical, broad, heavy, spinose and nodose.
Subsutural ramp broad, weakly sloping and convex. White
or light tan wtith pinkish or brown protoconch and 2 or
3 first teleoconch whorls. Primary, or primary and second¬
ary cords topped with narrow brown line. Aperture white.
Spire high with 2 protoconch w'horls and teleoconch up to
8 broad, weakly shouldered, spinose and nodose whorls.
Suture impressed. Protoconch small, whorls rounded,
smooth. Maximum height and width 1300 pm. Terminal
lip almost straight, partly eroded. Axial sculpture of
teleoconch whorls consisting of high, narrow, rounded,
nodose varices. Each varix with long, broad, open,
webbed, primary and short secondary spines. Shoulder
spine longest. P1-P3 and P4— s6 spines joined by thin
webbing. Lirst whorl already starting 3 small, rounded
varices with 2 or 3 nodose intervariceal ribs; third whorl
with broader ribs. Lourth to last whorl with 2 broad, high,
intervariceal ribs. Spiral sculpture of low, narrow, nodose,
primary, secondary and tertiary cords and numerous
nodose threads. Lirst whorl with visible IP, P1-P4; second
with IP, PI, P2, P3, s3, P4; third whorl starting spiral
threads; fourth to penultimate whorl with visible adis, IP,
abis, PI, P2, P3, s3, P4. Last whorl with adis, IP, abis, PI,
P2, s2, P3, s3, P4, s4, P5, s5, P6, s6, two additional tertiary
cords and numerous spiral threads over whole shell.
Siphonal canal with almost straight or straight ADP, MP
and ABP. PI and P3 spines webbed from second to last
whorl; P4 to s6 spines webbed. S3 not joined by webbing
to other spines. Aperture huge, broadly ovate. Columellar
lip narrow, smooth, with low parietal tooth at adapical
extremity. Rim adherent at adapical extremity, weakly
erect abapically. Anal notch shallow, broad. Outer lip erect,
crenulated, with strong, low, narrow lirae within: IP split,
Dl, D2-D7 occasionally split. Siphonal canal long, nar¬
row, lightly bent abaxially, weakly dorsally recurved at tip,
narrowly open, with 3 acute, straight or almost straight
spines: ADP, MP, ABP, decreasing in length abapically.
Operculum dark brown, broadly ovate, with subapical
nucleus and 17 concentric ridges (paratype B, Dharma).
Page 150
THE NAUTILUS, Vol. 129, No. 4
Figures 26-33. Chicoreus ( Chicopinnatus ) dharmai new species. 26-28. Indonesia, East Java, Bamyuwangi-Jember, 146 m,
in lobster nets, holotype MZB Gst. 18174, 74.5 mm. 29-30. Lombok Island, in lobster net, +/- 91 m, paratype BH, 67.3 mm.
31. East Java, Muncar, 165 m, in lobster nets, paratype B. Dharrna, 65.8 mm. 32. Operculum (paratype B. Dharma), scale bar 500 pm.
33. Protoconch (paratype BH), scale bar 500 pm.
R. Houart, 2015
Page 151
Figures 34-43. Species of Chicoreus. 34-35. Spiral sculpture (scale bar 5 nun). 34. Chicoreus ( Chicopinnatus ) arbaguil new
species, holotype MNHN-IM-2013-14388. 35. Chicoreus ( Chicopinnatus ) dharmai new species, paratype RH. 36-43. Chicoreus
( Chicopinnatus ) brianbaileyi (Miihlhausser, 1984). 36-37. Russell Island, Solomon Islands, holotype ZSM 1746, 74 mm (photo
E. Sehwabe). 38-40. Solomon Islands, SALOMONROA3, stn DW2855, 9° 45' S, 160° 50' E, 183 m, MNHN I M -2009-5858, 67.2 mm.
41^2. Vanuatu, MUSORSTOM 8, stn CP1071, 15°37' S, 167° 16' E, 180-191 m, MNHN-IM-2012-18002, 62.6 mm. 43. Protoconch,
scale bar 500 pm.
Page 152
THE NAUTILUS, Vol. 129, No. 4
Figures 44-60. Species of Chicoreus and Dermomurex. 44—46. Chicoreus ( Chicopinnatus ) miyokoae (Kosuge, 1979).
44-45. Philippines, Samal, Ligid Island, 58.5 mm, RH. 46. Protoconch. Philippines, Davao, RH, scale bar 500 pm. 47-48. Chicoreus
(Chicopinnatus) loebbeckei (Kobelt, 1879), Philippines, Davao, Samal Island, 53.6 mm. RH. 48. Protoconch, scale bar 500 pm.
49-50. Chicoreus ( Chicopinnatus ) orchidiflorus (Shikama, 1972), Philippines, Bohol, Caubian Island, 130 m, RH, 41.6 mm.
51-56. Dermomurex fitialeatai new species. 51-53. Papua New Guinea, north of Tadwai Island, outer slope, 145°47,7'E,
04°59,1'S, 22 m, holotype MM IN IM-2013-14300, 18.9 mm. 54. Protoconch (holotype), scale bar 500 pm. 55. Operculum
(holotype), scale bar 1 mm. 56. Detail of the intritacalx (holotype), scale bar 1 mm. 57-59. Dermomurex triclotae Houart, 2001.
57-58. New Caledonia, channel of Koumac pass, 20°40.7' S, 164° 14.7' E, holotype MNHN-IM -2000-0340, 13.10 mm (photo
MNI IN). 59. Detail of the intritacalx (paratype RH), scale bar 1 mm. 60. Dermomurex trondleorum Houart, 1990, French Polynesia,
Tuamotu Archipelago, Anaa Atoll, holotype MNHN-IM-2000-0223, 17 mm (photo MNHN).
R. Houart, 2015
Page 153
Attached surface with many growth lines and broad,
callused rim. Radula unknown.
Type Material: Holotype MZB Gst. 18174, 1 paratype
B. Dharma; 1 paratype collection RH (as listed below).
Material Examined: Indonesia: East Java, Bamyuwangi-
Jember, 146 m, in lobster net, January 2014, 1 dd (holo¬
type MZB); East Java, M uncar, 165 m, in lobster net, 1 Iv
(paratype coll. B. Dharma); Lombok Island, in lobster
net, about 90 m, 2005, 1 dd (paratype coll. RH).
Type Locality': Indonesia, East Java, Bamyuwangi-
Jember, 146 in, in lobster nets.
Distribution: Indonesia, east of Java and Lombok
Island, living at 165m.
Remarks: Chicoreus ( Chicapinnatus ) dharmai is simi¬
lar to C. arbaguil but differs in protoconch and other shell
morphology. The protoconch is comparatively larger,
higher and smooth rather than flatter and having a
strongly keeled last whorl in C. arbaguil. The sculpture of
the teleoconch is also similar but the spiral cords are
narrower in C. dharmai, and the spiral threads are more
numerous (Figures 34—35), IP starts from first teleoconch
whorl rather than the second and the space between
cords P4 and s6 is distinctly narrower (Figures 3-4). The
shell is also stockier than C. arbaguil, smaller for a same
number of teleoconch whorls, and has a lower spire and a
notably shorter siphonal canal with straight rather than
strongly backward curved ADP spine.
Chicoreus ( Chicopinnatus ) dharmai differs from the
other more or less similar species, C. brianbaileyi,
C. miyokoae and C. loebbeckei in having more strongly
spinose axial varices, lower and narrower intervariceal
nodes, less numerous, narrower secondary spiral cords
and a comparatively narrower and longer siphonal canal.
I earlier misidentified a specimen of C. dharmai as
C. brianbaileyi which was thus identified as such by
Dharma (2005).
Etymology: This species is named for Bunjamin
Dharma, who kindly donated the holotype and in acknowl-
edgment for more than 20 years of useful collaboration.
Genus Dermomurex Monterosato, 1890
Type Species: Murex scalarinus Bivona-Bernardi, 1832
{= Murex scalaroides Blainville, 1829), Mediterranean
Sea and Eastern Atlantic (Senegal) (original designation).
Remarks: There are currently fifteen Recent Indo-West
Pacific species assigned to Demwmurex Monterosato,
1890. Six belong in Dermomurex sensu stricto: D. agnesae
Vokes, 1995, D. angustus (Verco, 1895), D. chariest Houart
and Heros, 2013, D. goldsteini (Tenison Woods, 1876),
D. neglecta (Habe and Kosuge, 1971), and D. rayivalkeri
Houart, 1986; four in subgenus Takia Kuroda, 1953:
D. africanus Vokes, 1978, D. bobyini Kosuge, 1984,
D. infrons Vokes, 1974, and D. wareni Houart, 1990; two
in subgenus Trialatella Berry, 1964: D. triclotae Houart,
2001 and D. trondleorum Houart, 1990; and three in
subgenus Viator Vokes, 1974: D. antonius Vokes, 1974,
D. hoivletti Vokes, 1995, and D. pasi Vokes, 1993.
A simple key to separate the subgenera of Dermomurex
was given by Vokes (1985):
I. SPIRE SHORT
A Six varices
1. Moderate canal
2. Long, straight canal
B Three varices
II. SPIRE ELONGATE
A Two varices
B Three to six varices
Takia
Viator
Trialatella
Gracilimurex
Dermmuirex sensu stricto
Although the distinction between Dermomurex sensu
stricto, Dermomurex (Takia), and Dermomurex ( Viator )
is clear, it is not readily apparent how to separate some
species of Dermomurex sensu stricto from Dermomurex
(Trialatella) . For example, the species newly described
here could be allocated into Trialatella because it has
three axial varices per whorl from third to last teleoconch
whorl, but the spire is elongate rather than short as
in Trialatella and fits better in Demwmurex sensu
stricto. Other representatives of uncertain relationship
were recorded by Merle et al. (2011: 212-213), who also
doubted the need for Trialatella. Therefore the decision
was taken here to describe this new species without any
subgeneric distinction.
Dermomurex fitialeatai new species
(Figures 5, 51-56)
Description: Shell medium sized for the genus,
18.9 mm in length. LentghAvidth ratio 2.35. Slender, lan¬
ceolate, nodose, lightly built. Subsutural ramp narrow,
weakly sloping and convex. Shell covered by thick, white,
minutely reticulate intritaealx. Aperture bluish-white.
Spire high with 1.5 protoconch whorls and 6.5 weakly
convex, narrow, shouldered, nodose whorls. Suture
impressed, partially obscured by small, narrow buttresses
connecting preceding whorl. Protoconch small. Whorls
rounded, smooth, height 900 pm, width 800 pm. Terminal
lip lightly erect, narrow, opisthocline. Axial sculpture of
teleoconch whorls consisting of narrow lamellae and high,
strong, narrow, rounded varices. First and second whorl
with 6 axial lamellae; third whorl starting varices. Three
rounded varices and one narrow, low, intervariceal node
from fourth to last whorl. Spiral sculpture of low, rounded,
broad, nodose, primary cords, visible from fourth to last
whorl. Last whorl with P1-P6. PI, P2 and P3 broad,
moderately high, more obvious when connecting axial
varices, forming deep pits between PI and P2 and P2
and P3. P4, P5 and P6 almost obsolete, probably more
obvious when intritaealx removed. Siphonal canal with
very low ADP, MP and ABP. Aperture small, ovate. Colu-
mellar lip narrow, smooth, adherent. Anal notch shallow,
broad. Outer lip not fully adult, smooth within. Siphonal
Page 154
THE NAUTILUS, Vol. 129, No. 4
canal short, narrow, dorsally bent at tip, open. Operculum
dark brown, roundly ovate, inverted tear-shaped with
apical nucleus and numerous concentric ridges. Attached
surface with about 4 growth lines and very broad, large,
callused rim. Radula unknown.
Type Material: Holotype MNHN IM-2013-14300 (lv).
Material Examined: Papua New Guinea: PAPUA
NIUGINI, stn PB26, north of Tadwai Island, outer
slope, 145°47,7'E, 04°59,1'S, 22 m, 22 November 2012
(holotype MNHN).
Type Locality: Papua New Guinea, north of Tadwai
Island, outer slope, 145°47,7' E, 04°59,1' S, 22 m.
Distribution: Papua New Guinea, north of Tadwai
Island, living at 22 m.
Remarks. Dermomurex fitialeatai new species can only
be compared with D. triclotae Houart, 2001 from New
Caledonia (see discussion under Dermomurex). It differs
from all the other species in haring a trivaricate shell
whereas the other species from the Indo-West Pacific
bear lour to six varices.
Dermomurex trondleorum from French Polynesia also
has a trivaricate shell but differs in many ways and does
not need to be compared further here. It is only illus¬
trated for reference (Figure 60).
Dermomurex fitialeatai new species differs from
D. triclotae (Figures 57-59) in having a more elongate
spire, 7 varices on first teleoconch whorl and 6 on second
and third, while decreasing from 6 varices on first whorl
to 3 on all the subsequent whorls in D. triclotae. The
intritaealx also differs in being minutely reticulate in
D. fitialeatai (Figure 56) and faintlv striate in D. triclotae
(Figure 59).
Etymology': The name of this species is dedicated to
Christian Kesiano Fitialeata, who tragically passed away
during the first leg of BioPapua, another expedition to
Papua New Guinea, which took place in 2010.
ACKNOWLEDGMENTS
I am gratefnl to Philippe Bouehet for giving me
opportunity to study the material from the expeditions
organized jointly by MNHN and IRD over many years
and for his comments on the manuscript, to Enrico
Schwabe (Bavarian State Collection of Zoology, Miinchen,
Germany), for the digital images of the holotype of
Pterynotus brianbaileyi , to Virginie Heros and Philippe
Maestrati (Museum national d’Histoire naturelle, Paris)
for the loan of material and for other information, to
Bunjamin Dharma for the donation of the holotype of
Chicoreus ( Chicopinnatus ) dharmai new species and to
Manuel Caballer (MNHN) who provided the images
of the MNHN types, E-Recolnat Project: ANR-11-
INBS-0004. Thanks also to John Wolff, Lancaster,
Pennsylvania, USA, for checking the English text. I also
thank Bruce Marshall (Museum of New Zealand, Te
Papa Tongarewa, Wellington, New Zealand) and Marco
Oliverio (University of Rome, Italy) for their careful
review of the manuscript.
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Neogastropoda): importance of ontogenetic and topological
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Recent Muricidae of the World - Part Muricinae.
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(Mollusca: Gastropoda. Muricinae, Ocenebrinae. Bulletin
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THE NAUTILUS 1 29(4): 1 56-162, 2015
Page 156
On the phylogenetic relationships of the genus M exist rophia
and of the family Cerionidae (Gastropoda: Eupulmonata)
M.G. Harasewych
Amanda M. Windsor
Dept, of Invertebrate Zoology, MRC-163
National Museum of Natural History
Smithsonian Institution
PO Box 37012
Washington, DC 20013-7012 USA
H a [email protected] u
Estuardo Lopez- Vera
Institute de Ciencias del Mary Limnologia
Universidad Nacional Autonoma de Mexico
Circuito Exterior S/N
Ciudad Universitaria
Delegacion Coyoacan
CP: 04510 Mexico D.F. MEXICO
Fred G. Thompson
Florida Museum of Natural History
University of Florida
Gainesville, FL 32611 USA
ABSTRACT
Phylogenetic analyses of partial DNA sequences ot the mito¬
chondrial COI and 16S rDNA genes derived from Mexistrophia
reticulata Thompson, 2011, the type species of the genus
Mexistrophia , indicate that this genus is sister taxon to all
remaining living Cerionidae, and that the family Cerionidae is
most closely related to Urocoptidae. Relationships among repre¬
sentative cerionid taxa are consistent with the zoogeographic
hypothesis that Mexistrophia has been isolated from the remain¬
ing living Cerionidae since the Cretaceous, and suggest that the
near-shore, halophilic habitat that has commonly been associated
with this family is likely a Cenozoic adaptation that coincided
with the transition from continental to island habitats. The genus
Protocerion is described to include the Late Cretaceous species
Cerion acherontis Roth and Hartman, 1998, as its retention in
Cerion would render this genus paraphyletic.
INTRODUCTION
The family Cerionidae comprises a well-studied group
of terrestrial snails inhabiting islands of the tropical west¬
ern Atlantic, ranging from the barrier islands of southeast¬
ern Florida, throughout the Bahamas, Greater Antilles,
Cayman Islands, western Virgin Islands, and the Dutch
Antilles, but absent from Jamaica, the Lesser Antilles,
and coastal Central and South America. All are halophilic,
confined to terrestrial vegetation growing in close prox¬
imity of the shore, but occasionally occur further inland
in areas that can be reached by salt spray (Clench, 1957;
Woodruff, 1978). Rare fossil taxa extend the range of
Cerionidae to the Upper Cretaceous of Montana (Roth
and Hartman, 1998) and the Paleoeene of the Itaboraf
Basin of Brazil (Salvador et ah, 2011). More recently, the
genus Mexistrophia was proposed within the Cerionidae
to include three new species inhabiting cool coniferous
forests in the highlands (2000-2600 m) of central Mexico
(Thompson, 2011). Thompson (2011) compared the shell
morphology, anatomy, and radula of Mexistrophia
reticulata, the type species of Mexistrophia, with those of
several species of Cerion, including Cerion uva (Linnaeus,
1758), the type species of the type genus of Cerionidae.
He concluded that anatomical features of Mexistrophia
reticulata are typical of Cerionidae and that radular mor¬
phology differs only slightly. However, Mexistrophia may
be distinguished from species of Cerion in lacking lamellae
and denticles along the columella at all stages of growth.
Harasewych (2012) reviewed the diversity of living and
fossil Cerionidae from geographic and temporal perspec¬
tives and combined these data with paleogeographie recon¬
structions of the Caribbean region (Iturralde-Vinent, 2006)
and a COI based phylogenv of a selection of cerionid taxa
(Harasewych et ah, 2011: fig. 17) to formulate a hypothesis
for the zoogeographic history of the family Cerionidae
from the earliest fossil record in the Late Cretaceous of
Montana to the more widespread modern fauna. Accord¬
ing to this hypothesis (Figure 1), Mexistrophia was an early
offshoot that was isolated from the South American
ancestors of all remaining cerionids during the Late Cre¬
taceous by the formation of a seaway separating the faunas
of North and South America.
The family Cerionidae lias been assigned to a variety of
superfamilies, among them Orthalicoidea (Thiele, 1931;
Bouchet et ah, 2005) Clausilioidea (Baker, 1961; Solem,
1978; Tillier, 1989) Cerioidea (Baker, 1955; Shileyko,
1979, 1999) and most recently Urocoptoidea (Uit de Weerd,
2008). In this paper we investigate the phylogenetic rela¬
tionships of the genus Mexistrophia to other members of
the Cerionidae as well as the placement of the family
Cerionidae within Eupulmonata based on partial sequences
for the COI and 16S genes.
MATERIALS AND METHODS
Specimens of Mexistrophia reticulata Thompson, 2011
were collected west of Pinal de Amoles, Queretaro State,
Mexico (21°07' 26.52" N, 99°40'59,58" W), not far from
the type locality for this species. The shells were cracked.
M.G. Harasewych et al., 2015
Page 157
Figure 1. Hypothesized zoogeographic history of the family Cerionidae (after Harasewych 2012:fig. 12). Grey portions based on
fossil taxa. ABC, Aruba, Bonaire and Curasao; B, Brazil; C, Cuba; Cl, Cayman Islands; F, Florida; GBB, Great Bahama Bank;
H, Hispaniola; LBB, Little Bahama Bank; M, Mexico; PR, Puerto Rico, VI, western Virgin Islands.
and die animals preserved in RNAlater and stored at — 70°C
(voucher material USNM 1283835). As the Cerionidae
had most recently been included in Uroeoptoidea on the
basis of molecular data (Uit de Weerd, 2008), tissue from
living specimens of Microceramus pontificus (Gould, 1848)
from South Miami, Florida (voucher specimens USNM
1283834) were also sequenced in order to include a mem¬
ber of the family U rocoptidae in this analysis.
Genomic DNA was extracted from buccal muscle dis¬
sected from preserved or living specimens using the
DNAeasy Tissue Kit (Qiagen) according to the manufac¬
turer’s animal tissue protocol.
Portions of two mitochondrial genes were amplified: a
655 bp region of the cytochrome c oxidase I gene using
the primers JgLCO1490 (Geller et al., 2013) and Cl-N-
2191R (ilka NaneyCOIR) (Simon et id., 1994) and a 510 bp
region of the 16S ribosomal gene using the primers 16S-ar
and 16S-br (Palumbi, 1996). For each gene, the Promega
GoTaq hot start master mix (Promega M7132) was uti¬
lized at concentrations according to manufacturer’s
instructions, but modified to reduce reaction volume to
20 pL. Cycling parameters for each gene region were opti¬
mized as follows: COI - initial denaturation for 7 min at
95° C + 45 cycles (30 sec at 95°C + 45 sec at 42°C +1 min
at 72°C) + 3 min at 72°C; 16S - initial denaturation for
7 min at 95°C + 35 cycles (30 sec at 95° C + 45 sec at
48°C +1 min at 72°C) + 5 min at 72°C. PCR products
were visualized by agarose gel electrophoresis (1.5% aga¬
rose) and purified with ExoSAP-IT (Affymetrix) accord¬
ing to manufacturers protocols prior to sequencing.
Page 158
THE NAUTILUS, Vol. 129, No. 4
Sequencing reactions for 16S were performed using
1 pL of purified PCR product in a 10 pL reaction con¬
taining 0.5 pL primer, 1.75 pL Big Dye buffer and 0.5 pL
Big Dye (Life Technologies); for COI the volume of Big
Dye was increased to 0.75pL. The sequencing reaction
was carried out under standard cycling conditions (25 cycles
of 5 sec at 95 °C + 10 sec at 50 °C + 4 min at 60 °C).
Reactions were purified using Millipore Sephadex plates
(MAHVN-4550) according to the manufacturers instruc¬
tions and sequenced on an AB1 3730XL automated DNA
sequencer. Sequencher v. 4.7 (GeneCodes, Ann Arbor,
MI, USA) was used to visualize, trim, edit, and assemble
contigs from forward and reverse sequences. All PCR,
sequencing, and analytics were carried out at the Labora¬
tories of Analytical Biology at the National Museum of
Natural History. The sequences have been deposited in
GenBank (NCBI). Accession numbers are listed in Table 1.
Partial sequences for the mitochondrial COI and 16S
genes of Mexistrophia reticulata and Microceramus
pontificus were aligned against a range of euthyneuran
taxa for which both COI and 16S sequences were available
(Table 1), most derived from complete mitochondrial
genomes. Representative species of Cerionidae were
selected to span the previously documented phyloge¬
netic diversity within the family (Harasewyeh et ah,
2011: fig. 17).
Alignments of COI and 16S were obtained using the
L-INS-i alignment strategy in MAFFT (Katoh et ah, 2002)]
for 16S anti MUSCLE (Multiple Sequence Comparison
by Log- Expectation) (Edgar, 2004) for COI. The aligned
sequences were concatenated using Geneious version 7. 1 .2
(Kearse et ah, 2012) In the concatenated data set, posi¬
tions 1-554 are 16S, and positions 555 to 1212 are COI.
A best-fit model of nucleotide sequence evolution
(compatible with MrBayes) and partitioning arrangement
for each locus was determined using MrAIC (Nylander,
2004). The GTR+I + G model was chosen for both loci.
Phylogenetic analyses were performed on a con¬
catenated dataset (16S + COI) using Bayesian Infer¬
ence (BI) performed with MrBayes 3.1.2 (Ronquist and
Huelsenbeek, 2003) and Maximum Likelihood (ML)
with RAxML (Stamatakis, 2006). All analyses were run on
the Smithsonian Institution high performance computing
cluster (SI/HPC). BI analysis was carried out for 10 million
Table 1. List of taxa and their GenBank reference numbers for the gene sequences used in phylogenetic analyses. Asterisk (*)
denotes that both COI and 16S sequence data were derived from a complete mitochondrial genome sequence.
M.G. Harasewych et al., 2015
Page 159
generations with two independent runs, each with four
chains, and with trees sampled every 1000th generation.
Model parameters (tratio, statefreq, shape, pinvar) were
unlinked among partitions, and the rate prior (prset
ratepr) was set to “variable”. Convergence was determined
when the average standard deviation of split frequencies
was <0.01 and the potential scale reduction factor (PSRF)
was 1.00. To calculate posterior probabilities, a “burn-in”
of 25% of the total trees sampled per run adequately
removed trees prior to convergence. ML options for
RAxML included the GTRCAT model of nucleotide
evolution (-m), rapid bootstrap analysis, and search for
best-scoring ML tree (-fa), and 1000 bootstrap replicates.
RESULTS
The region of the 16S gene sequenced for Mexistrophia
reticulata and Microceramus pontificus corresponded
to positions 564-1069 of the 16S gene in Cerion
incanum. The length was 486 bp in Mexistrophia and
476 bp in Microceramus. The alignment containing
the taxa in Table 1 spanned 554 positions, of which
174 (31.4%) were constant and 326 (58.8%) were parsi¬
mony informative. The 655 bp portion of the COl
gene sequenced for Mexistrophia reticulata and
Microceramus pontificus corresponded to positions 39-
693 of the COI gene in Cerion incanum. The COI
alignment of the taxa in Table 1 spanned 658 bp, of
which 310 (47.1%) were constant and 302 (45.9%) were
parsimony informative.
Phylogenetic analyses of the concatenated 16S + COI
data using maximum likelihood and Bayesian inference
resulted in a single, fully resolved and well supported
tree (Figure 2). Mexistrophia emerged as the sister taxon
to all remaining living Cerionidae, and the family
Cerionidae as sister taxon to the single representative
Aplysia californica
Siphonaria pectinata
Salinator rhamphidia
■Ovatella vulcani
■Trimusculus reticulatus
• Plativendix mortoni
• Onchidella celtica
■ Peronia peroni
■ Biomphalaria glabrata
■ Physella acuta
Pupilla muscorum
Albinaria caerulea
Achatina fulica
Mastigeulota kiansinensis
Camaena cicatricosa
Cylindrus obtusus
Cepaea nemoralis
Cerion uva
Figure 2. Molecular phylogenetic tree from analyses of concatenated 16S and COI sequence data, represented as a maximum
likelihood phylogram with maximum likelihood bootstrap values/ Bayesian posterior probabilities. (* > 95% support, - < 50% support).
The family Cerionidae is in gray. Letters to the right of the bar correspond to the geographic localities / island groups identified in figure 1 .
Page 160
THE NAUTILUS, Vol. 129, No. 4
of the family Urocoptidae. Cerionidae and Urocoptidae
(both in Uroeoptoidea) are most closely related to the
Helicoidea, a clade represented by multiple families in
our study.
DISCUSSION
Molecular data strongly supports the inclusion of the
genus Mexistrophia in the Cerionidae, an assignment
originally based on anatomical data and shell morphol¬
ogy. The topology of the phylogram of the Cerionidae is
concordant with the branching patterns in the zoogeo¬
graphic hypothesis for the distribution of Cerionidae
(Figure 1), although several geographic regions are not
yet represented by molecular data.
Results of this analysis also recovered a sister group
relationship between Cerionidae and the urocoptid
Microceramus pontificus , supporting the inclusion of
Cerionidae within Uroeoptoidea, as advocated bv Uit de
Weerd (2008), and contradict its placement within
Clausilioidea (Baker, 1961; Solem, 1978; Tillier, 1989).
To date, Cerionidae have been included in verv few of
the broader molecular studies of pulmonate phytogeny.
Based on ribosomal RNA sequences, Wade and co-authors
(2001 : fig. 1) show Cerion as the sister taxon to Helicoidea +
Spiraxidae + Haplotremidae, but do not include
Urocoptidae among their sampled taxa. Topology of the
more basal portions of our tree are generally consistent
with results of other molecular studies on phylogenetic
relationships among Pulmonata (e.g., Wade et ah, 2006;
Dayrat et ah 2011; White et al. 2011) when adjusted for
taxon sampling and rooting.
The phylogenetic relationships within Cerionidae sug¬
gest that the near-shore, halophilie habitat that has com¬
monly been associated with this family is likely a Cenozoie
adaptation that coincided with the transition from conti¬
nental to island habitats.
As reported by Thompson (2011: 190), Mexistrophia
species inhabit cool mesic or submesic temperate forests,
at elevations greater than 2000 m and distances in excess
of 200 km from the nearest coastline. Mexistrophia is an
early offshoot of a lineage that dates back to “ Cerion ”
acherontis from the Hell Creek Formation [Upper Creta¬
ceous (Maastrichtian)] of northeastern Montana. This
species was part of a faunule that consisted almost entirely
of fresh water forms (Roth and Hartman, 1998: Table 1).
The habitat for this faunule has been interpreted as a
subtropical, flat, forested floodplain. Similarly, the genus
Brasilennea (Maury, 1935), recently transferred to the
Cerionidae (Salvador et al, 2011), was endemic to the
Middle to Late Paleocene Itaboraf Basin of Brazil. This
genus was part of a fauna that included numerous terres¬
trial snails and mammals. Maury (1935: 4) commented
that a crocodilian jaw was associated with the fossils she
described. Salvador and Simone (2012: 49) noted that
little is known about the paleoenvironment of the Itaboraf
Basin, other than its high calcium carbonate availability,
and cited reports that this basin had a wet and warm
climate with copious vegetation.
Cerion uva represents the most basal lineage within
the Cerionidae that is limited to island habitats. This
species is endemic to the islands of Aruba, Bonaire and
Curasao, and is widely distributed throughout these
islands (Harasewych, 2014: fig. 4), occurring on lime¬
stone plateaus along the coast, as well as further inland,
even at elevations of 200 m or more. There are few
places on these hot, arid islands that are more than 5 km
from the ocean. Windborne salt spray and salt particles
reach most or all parts of these islands. All remaining
species of Cerionidae are limited to islands ranging from
southern Florida throughout the Bahamas, Cuba, Cayman
Islands, Hispaniola, Puerto Rico and the western Virgin
Islands, where they occur on or near living or dried
terrestrial vegetation, generally at low elevations and in
close proximity to the shoreline. Some populations occur
at elevations of tens of meters, usually near the edges of
coastal cliffs. Populations of some normally coastal spe¬
cies may occasionally be found several kilometers inland,
generally on the windward sides of islands.
When the progenitors of Cerion were first isolated on
small, arid islands, most likely in the early to mid-Tertiary,
selection favored animals that were salt tolerant and able
to withstand heat, exposure to sun, and prolonged periods
of desiccation. Descendants of these animals colonized
the islands of the Greater Antilles along the GAARlandia
land bridge during the late Eocene-early Oligocene and
later the Bahamas via a stochastic accumulation of
hurricane-borne propagules (Ituarralde-Vinent, 2006:
figs. 6, 13; Harasewych, 2012: 123).
Although Cerion acherontis Roth and Hartman, 1998,
the Cretaceous ancestor of Cerionidae, was provisionally
described in the genus Cerion , its inclusion in Cerion
would render this genus paraphyletic due to the interven¬
ing phylogenetic positions of the genera Mexistrophia and
Brasilennea. We therefore propose Protocerion as a new
genus for this Cretaceous species.
Protocerion new genus
Type Species: Cerion acherontis Roth and Hartman,
1998. (By original designation)
Diagnosis: Shell of moderate size (~ 23 mm), pupiform
in shape, apically rounded, elongate-ovate (shell length/
shell width ~ 2.7). Protoconch and early whorls unknown.
Teleoconch of 4+ smooth, weakly convex whorls, with
impressed suture. Sculpture of faint growth lines. Umbili¬
cus imperforate. Aperture rounded, slightly higher than
wide, ~ 3/8 of shell length. Peristome adpressed, form¬
ing thin parietal callus and thickened, axial columellar
lip, smoothly rolled outward. Lacking columellar or pari¬
etal folds.
Distribution: Known only from a single, fractured and
compressed specimen (Figure 3) collected ~ 29.8 m above
the base of the Hell Creek Fonnation, Garfield County,
Montana (106°56'47" N, 47°34'10" N). Late Cretaceous
(Late Maastrichtian).
M.G. Harasewych et al., 2015
Page 161
Figure 3. The holotype and only known specimen of Cerion
acherontis Roth and Hartman, 1998, the type species of
Protocerion new genus. USNM 491763, Hell Creek Formation,
Garfield County, Montana [106°56'47" N, 47°34'10" W], Late
Cretaceous (Late Maastrichtian).
Remarks: As noted by Roth and Hartman (1998),
Protocerion acherontis more closely resembles the smooth-
shelled morphotype exemplified by the Recent species
Cerion incanum (Leidy, 1851) titan any of the ribbed
morphotypes (e.g., Cerion uva). Both Protocerion and
Mexistrophia lack columellar or axial folds, which are
present in Brasillenea and all species of Cerion. These
folds are an apomorphy for Brasillenea + Cerion that is
lacking in Protocerion and Mexistrophia. Their absence in
Protocerion distinguishes this genus from Cerion.
ACKNOWLEDGMENTS
We are most grateful to Dr. John Park for providing the
specimens of Microceramns pontificus used in this study.
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THE NAUTILUS 129(4): 163-168, 2015
Page 163
A new species of the genus Mexipyrgus Taylor, 1966
(Caenogastropoda: Truncatelloidea: Cochliopidae) from late
Holocene spring deposits in Viesca, Coahuila, Mexico
Alexander Czaja
Jose Luis Estrada-Rodriguez
Ulises Romero-Mendez
Faculty of Biological Sciences
Juarez University of the State of Durango (UJED)
35010 Gomez Palacio, Durango, MEXICO
aaczaja@hotmail .com
ABSTRACT
A new species of the genus Mexipyrgus Taylor, 1966
(Caenogastropoda: Truncatelloidea: Cochliopidae) is described
from two Holocene springs in Viesca, Coahuila, Mexico. Previ¬
ously, only one species was known in this genus, Mexipyrgus
carranzae Taylor, from freshwater springs and streams in the
Cuatrocienegas Valley in Coahuila, Mexico. Similar to the extant
species, the shells of Mexipyrgus viescaensis new species show
a high level of morphological variability.
Additional Keywords: Mexipyrgus carranzae, freshwater
gastropod, subfossil endemic
INTRODUCTION
The Chihuahuan Desert in United States and Mexico is
considered as a hotspot of molluscan diversity, with a high
number of endemic gastropod species (Hershler, 2011).
One of the most interesting sites in this desert is
Cuatrocienegas Valley in Coahuila, Mexico. The relatively
small valley contains more than 70 endemic species of
animals and plants, and is biologically tire most diverse
site in North America, in relation to endemism (Stein
et ah, 2000). The malacologist Dwight Willard Taylor,
who had presented a first monograph of the snails of
Cuatrocienegas, considered the mollusks of this site as
. .the most spectacularly endemic fauna of freshwater
snails known in the Western Hemisphere. . (Taylor, 1966).
The freshwater ecosystem of Cuatrocienegas includes
five endemic genera of hydrobiid snails ( Paludiscala ,
Coaim ilix , Mexithauma, Nymphophilus and Mexipyrgus) .
Fossil records of Cuatrocienegas endemic species from
other sites were reported only by Czaja et ah (2014a).
The authors discovered in die Valley of Sobaco, Coahuila,
a system of paleo-lakes with a malacofauna similar to
the modem snail communities of Cuatrocienegas. This
includes die endemic genus Coahuilix.
Mexipyrgus is a Mexican endemic freshwater hydro¬
biid gastropod that lives in springs, lakes, and the
Mezquites River in the Valley of Cuatrocienegas. Its main
morphological characteristic is a thickened, strongly sculp¬
tured shell widi color banding. All these shell features are
unusual for freshwater snails, especially within the family
Cochliopidae. Geographic variation in shell morphom¬
etry of Mexipyrgus was analyzed by Hershler and Hayek
(1988). Aspects of ecology and coevolution between
Mexipyrgus carranzae and its fish predator, Herichthys
minckleyi, have been subject of various investigations
(Smith, 1982; Tang and Roopnarine, 2003; Johnson et ah,
2007; Covich 2010; Chaves-Campos et ah, 2012).
In sediments of two recently dried-up springs near the
town Viesca, Coahuila, specimens of a second species of
the genus Mexipyrgus Taylor were collected (Figure 1).
The springs are located at the foot of the Sierra la
Cadena and belong to a system of several water bodies
that have provided water to small settlements at the
vicinity. Both sites contain an abundance of shells includ¬
ing several possibly new species of Tryonia, Pyrgulopsis
and Pyrgophorus (Czaja, personal data). Similar to
Mexipyrgus carranzae from Cuatrocienegas, the shells
from Viesca possess a high level of phenotypic variability.
The aim of the present study is to describe the new
species of Mexipyrgus and to show its similarities and
differences to the living species from Cuatrocienegas.
MATERIALS AND METHODS
Specimens of Mexipyrgus viescaensis new species were
collected from two recently dried-up springs near the
town Viesca, Coahuila, Mexico, approximately 2 km south
of the locality Venustiano Carranza (spring El Molino:
25°19/55" N, 102°55,49"W; spring Hacienda de Homos y
Carranza: 25°20'05" N, 102o57'39" W). The material
derives from outcrops on the edges of the springs. The
THE NAUTILUS, Vol. 129, No. 4
Page 164
103°0'0"W
102°45'0"W
102°30'0"W
102°15'0"W
102°0'0"W
103°30'0"W
. ^
103°15'0"W
SAN PEDRO
Legend
103°30'0"W 103°15'0"W 103°0'0"W 102°45'0''W 102°30'0"W 102°15'0"W 102°0'0"W
Figure 1. Map of the study area with localization of the springs of Viesca.
section is approximately 1.5 in thick and contains uncon¬
solidated sediments, small travertine pieces and shell
debris. The sub-recent (late Holocene) age of the super¬
ficial deposits is confirmed by severed reports and photo¬
graphs which document the desiccation of the springs
during the drought of 1958/59.
The sediments of both sites were screened through
0.5 mm and 0.3 mm sieves. The shells were photographed
with a Zeiss AxioCam ERe5s microscope-camera. The
shells of the new species were compared directly with
Mexipyrgus carranzae from the Malacologieal Collection
of the Faculty of Biological Sciences, Juarez State Univer¬
sity of Durango. All collected material is housed at the
same Faculty.
SYSTEMATICS
Class Gastropoda Cuvier, 1795
Subclass Caenogastropoda Cox, 1960
Superfamily Truneatelloidea Gray, 1840
Family Cochliopidae Tryon, 1866
Genus Mexipyrgus Taylor, 1966
Type Species: Mexipyrgus carranzae Taylor, 1966
(by original designation).
Mexipyrgus viescaensis new species
(Figures 2-11)
Diagnosis: Shell medium-sized, thick, white to beige
colored, conical to turriform, height 3.18-4.90nnn, width
1.99-2. 45mm (shell measurements in Table 1), with 5.25
to 6.25 whorls, whorls flattened, aperture elongate-ovate,
outer lip prosoeyrt, inner lip not adnate to parietal wall,
sculpture with strong spiral and axial elements, with prom¬
inent knobbed ribs or spines, sculpture variable.
Description: Shell medium-sized, thickened, conical
to turriform, with 5.00-6.25 whorls. Whorls flattened
with sutures not very impressed; lower whorls develop
a prominent spiral swelling anterior to suture (subsu-
tural cord). Height 3.18-4.90 mm, width 1.99-2.45 mm.
Aperture elongate-ovate, somewhat pyriform, angulate
25 WN 25°15'0"N 25°30'0"N 25°45'0"N 26 WN
A. Czaja et al., 2015
Page 165
Figures 2—11. Mexipyrgus viescaensis new species. 2-4. Holotype, UJMC-200. 5-6. Paratype 1, UJMC-201. 7. Specimen showing
spines, UJMC-202. 8-9. Paratype 2, UJMC-203. 10-11. Specimens with shovel-shaped spines. 10. UJMC-204. 11. UJMC-205.
above; inner lip not adnate to parietal wall forming a
narrow gap between lip and pariental wall; outer lip
strongly prosoeyrt; aperture 1.90 mm height and 1.25
mm wide (holotype). Protoeoneh and second whorl
smooth. Sculpture with spiral and axial elements begin¬
ning at third whorl; spiral cords close to suture; wave-like
axial ribs developing on third whorl, with prominent
knobs where they cross spiral cords; knobs at lower part
of whorls prominent, sometimes with shovel-shaped
spines emerging from sub-sutural knobs; after third
whorl knobbed ribs very prominent; sculpture reduced
on last part of body whorl. Axial growth lines prosoeyrt
to sinusoidal, prominent. Color beige or white, origi¬
nal coloring (periostracal color bands?) and operculum
not preserved.
Type Material: Holotype (Figures 1-3, 15), UJMC-
200, 4.15 mm height, 2.05 mm width, 6.25 whorls.
Paratype 1 (Figures 5, 6), UJMC-201, 4.22 mm height,
2.39 mm width, 6.00 whorls. Paratype 2 (Figures 8, 9),
UJMC-203, 4.09 mm height, 2.50 mm width, 5.25 whorls.
Alexander Czaja and Jose Luis Estrada-Rodrfguez coll.,
2014. All from type locality.
Type Locality: Spring Hacienda de Homos y Carranza ,
ea. 15 km west of the town Viesca, Coahuila, Mexico
(25° 20' 05" N, 102°57'39" W).
Stratum Typicum: Holocene (sub-recent).
Other Material Examined: More than 500 speci¬
mens from tlie type locality and 215 specimens from
spring El Molino.
Etymology: The new species is named after Viesca,
Coahuila, a small town near the dried springs.
Geographic Distribution: Endemic to the springs
Hacienda de Homos ij Carranza and El Molino near Viesca,
Coahuila, Mexico.
Remarks: The new species differs from other mem¬
bers of the family Cochliopidae by their strongly thick¬
ened and highly sculptured shells with spiral cords
and knobbed ribs, which allocate the species into the
genus Mexipyrgus. Some species of the genera Tryonia,
Pyrgophoms, and Lithococcus also have sculptured shells
but differ by the following characters: species of the first
Page 166
THE NAUTILUS, Vol. 129, No. 4
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two genera do not have knobbed ribs and the shells of
Lithocpccus are globose-trochoid.
Considerable variation of shell features occurs among
the populations of both sites where the species was
found. Variation showing ribs with reduced knobs is
shown on Figure 13. That morphotype possesses two
spiral cords with knobbed ribs close to the sutures. The
number of knobbed ribs on the whorls varies between 8
and more than 20 (Figures 5, 6, and 8) and, in particular,
specimens from Hacienda de Homos y Carranza Spring
have shells with numerous fine ribs (Figures 5 and 6).
Figures 7, 10, and 11 show another variation with pro¬
longed spines on the spiral cords. In some specimens
from the El Molino Spring these shovel-shaped spines
on the knobs are sometimes connected between the
sutures (Figure 10). This phenotype is the least similar
to the holotype and shells with this kind of sculpture are
smaller, reaching just 3.5 cm in length. Intergradation
occurs among the morphotypes from the same springs
but most shells closely resemble the holotype.
Originally six nominal species of Mexipyrgus were
described by Taylor (1966), based on characters like shell
size, shell sculpture and periostracal bands. But Hershler
(1985) could prove that all specimens of Mexipyrgus
from Cuatrocienegas belong to a single variable species
and that the differences in the shell features are depen¬
dent on environmental factors. According to Vermeij and
Covich (1978) and Covich (2010) Mexipyrgus and other
snails from Cuatrocienegas evolved in coevolution with
snail-eating cichlid fishes. Also the sub-recent material of
both springs of Viesca show a high level of phenotypic
variability among the populations in time and space. We
find in the same horizons with specimens of Mexipyrgus
pharyngeal teeth of fishes (probably cichlids). Approxi¬
mately 10% of all shells in the springs were perforated
(Figures 13-14). Similar holes in shells were described
recently by Rasser and Covich (2014) from Miocene
Lake Steinheim, Germany, and interpreted as perforations
of a fish predator. We suppose that the thickened and
strongly sculptured shells of Mexipyrgus of both springs
could be a direct adaptive response to strong fish preda¬
tion. Further studies on shells of the new Mexipyrgus
from Viesca will show more details of the predator-prey
interaction during the coevolution of these species.
Mexipyrgus viescaensis is clearly distinguished from
Mexipyrgus carranzae mainly by its size and by the sculp¬
ture of the shells. While shells of M. carranzae attain a
length up to 8.45mm (Hershler, 1985, p. 99), the largest
specimens from Viesca reach just 4.85mm. Shells of
Mexipyrgus carranzae are thicker (more than 0.25mm
thickness) than specimens from Viesca (less than 0.20mm
thickness). Most diverse and stronger is the sculpture of
shells of M. viescaensis with very prominent and strong
knobs on the spiral cords and shovel-shaped spines.
Spines on whorls have never been observed on living
snails from Cuatrocienegas. Another difference is that by
M. viescaensis the inner lip are not adnate to the parietal
wall forming a narrow gap between the lip and pariental
wall. Shells of M. carranzae lack this gap.
A. Czaja et al., 2015
Page 167
Figures 12-17. Mexipyrgus viescaensis new species, Mexipyrgus carranzae Taylor, from Poza Becerra, Cuatrocienegas Valley,
Coahuila, Mexico, and Tryonia hershleri Czaja and Estrada-Rodnguez from Paleolake Irritila, Coahuila, Mexico. 12. Mexipyrgus
carranzae Taylor, hole on the ab-apertural side on the shell. 13. Mexipyrgus viescaensis new species, hole on the apertural side on the
shell. 14. Mexipyrgus viescaensis new species, double holes on the ab-apertural side on the shell. 15. Mexipyrgus viescaensis new
species, Holotype (UJMC-200), protoconch view. 16. Mexipyrgus viescaensis new species from Viesca, Coahuila. 17. Tryonia
hershleri Czaja and Estrada-Rodnguez from Pleistocene Paleolake Irritila. Coahuila, Mexico.
The findings of a new species of Mexipyrgus shed new
light on tlie origin of the endemic M. carranzae from
Cuatrocienegas. Of die five endemic genera from Cuatro¬
cienegas Valley, four, Coahuilix, Paludiscala, Nymphophilus,
and Mexipyrgus, have already been found as (sub-) fossils
outside the basin (Hershler, 1985; Czaja et al., 2014a;
Czaja, personal data). Mexipyrgus is possibly a relict genus
which originally had a wider distribution with different
species. Interestingly, some morphotypes of M. viescaensis
show great similarity with shells of Tryonia hershleri
(Figures 16-17). This species with diiekened and strong
sculptured shells was newly reported from Late Pleisto¬
cene deposits near Viesca (Czaja and Estrada-Rodnguez,
2015). This similarity may be due to convergence but on
the odier hand, there are also molecular evidences that
both genera are closely related (Hershler et al., 1999;
Hershler et id., 2005). This might be proved only by fossil
findings in Cuatrocienegas Valley and Viesca. Future work
on sub-fossil shells in bodi localities will allow us to know
more on the evolutionary patii of the enigmatic genus
Mexipyrgus from northern Mexico.
ACKNOWLEDGMENTS
We thank Dr. Robert Hershler (Smithsonian) for valuable
discussions. A special thank you goes to Allen Kosinsld,
New Jersey, USA, for his assistance in the revision of the
text in English.
LITERATURE CITED
Chaves-Campos J., L.M. Coghill, M.A. Al-Salamah, T.J.
DeWitt, and S.G. Johnson. 2012. Field heritabilities and
lack of correlation of snail shell form and anti-predator
function estimated using Bayesian and maximum likelihood
methods. Evolutionary Ecology Research 14: 743-755.
Covich, A.P. 1976. Recent changes in molluscan species diver¬
sity of a large tropical lake (Lago de Peten, Guatemala).
Limnology and Oceanography 21: 51-59.
Covich, A.P. 2010: Winning die biodiversity arms race among
freshwater gastropods: competition and coexistence through
shell variability and predator avoidance. Hydrobiologia
653: 191-215. '
Czaja, A., J.L. Estrada-Rodnguez, and U. Romero-Mendez.
2014a. Freshwater mollusks of the Valley of Sobaco,
Coahuila, Northeastern Mexico - a subfossil ecosystem
similar to Cuatrocienegas. Boletin de la Sociedad Geologica
Mexicana 66 (3): 459-469.
Czaja, A., M.R. Palacios-Fest, J.L. Estrada-Rodnguez,
U. Romero-Mendez, and j.A. Alba-Avila. 2014b. Inland
Dunes mollusks fauna from the Paleolake Irritila in the
Comarca Lagunera, Coahuila, Northern Mexico. Boletin
de la Sociedad Geologica Mexicana, Vol. 66 (3): 541-551.
Czaja, A. and J.L. Estrada-Rodnguez. 2015. Two new species
of Tryonia (Caenogastropoda: Coehliopidae) from the Late
Pleistocene of Coahuila, Northern Mexico. The Nautilus
129: 83-89.
Hershler, R., 1985. Systematic revision of the Hvdrobiidae
(Gastropoda: Rissoacea) of the Cuatro Cienegas Basin,
Coahuila, Mexico. Malacologia 26: 31-123.
Hershler, R., and L.-A.C. Hayek. 1988. Shell Variation of
Springsnail Populations in the Cuatro Cienegas Basin,
Mexico: Preliminary Analysis of Limnocrene Fauna. The
Nautilus 102: 56-64.
Hershler, R., H.-P. Liu, and M. Mulvey. 1999. Phylogenetic
relationships within the aquatic snail genus Tryonia :
implications for biography of the North American
Southwest. Molecular Phylogenetics and Evolution 13:
377-391.
Hershler, R., M. Mulvey, and H.-P. Liu, P. 2005. Genetic varia¬
tion in die Desert Sringsnail (Tryonia porrecta): implicaeion
for reproductive mode and dispersal. Molecular Ecology
14: 1755-1765.
Hershler, R., H.-P. Liu, and J.J. Landye. 2011. New species and
records of springsnails (Caenogastropoda: Coehliopidae:
Tryonia) from the Chihuahuan Desert (Mexico and
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United States), an imperiled biodiversity hotspot. Zootaxa
3001: 1-32.
Johnson, S.G., C.D. Hulsey, and F.J. Garcia de Leon. 2007.
Spatial mosaic evolution of snail defensive traits. BMC
Evolutionary Biology 7 (50): 1-11.
Rasser, W. M. and A.P. Covich. 2014. Predation on freshwater
snails in Miocene Lake Steinheim: a trigger for intra-
lacustrine evolution? Lethaia 47 (6): 524-532.
Smith, D.C. 1982. Trophic ecology of the cichlid morphs of
Cuatro Cienegas, Mexico. M aster’s Thesis, University ol
Maine at Orono, pp. 46. http:/Avww.nativefishlab. net/library/
textpdf/21 141 .pdf
Stein, B.A., L.S. Kutner, and J.S. Adams. 2000. Precious Heri¬
tage: The Status of Biodiversity in the United States.
Oxford University Press, Oxford, pp. 399.
Tang, C.M. and P.D. Roopnarine. 2003. Complex morphological
variability in complex evaporitic systems: Thermal spring
snails from the Chihuahuan Desert, Mexico. Astrobiology
3: 597-607.
Taylor, D.W. 1966. A remarkable snail fauna from Coahuila,
Mexico. The Veliger 9: 152-228.
Vermeij, G.|. and A.P. Covich. 1978. Coevolution of fresh¬
water gastropods and their predators. American Naturalist
112:833-843.
THE NAUTILUS 129(4): 169-1 71 , 2015
Page 169
Intracapsular development in the freshwater gastropod
Chilina dombeiana (Bruguiere, 1789) (Gastropoda:
Hygrophila: Chilinidae)
Jessica Borquez
Departainento de Ecologfa
Facultad de Ciencias
Universidad Catolica de la Ssma.
Concepcion, CHILE
Universidad Catolica de la Ssma.
Concepcion, CHILE
[email protected]
Claudio Valdovinos
Departainento de Sistemas Acuaticos
Facultad de Ciencias Ambientales y
Centro de Ciencias Ambientales (EULA)
Universidad de Concepcion, CHILE
Antonio Brante1
Departainento de Ecologfa
Facultad de Ciencias y
Centro de Investigation en Biodiversidad
y Ambientes Sustentables (Cl BAS)
ABSTRACT
Chilina dombeiana (Bruguiere, 1789) is a native Chilean species
inhabiting freshwater and estuarine environments. In tire pres¬
ent study, a series of stages for embryonic development of the
embryo is described. The snails lay gelatinous and transparent
zig-zag-like string egg masses, ranging between 10 and 130 mm
in length. Each egg mass contains a variable number of embryos
ranging from 60 to 298 eggs with a mean density of 2.9 ■ mm-2
(± 0.7 SD). Embryos inside egg masses are individually encapsu¬
lated and embedded in a jellv matrix. Hatching as crawling juve¬
niles took place after 28 days, indicating that direct development
occurs in this species. In contrast to other freshwater pulmo-
nates, the well-developed operculum observed in C. dombeiana
from the veliger stage, suggest a marine ancestry for this species.
Additional Keywords: Chilinidae, Bio-Bio river, hermaphroditism
INTRODUCTION
Chilinidae is an ancient family of freshwater gastropods
endemic to South America (Jame et ah, 2010). The family
is monotypic, with Chilina including the primitive pul-
monate snails described by Gray in 1828 (Brace, 1983).
Chilina dombeiana (Burguiere, 1789) is a native Chilean
species that inhabits freshwater and estuarine environ¬
ments from approximately 35° S to 37° S (Valdovinos,
2006). Although this species is highly abundant in some
rivers and hikes, and may play an important ecological role
in freshwater ecosystems (Valdovinos et al. 2006), there is
little information about the basic biology of this organism.
Species of the order Hygrophila shows simultaneous
hermaphroditism and are capable of self-fertilization and/
or biparentality through cross-fertilization (Jame et al., 2010;
Nakadera et ah, 2014). However, there is no published
information on the reproductive behavior and embryonic
1 Corresponding Author
development of C. dombeiana. We describe herein the
intracapsular development of individuals of C. dombeiana
inhabiting a riverine environment in south-central Chile.
MATERIALS AND METHODS
At one site in the Bio-Bio river (36°49'22.43" S,
73° 6' 3 1.41" W) in Concepcion, Chile, 50 individuals of
C. dombeiana were collected by hand and transported to
the Facultad de Ciencias, Universidad Catolica de la Ssma.
Concepcion. In the laboratory, the C. dombeiana adults
were cultivated in individual plastic boxes (200 ml) filled
with freshwater collected from the sample site. Boxes
were constantly aerated with an air pump and the temper¬
ature was maintained at 18°C. The water inside the boxes
was changed every ten days. Once the individuals had laid
egg capsules, the relative duration of each developmental
stage (in days), a range of total number of egg by egg mass,
egg and embryo size, and embryonic development traits
were recorded. In order to make these measurements,
20 egg masses from different individuals were haphazardly
chosen and photographed every three days under a micro¬
scope ( lOx ; Olympus CX31) connected to a tablet com¬
puter equipped with a digitiil camera (Smart Pad 500).
Five stages of embryonic development were observed:
early embryo, trochophore, early veliger, late veliger, and
pre-hatching juvenile. Embryo size was measured with the
software Image measure (VMS3.5). In addition, when the
egg mass showed a string formation (Figures 1, 2), the total
number of embryos per clutch was estimated by counting
the number of embryos in a section of the egg mass and
extrapolating to the total egg mass size. This was done
using a binocular microscope (0.67x; Olympus CX31).
RESULTS AND DISCUSSION
Sexual maturity of the freshwater snail C. dombeiana is
reached within the first year of the individual’s life cycle
Page 170
THE NAUTILUS, Vol. 129, No. 4
Figures 1-10. Chilina dombeiana, egg capsules and intracapsular development. 1—4. Egg capsules. 1. Gelatinous egg mass (em)
attached to hard substrate. 2. Egg mass showing the general zig-zag string appearance. 3. Light micrograph showing the embryos (e)
contained within each egg capsule (ec). 4. Detail of the external layer (el) of the egg mass surrounding the egg capsules.
5-10. Intracapsular development. 5. Segmented egg. CA: egg capsule; EM: embryo. 6. Trochophore stage. EM: embryo. 7. Early
veliger stage. DG: digestive gland; EC: intracapsular fluid; SH: shell, VE: velum. 8. Veliger stage. DG: digestive gland; HE: heart;
MO: eyespot; ST: stomach. 9. Late veliger stage. FO: foot; SH: shell. 10. Pre-hatching juvenile. OP: operculum.
(C. Valdovinos; unpublished data). In our observations,
the smallest individual laying egg masses was 11.7 mm in
shell size. Individuals lay gelatinous and transparent egg
masses on rocks.
The egg masses had a zig-zag-like string formation
ranging between 10 and 130 mm in length (Figures 1, 2).
Witliin each egg mass, embryos were observed individually
encapsulated and embedded within a gelatinous matrix
(Figures 3, 4). The total number of embryos per egg
mass ranged from 60 to 298 with a mean density of
2.9 eggs • mm- (dt 0.7 SD). In the first 4 days of develop¬
ment, the early embryos were characterized by cleaved
eggs and embryos at the pre-trochophore stage with an
average size of 120.9 pm (± 11.8 SD; Figure 5; Table 1).
The embryos were yellow in color and were embebed in a
transparent intracapsular fluid. No aparent major move¬
ment of the embryos was noticed. Between days 5 and 6 of
cultivation, die embryos developed into early and late
trocophores had a mean average diameter of 139.4 pm (±
11.1 SD; Figure 6; Table 1). Between days 12 and 15, early
veliger embryos widi velum and noticeable development of
a very soft shell (average size of 235.5 pm ±8.1 SD;
Table 1) at die apical end were observed (Figure 7). In
addition, at this stage, digestive gland and active movement
of the larvae inside die capsules were visible. After 16-
19 days of development, the intracapsular fluid became
more transparent. Embryo at this late veliger stage had
a well-developed velum with an average shell size of
J. Borquez et al., 2015
Page 171
Table 1. Developmental time (days) and average size (pm) of embryos of the freshwater gastropod Chilina dombeiana at different
developmental stages. Thirty embryos were measured at each stage (excepting stage 2 were only 10 embryos were recorded).
394.5 pm (± 12.5 SD) (Figures 8, 9; Table 1). In addition,
coiling of the calcified shell, the dark pigmented eyespots,
the transparent operculum, and the digestive gland were
also evident. At this stage, the embryo had a high heart
rate. Between days 23-29, pre-hatching juveniles were
observed inside capsules. The shells were well calcified
and had a reduced velum (Figure 10). The foot muscle
and operculum were fully developed and individuals
actively moved inside the capsule. At this stage, the mean
shell size was 505.6 pm (± 11.4) (Table 1). Hatching as
crawling juveniles took place after 28 days, indicating that
direct development occurs in this species.
Adult freshwater pulmonates usually lack an opercu¬
lum, the exception being the family Amphibolidae (see
Golding, Ponder, and Byrne, 2007). Chilina dombeiana
differs completely from other non-amphibolid freshwater
pulmonates by having a well-developed operculum. This
suggests that C. dombeiana could have an evolutionary
origin from marine ancestry. Phylogenetic studies on this
group would shed light on the evolution of this species
and the potential adaptation of populations of these snails
to freshwater habitats (see Harry, 1964; Barker, 2001).
Although the range of C. dombeiana is restricted
latitudinally from 35° to 37° S (Valdovinos, 2006), this
species can be found abundantly at estuarine river
mouths all the way to upper river basins characterized
by well-oxygenated, clean, and cold water. As has been
suggested for other gastropod species (e.g., Pechenik,
1983; 1986; Rawlings, 1996; Pande et al., 2010), the direct
and encapsulated development of C. dombeiana would
allow this organism to tolerate extreme environmental
conditions during its entire embryonic phase. Although
direct development may lead to restricted dispersal of this
species, potentially dislodged egg masses or capsules from
the substrata would favor population connectivity and
gene flow via river currents. Considering that human use
of freshwater ecosystems causes the greatest negative
impact on these ecosystems (Strayer and Dudgeon,
2010), large anthropogenic perturbations could expose
freshwater invertebrates to specific threats that may result
in local extinction processes. Accordingly, Valdovinos
(2006) highlighted the vulnerability of the Chili nidae fam¬
ily due to significant habitat lost. In that study, the author
further emphasized the importance of conducting more
studies that contribute to the understanding of these
freshwater gastropods.
ACKNOWLEDGMENTS
This work was supported by the Postdoctoral
FONDECYT 3140561 to JB, FONDECYT 1130868 to
AB and the EULA Center, University of Concepcion
(PMBB Project). We thank Fernanda Oyarzun and Jose H.
Leal for the assistance in editing the photographs.
LITERATURE CITED
Barker, G.M. 2001. Gastropods on land: phylogeny, diversity and
adaptative morphology. In: Barker, G.M. (ed.) The Biology of
Terrestrial Molluscs. CABI Publishing, Wallingford, 146 pp.
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on the early Evolution of Pulmonate Gastropods. Philo¬
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reexamined, with prolegomena on the phylogeny of the
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THE NAUTILUS 129(4):172-174, 2015
Page 172
Trophon geversianus (Pallas, 1774): the first record of communal
egg masses in the muricid subfamily Trophoninae (Gastropoda)
Maria Jose Pio
Guido Pastorino
Museo Argentine) de Ciencias Naturales,
Av. Angel Gallardo 470 3° piso. Lab. 57
C1405DJR Buenos Aires, ARGENTINA
[email protected]
[email protected]
Gregory S. Herbert
School of Geosciences
University of South Florida
4202 E. Fowler Avenue
Tampa, FL 33620 USA
[email protected]
ABSTRACT
Reproductive strategies are thought to have played a major role in
the adaptive radiation of neogastropods, but data are still lacking
for important taxa, and the extent of plasticity in reproductive
strategies is not well understood. Females of the Patagonian
neogastropod Trophon geversianus are here shown for the first
time to have plastic egg capsule-laying strategies. Past work has
recorded only egg laying by isolated females, both in the intertidal
and in the lab. New observations of egg laying in subtidal
T. geversianus show that communal opposition also occurs in this
species, a first record of its land for the species and for the
subfamily Trophoninae. Given that communal opposition in other
species is commonly regarded as an adaptation to predation cues
or changing environment, we propose this reproductive strategy
in trophonines is an inducible response to environmental risk.
INTRODUCTION
In the course of gastropod evolution, many species have
evolved reproductive adaptations related to the encapsu¬
lation of embryos in response to diverse environmental
threats, such as predation (Rawling, 1994). Selective
pressures such as predation can affect gastropod embry¬
onic development, capsule morphology, and also opposi¬
tion behaPors, including effects that are phenotypically
inducible responses (among others: Harasewych, 1990;
Rawlings, 1990; 1994; Dumont et al., 2008; Bigatti et ah,
2010; Roche et ah, 2011).
Communal oviposition, which has evolved indepen¬
dently' in many animal groups, consists of several eonspe-
cific females depositing egg capsules on a shared substrate,
often resulting in large masses of eggs (D’Asaro, 1970;
D’Asaro, 1991; Swanson, 2004). Adaptive explanations for
this behavior include predator deterrence for embryos
and facilitation of social foraging (Waldman, 1982; Dumont
et ah, 2008). Communal opposition is a phylogenetically
widespread behaPor in gastropods (Thorson, 1940;
Giglioli, 1955; Gohar, and Eisawy, 1967; D’Asaro 1970;
Bandel 1975; Soliman, 1987; D’Asaro, 1991) and is
particularly well documented in the neogastropod
family Muricidae, including the subfamilies Muricinae,
Oeenebrinae and Rapaninae.
Trophon geversianus (Pallas 1774), type species of the
type genus of the subfamily Trophoninae and a good rep¬
resentative taxon of the Magellanic malacological prov¬
ince, occurs, lives, and feeds, on banks of mytilid bivalves
and barnacles on the rocky intertidal and subtidal zones
along the Patagonian coast (Pastorino, 2005). Its egg cap¬
sules are easily identifiable in the field due to their abun¬
dance and characteristic erect, rounded form and bright
yellow color. Capsules are variable in size (up to 22 mm in
height) and are usually attached by the female to a hard
substratum by a short and wide peduncle that rises from a
common basal membrane (Zaixso, 1973; Penehaszadeh,
1976). Observations on the number of capsules from a
single opposition event range from 2 to 22 per female
(Penehaszadeh, 1976). Cumplido et ah (2010) published
observations on the spawning behaPour of T. geversianus
in lab aquaria and reported no events of communal
spawning. Communal opposition has also not been
reported previously for T. geversianus in the wild.
MATERIALS AND METHODS
In this work, we report observations on communal spawn¬
ing of T. geversianus found living at approximately 20 m
depth on a mixed bottom of thick sand and pebbles near
to Puerto Almanza at Beagle Channel (54°53' 10.56" S,
67°42'6.55" W), Tierra del Fuego, Argentina on March
2010. One communal spawn mass, which was attached on
a sub-quadrate rock about 20 cm in width, was collected by
SCUBA and fixed with 5% formalin in seawater (Figure l).
RESULTS
Several communal spawn masses were observed depos¬
ited on rocks partially covered by barnacles. One of the
M.J. Pio et al., 2015
Page 173
Figures 1, 2. Trophon g eversianus egg masses from Tierra del
Fuego, Argentina. 1. Suhtidal communal spawn, fixed in 5%
formalin in seawater. 2. View of a subtidal communal oviposition
in situ.
largest masses consisted of a total of 250 capsules. Each
capsule had the escape aperture (mucoid plug) oriented
toward the upper exterior of the spawn mass and was
attached to the substrate by its own basal membrane.
The basal membrane of a capsule was never superimposed
over the basal membranes of adjacent capsules. A mini¬
mum of twelve females would have been required to form
a mass of 250 capsules, assuming each female produced
the maximum number of capsules recorded for a single
female T. geversianus at a single oviposition event (Zaixo,
1973; Penchaszadeh, 1976; D’Asaro, 1991; Cumplido et al.
2010) (Figure 2).
DISCUSSION
This study provides new information about the repro¬
ductive strategy of T. geversianus , which can be coded
for phylogenetic analyses or used to map the evolution of
reproductive strategies across M urieidae. Moreover, com¬
munal spawn masses of this type have not been observed
previously at the intertidal and subtidal levels where
T. geversianus usually occurs, indicating that females
may modify their oviposition habit in response to envi¬
ronmental cues.
The area where these egg capsules were collected is
not exposed, is relatively quiet, and moderately shallow.
No strong waves or active winds affect the coast at
Puerto Almanza, which is very different from the typical
intertidal conditions of the rocky shores along the
Patagonian coast. Also, the substrate is a mix of free
boulders, fine-grained sand, and large and immobile rocks.
In this habitat, large rocks are the only suitable substrate
on which to attach the capsules. This difference in envi¬
ronmental conditions in between intertidal anti subtidal
areas may exphiin the occurrence of communal spawning.
The conditions are also favourable for gastropod preda¬
tors, and it would be valuable to test whether the unusual
spawning behaviour for T. geversianus influences repro¬
ductive success of adults.
AC K N OWLE DGMENTS
Authors would like to thank P. Penchaszadeh for his
helpful advice on a first draft of the manuscript. To
reviewers Yuri Kantor and Jerry Harasewych, for the
thoughtful comments that contributed to improve the
manuscript. To Diego Urteaga for his assistance during
field work. We acknowledge funding by National
Scientific and Technical Research Council (CONICET)
from Argentina.
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Determining the date of publication for
Contradens Haas and Uniandra Haas
(Bivalvia: Unionidae)
Establishing the date of publication for a genus or species
name can be a case for Sherlock Holmes, especially if
the work was published in severed sections over time. The
dates of publication of the Sytematisches Conchylien-
Cabinet von Martini and Chemnitz are confusing, but
dates of publication for Haas’s (1910-1920) volume in this
set have been presented by Haas (1923: 203) listed by
lieferung, signatures (Bogen) in each lieferung, page num¬
bers, and the year published. These data were summa¬
rized and repeated by Smith and England (1937), Johnson
(1968), Welter-Schultes (1999: 190) and Coan and Kabat
(2015). However, these authors used the dates of the
Lieferung printed on the cover and not the dates printed
on each signature. Solem (1967: 80) examined the copy of
Die Unioniden available in Chicago and reported dates
listed on the first page of each of the 43 signatures that
make up the complete volume, but combined some of the
signatures into a single citation. The copy of Dig Unioniden
in the Library of the Academy of Natural Sciences of
Drexel University of Philadelphia was examined for the
dates of publication found on the first page of each of the
43 signatures, combined into 13 Heft, or numbered parts,
and lieferung. Each Heft/liefening was published with a
separate title page and year of publication and each
contained 2 or more signatures and plates (Table 1). Dates
of publication on the title page match the dates of the
included signatures for all but three of the heft (47, 51,
and 52), the date on the title page does not correspond to
dates on the signatures (Table 1). The back cover of each
Heft/liefening listed the plates and figures contained and
the signature numbers included in the Heft.
There is confusion around the original date of publica¬
tion of the generic names Contradens Haas and Uniandra
Haas. Many authors have used different dates for the
description for both genera. The following ehresonymy
summarizes the various dates used for both Contradens
and Uniandra-.
Contradens Haas, 1911: Graf and Cummings, 2015
[4 September, 2015],
Contradens Haas, 1912: Simpson 1914: 1005; Van Benthem-
Jutdng, 1953: 33.
Contradens Haas, 1913: Ortmann, 1917:107; Thiele, 1935:
821; Neave 1939: 825; Mode!’., 1942: 189; Stansbery
and Soehngen, 1964: 17; Solem 1967: 109; Haas 1969a:
96; Haas 1969b: N420; Starobogatov, 1970: 64; Brandt,
1974: 289; Vokes, 1980:73; Vaught, 1989: 122; Millard,
2003: 1120; Graf and Cummings, 2006: 393; Graf and
Cummings, 2007: 310; Bouchet and Rocroi, 2010: 30;
AnimalBase, 2015; Graf and Cummings, 2015 [12 May
2015]; ITIS, 2015 [11 September 2015]; GB1F, 2015
[11 September 2015].
Uniandra Haas, 1912: Neave, 1940: 612; Haas, 1969a:
73; Haas, 1969b: N423; Brandt, 1974: 289; Vokes,
1980: 78; Vaught, 1989: 122; Millard, 2003: 1124;
GBIF, 2015 [11 September 2015],
Uniandra Haas, 1913: Solem, 1967: 132; Starobogatov,
1970: 64; Graf and Cummings, 2015 [12 May, 2015;
4 September 2015]; ITIS, 2015 [11 September 2015];
FADA, 2015 [11 September 2015],
Model! (1942:189) recognized the genus Contradens
Haas, 1913 and erected the new subfamily Contradentinae
Model!, 1942. Confusion was further compounded by
Brandt (1974: 289) as to the priority of the two names
and the type species of Uniandra. Brandt used Uniandra
Haas, 1912 and included Contradens Haas, 1913 as a
junior synonym, incorrectly listing the type species of
Uniandra as Unio contradens Lea, 1838. However, neither
genus was listed in the online searchable version of Neave
(2015: 14 May 2015).
Graf and Cummings (2015: 12 May 2015) use Contradens
Haas, 1913, noting Haas (1911: pi. 18) as a nomen nudum,
referring to the description by I Lias, 1913 in Conehylien
Cabinet (page 173), but take the date of publication from
Haas (1913). They subsequently changed the date of
Contradens to Haas, 1911 in the version on the web with
no explanation (4 Sept 2015).
The generic name Contradens Haas first appears in
plate captions for plates 18-22 in Conehylien Cabinet
(Haas, 1910-1920). Plates 18-23 and pages 81-88 were
published on 1 July 1911 (Table 1). There are no dates
on any of the plates, so they take the latest date of publi¬
cation of the last signature in Heft 44 (Table 1). The
generic name Contradens was associated with 15 previ¬
ously published species and subspecies names. The
generic name Contradens was thus available from the
date of publication of the plates. This is considered an
indication for a genus named before 1931 under ICZN
Code articles 12.1, 12.2. The plates do not have a date on
them but are included in Heft 44 and the last signature in
this heft has a date of 1 July 1911.
Additionally, Heft 44 was stamped on the cover as
received by the Library at the Academy of Natural
Sciences of Philadelphia as received 16 October 1911.
The plates could not have been published after the date
stamped received in Philadelphia.
Contradens was fully described and Unio contradens
Lea, 1838 subsequently designated as the type species by
Haas (1913a: 35-36). Date of publication of this article
was listed as 27 January, 1913 (Solem, 1967: 83). Haas
A. E. Bogan, 2015
Page 177
acknowledged for his helpful comments. The Library at
the Academy of Natural Sciences of Drexel University,
Philadelphia, PA is thanked for access to their copy of
Haas (1910-1920).
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Arthur E. Bogan
North Carolina Museum of Natural Sciences
11 West Jones Street
Raleigh NC 27601 USA
[email protected]
Oscar Garcia- Alvarez, Luitfried von Salvini-Plawen,
Victoriano Urgorri, and Jesus S. Troncoso. 2014.
Fauna Iberica Volume 38 - Mollusca: Solenogastres,
Caudofoveata, Monoplacophora. Museo Nacional de
Ciencias N at u rales & Consejo Superior de Investigaeiones
Cientificas, Madrid 2014, 295 pp. 978-84-00-09870-4,
27 €. [In Spanish]
This impressive volume compiles vast amounts of data
on three of the least understood classes of molluscs,
Solenogastres (= Neomeniomorpha), Caudofoveata
(= Chaetodennomorpha) (together constituting Aplaeophora)
and Monoplacophora, and has been compiled by lead¬
ing experts O. Garcfa-Alvarez, L. v. Salvini-Plawen, J. S.
Troncoso and V. Urgorri, all of whom have published
extensively on the Iberian fauna of these groups. Sadly,
Professor Luitfried von Salvini-Plawen passed away on
October 20tl', 2014, while the book was in production,
so he never got to see this gem of malacology. Beautifully
bound in crimson cloth with the elegant black book cover
of the Iberian Fauna series, this book presents a state-
of-the-art compilation on these three molluscan classes.
Written in Spanish and profusely illustrated, this consti¬
tutes the most comprehensive treatment of each of these
classes of mollusks, synthesizing anatomical, ecological
and systematic aspects of the three groups.
The book begins with a 20-page general introduction
to the phylum Mollusca, including historical accounts of
the knowledge of the group and utilization by humans,
including a brief section introducing the aplacophoran
mollusks. These first pages make for very good gen¬
eral reading for those interested in delving into the
phylum Mollusca. The book is then divided into its three
main sections (with different authors), Solenogastres
(pp. 31-163), Caudofoveata (pp. 165-220) and Mono¬
placophora (pp. 221-252), followed by an Appendix
and four colour plates. The three main sections are
thorough taxonomic treatments of Solenogastres,
Caudofoveata, and Monoplacophora in the Iberian
waters down to 3,000 m depth. Each section com¬
prises anatomical and taxonomic treatments including
accounts for the natural history, collection, and preserva¬
tion methods. The book treats 42 Iberian Solenogastres
species grouped in 29 genera belonging to 14 families,
with taxonomic keys navigating through the taxonomic
ranks down to each species. All the taxa are plentifully
illustrated, including line drawings of cross sections,
sclerites and radula, often accompanied by scanning
electron micrographs. Descriptions, geographic distri¬
bution, and the biology of each species are provided.
For those unfamiliar with locality names, it would have
been very useful to have maps with distribution data.
The next section comprises 14 Iberian Caudofoveata
in four genera and three families, following the same
structure as the previous section, including grayscale
photographs of the animals (sadly, these are not of the
best quality), and line drawings of sclerites and radular
apparatus. Unfortunately, the quality of the illustra¬
tions for the Caudofoveata section is not comparable
to that of the Solenogastres part, despite both sharing
two authors. Finally, the two Iberian Monoplacophora
are described, with their key to species; Laevipilina
cachuchensis is described and illustrated based on
scanning electron micrographs of the shell and radula
(curiously, the radula is also presented as a line draw¬
ing), while L. rolani is illustrated by line drawings only
(both, the shell and the radula). Appendix 1 provides an
extensive nomenelatural list of synonyms and prior name
combinations, and is co-authored by M.A. Alonso-
Zarazaga, a member of the International Commission
on Zoological Nomenclature. The book ends with four
color plates with live photographs of 13 Solenogastres,
4 Caudofoveata, and L. cachuchensis .
This is a book that many molluscan workers, and all of
those interested in aplacophorans, must have. No com¬
parative thorough modern treatment of these groups is
available for any region of the world, and perhaps the
only one that comes close is |ones and Baxter (1987), but
the Iberian Fauna volume includes much more infor¬
mation for each species and many more species. This
volume can be purchased in the Spanish Consejo Supe¬
rior de Investigaeiones Cientificas publications portal.
A good level of Spanish is necessary both for reading
the book and for navigating through the website where
it can be purchased.
LITERATURE CITED
Jones, A.M. and J.M. Baxter. 1987 Molluscs: Caudofoveata,
Solenogastres, Polyplacophora and Scaphopoda: Keys and
Notes for the Identification of Species. Synopses of the
British Fauna, no. 37. The Linnean Society of London &
The Estuarine and Brackish-Water Sciences Association
by E. J. Brill/Dr. W. Baekhuys, 123 pp.
Gonzalo Giribet
Department of Organismic and Evolutionary Biology
Harvard University
Cambridge, MA 02138 USA
[email protected]
THE0NAUTILUS
Volume 129
2015
AUTHOR INDEX
Amanoj K . 43
Araujo, R . 126
Auffenberc, K . 54
Bogan, A. E . 175
Borquez, J . 169
Brante, A . 169
Campbell, K . 43
Campos, M . 126
Chrpa, M.E . 23
Czaja, A . 83, 163
Estrada-Rodri'guez, J.L . 83,163
Feo, C . 126
Fraussen, K . 95
Garcia, E.F. . 90, 136
Giribet, G . 179
Goddard, J.H.R . 31
Grimm, B.L . 77
Guller, M . 71
Harasewych, M.G . 156
Harding, j.M . 77
Hartman, J.H . 1
Houart, R . 143
Jenkins, R.G . 43
Little, C.T.S . 43
Lopez-Vera, E . 156
Marshall, B.A . 140
McLean, J.H . 118
Oleinik, A.0 . 23
Portell, R.W. . 54
Pou, Q . 126
Romero-Mendez, U . 163
Saether, K.P. . 43
Slapcinsky, J . 54
Spero, H.J . 77
Squires, R.L . 63
Straube, B.A . 77
Thompson, F.G . 156
Tracey, D . 140
Valdes, A . 31, 95
Valdovinos, C . 169
Vermeij, G.J . 77
Waiters, G.T. . 118
Windsor, A. M . 156
Zelaya, D.G . 71
NEWTAXA PROPOSED IN VOLUME 129
GASTROPODA
Bastropia llajasensis Squires, 2015, new species (Columbellidae, fossil) . 65
Campeloma acroterion Hartman, 2015, new species (Viviparidae, fossil) . 4
Chicoreus ( Chicopinnatus ) arbaguil Houart, 2015, new species (Muricidae) . 147
Chicoreus {Chicopinnatus) dhannai Houart, 2015, new species (Muricidae) . 149
Latiromitra niveobabelis Garcia, 2015, new species (Ptychatractidae) . 90
Mexipmgus viscaensis new species, Czaja, Estrada-Rodn'guez, and Mendez, 2015 (Cochliopidae) . 164
Protocerion Harasewych, Windsor, Lopez-Vera, and Thompson, 2015, new genus (Cerioniidae, fossil) . 160
Timbellus corbariae Houart, 2015, new species (Muricidae) . 144
Tocobaga Auffenberg, Slapcinsky, and Portell, 2015, new genus (Odontostomidae, fossil) . 55
Tryonia hershleri Czaja and Estrada-Rodn'guez, 2015, new species (Cochliopidae, fossil) . 84
Tnjonia pseudocircumstriata Czaja and Estrada-Rodn'guez, 2015, new species (Cochliopidae, fossil) . 86
Vivipanis purgatorius Hartman, 2015, new species (Viviparidae, fossil) . 8
Vivipanis codomorphus Hartman, 2015, new species (Viviparidae, fossil) . 14
BIVALVIA
Thyasira (Thyasira) beui Amano, Little, Campbell, Jenkins, and Saether, 2015, new species (Thyasiridae, fossil)
47
REVIEWERS FOR VOLUME 129
Beu, Alan G.
Rieler, Rudiger
Bigatti, Gregorio
Bogan, Arthur E.
Goan, Eugene V.
Cummings, Kevin S.
DeMaintenon, Marta
Fallon, Philip
Fedosov, Alexander
Garcia, Emilio F.
Groves, Lindsey
Harasewych, M.G.
Healy, John
Hershler, Robert
Kan tor, Yuri I.
Kiel, Steffen
Koehler, Frank
Malaquias, Manuel
Marquardt, William H.
Marshall, Bruce A.
Mikkelsen, Paula M.
Miloslavich, Patricia
Oliveira, Cleo
Oliverio, Marco
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CULTURE
BUILDS
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FLORIDA DEPARTMENT 0/ STATE
DIVISION oE CULTURAL AFFAIRS
Roopnarine, Peter
Roth, Barry
Ter Poorten, Jan Johan
Thompson, Fred G.
Valdes, Angel
Valentich-Scott, Paul
Verineij, Geerat J.
Von Cosel, Rudo
Von Rintelen, Thomas
Watters, G. Thomas
Wellington, A.
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