THE NAUTI
4 | Volume 133, Number 2
‘ NS August 8, 2019
CAT 99.132
INVZ ISSN 0028-1344
A quarterly devoted
to malacology.
EDITOR-IN-CHIEF
José H. Leal
The Bailey-Matthews National
Shell Museum
3075 Sanibel-Captiva Road
Sanibel, FL 33957 USA
EDITOR EMERITUS
M. G. Harasewych
Department of Invertebrate Zoology
National Museum of
Natural History
Smithsonian Institution
Washington, DC 20560 USA
CONSULTING EDITORS
Ridiger Bieler
Department of Invertebrates
Field Museum of
Natural History
Chicago, IL 60605 USA
Arthur E. Bogan
North Carolina State Museum of
Natural Sciences
Raleigh, NC 27626 USA
Philippe Bouchet
Laboratoire de Biologie des
Invertébrés Marins et Malacologie
Muséum National d'Histoire Naturelle
55, rue Buffon
Paris, 75005 FRANCE
Robert H. Cowie
Center for Conservation Research
and Training
University of Hawaii
3050 Maile Way, Gilmore 409
Honolulu, HI 96822 USA
Kenneth A. Hayes
Department of Biology
Howard University
Washington, DC 20001 USA
Steffen Kiel
Department of Paleobiology
Swedish Museum of Natural History
Box 50007
104 05 Stockholm, SWEDEN
Harry G. Lee
4132 Ortega Forest Drive
Jacksonville, FL 32210 USA
Charles Lydeard
Biodiversity and Systematics
Department of Biological Sciences
University of Alabama
Tuscaloosa, AL 35487 USA
Bruce A. Marshall
Museum of New Zealand
Te Papa Tongarewa
P.O. Box 467
Wellington, NEW ZEALAND
Paula M. Mikkelsen
Paleontological Research
Institution
1259 Trumansburg Road
Ithaca, NY 14850 USA
Diarmaid O Foighil
Museum of Zoology and Department
of Biology
University of Michigan
Ann Arbor, MI 48109-1079 USA
Gustav Paulay
Florida Museum of Natural History
University of Florida
Gainesville, FL 32611-2035 USA
Gary Rosenberg
Department of Mollusks
The Academy of Natural Sciences
1900 Benjamin Franklin Parkway
Philadelphia, PA 19103 USA
Elizabeth Shea
Mollusk Department
Delaware Museum of
Natural History
Wilmington, DE 19807 USA
Angel Valdés
Department of Malacology
Natural History Museum
of Los Angeles County
900 Exposition Boulevard
Los Angeles, CA 90007 USA
Geerat J. Vermeij
Department of Geology
University of California at Davis
Davis, CA 95616 USA
G. Thomas Watters
Aquatic Ecology Laboratory
1314 Kinnear Road
Columbus, OH 43212-1194 USA
SUBSCRIPTION INFORMATION
The subscription rate for volume
133 (2019) is US $65.00 for
individuals, US $102.00 for
institutions. Postage outside the
United States is an additional US
$10.00 for regular mail and US
$28.00 for air delivery. All orders
should be accompanied by payment
and sent to: THE NAUTILUS, P.O.
Box 1580, Sanibel, FL 33957, USA,
(239) 395-2233.
Change of address: Please inform
the publisher of your new address at
least 6 weeks in advance. All
communications should include both
old and new addresses (with zip
codes) and state the effective date.
THE NAUTILUS (ISSN 0028-1344)
is published quarterly by The Bailey-
Matthews National Shell Museum,
3075 Sanibel-Captiva Road, Sanibel,
FL 33957.
Periodicals postage paid at Sanibel,
FL, and additional mailing offices.
POSTMASTER: Send address
changes to: THE NAUTILUS
P.O. Box 1580
Sanibel, FL 33957
fer Fi LU 5
Volume 133, Number 2
August 8, 2019
ISSN 0028-1344
CONTENTS
Alan R. Kabat The Red Sea Mollusca described by Deshayes in Laborde’s Voyage de
Eugene V. Coan Arable PE rZe (MBBOAN BBD) cocoocovsooovcancavsbovevonnconseeaneoononsononeesseeonaeaDEeebREeAbod00N00006 31
J.G.M. Raven Crepidula fornicata (Linnaeus, 1758) (Gastropoda: Calyptraeidae) as
a hermit crab commensal in the North Sea wee cccccccecececeseeeeessesceccececeeevenssttseseees 40
Kazutaka Amano The Miocene to Recent biogeographic history of vesicomyid bivalves in
Yusuke Miyajima [iDEN Weill {Seo IER TECONCIS OE UNS TANTTETT cncococoscensocoencos0zos5056804000¢90000000909000090000 48
Robert G. Jenkins
Steffen Kiel
Kazutaka Amano Two warm-water species of Trochoidea (Gastropoda) from Pliocene
deposits on the Japan Sea side of Honshu, Japan, with remarks on the
influence of the onset of Northern Hemisphere glaciation ...........0....0...0000 57
2
THE NAUTILUS 133(2):31-39, 2019
Page 31
The Red Sea Mollusca described by Deshayes in Laborde’s
Voyage de l'Arabie Pétrée (1830-1834)
Alan R. Kabat!
Museum of Comparative Zoology
Harvard University
Cambridge, MA 02138 USA
alankabat@aol.com
Eugene V. Coan”
genecoan@gmail.com.
Santa Barbara Museum of Natural History
Santa Barbara, CA 93105 USA
ee ————E—E————EEEE=
ABSTRACT
The French malacologist Gérard Paul Deshayes described 13
new species of marine mollusks collected in 1828 from the
northern Red Sea by two French explorers, Léon de Laborde
and Louis Maurice Adolphe Linant de Bellefonds. His work was
published in an expedition volume authored by Laborde. We
determine the correct publication date and authorship attri-
bution for these new species: Deshayes, in Laborde, 1833. The
current systematic status of these species 1s discussed: seven are
currently accepted as valid, five are junior synonyms, and one is a
nomen dubium. Of the twelve known species, four are endemic
to the Red Sea, four are limited to the western Indian Ocean, and
four have broader Indo-Pacific distributions. This paper also
discusses the publication history of the botanical chapter of this
work, authored by Alire Raffeneau-Delile.
Tn
INTRODUCTION
An important but overlooked and frequently misdated
expedition volume is Laborde’s Voyage de l’Arabie Pétrée,
which contains the results of an expedition in 1828 from
Cairo, Egypt, to Petra, then in the Ottoman Empire and
now in modern Jordan. This expedition was undertaken by
two French explorers, Léon de Laborde (1807-1869) and
Louis Maurice Adolphe Linant de Bellefonds (1799-
1883). Laborde alone authored the expedition narrative,
published in a lavishly illustrated folio volume, which also
included a list of 13 new species of marine mollusks from
the Red Sea illustrated on single plate and described by
Gérard Paul Deshayes (1796-1875) and seven pages on
the botany of that region, including several new plant
species, some illustrated, described by Alire Raffeneau-
Delile (1778-1850).
We demonstrate that the natural history section of
Laborde was published in 1833, so that the new taxa
should be dated to 1833 (and not either 1830 or 1834,
1] .
Research Associate
9 .
~ Research Associate
as usually stated, see below), and should be attributed
to either “Deshayes, in Laborde, 1833” (mollusks) or
“Delile, in Laborde, 1833” (plants). Although Linant was
one of the two leaders of the expedition, Linant was
neither an author nor an editor of the expedition volume,
so that Linant should not be credited in that capacity,
contrary to numerous researchers who attributed this
work and its new species to “Laborde & Linant.”
Laborde’s Voyage
As this volume was published in a large folio size, 15 « 22
inches (approximately 43 59 cm), and natural history
comprised less than 10% of the text pages, and only three
of 69 plates, it is relatively rare in natural history libraries
and was inaccessible to most researchers until recently
digitized. Another complicating factor is that while the title
page is dated 1830, two of the maps at the end are dated
1833 and 1834, indicating that it was published over a
period of three or four years, so that subsequent re-
searchers have variously cited this work as “1830,” “1833,”
or “1834.”
The primary purpose of the Voyage was to provide a
narrative of Laborde’s and Linant’s travels across the
Sinai Peninsula to the archaeologically significant mon-
uments and ruins in Petra, a region that had then been
little explored by Europeans (Augé and de Bellefonds,
1990; 1994). The text of the Voyage has the following
components:
Half-title page, with “explication du frontispiece,” two
unpaginated pages (1 leaf);
Title page, two unpaginated pp. (1 leaf);
Dedication to “Guillaume II” (William II (1777-1847),
Elector of Hesse from 1821 to 1847), two unpaginated
pp. (1 leaf, verso blank) [= iil:
Preface, two unpaginated pp. (1 leaf) [= iii-iv];
“Introduction” (pp. 1-36);
“Précis du voyage et explication des planches” (pp. 37—72)
[page 65 has the text of the molluscan species]:
“Cartes de l Arabie Pétrée — Journal topographique de Suez
a lAccabah” (pp. 73-80):
Page 32
“Flore de lArabie Pétrée” (pp. 81-87) [page 88 is blank
except for the printer's mark]; and
“Liste des planches et indication ot elles se trouvent
expliquées” (one unpaginated page following page 88).
The book is accompanied by 69 full-page plates as well as a
number of text figures. In most of the copies examined,
including the online (scanned) version, the 69 plates are
bound at the end. One of the two copies in the New York
Public Library has the plates interspersed throughout the
text, with the plate(s) bound following the specific page on
which each plate is cited, with the maps alone at the very
end. Plate 1 is of Laborde himself, in “Arabian desert”
costume: “Arabe du désert (portrait de auteur dans son
costume de voyage ).” Plates 2 to 62 are archaeological and
scenery plates showing various sites in the Sinai and Petra.
Plates 63 to 65 are the three natural history plates, de-
voted to a mammal, the rock hyrax (plate 63), plants (plate
64) and mollusks (plate 65). Plates 66 to 69 are maps of the
region, including one showing the routes of earlier
expeditions.
The only indications of publication dates in the entire
volume are on the title page (dated 1830), Plate 67 (a
map with a date of 1833), and Plate 69 (an oversize map
not included in the online version, which has a date of
1834). Biographical accounts of Laborde and/or Linant
have usually cited this publication as “1830” (e.g., Goby,
1981: 474; Kornrumpf and Kornrumpf, 1998: 219; Kurz
and Linant de Bellefonds, 1998: 68: Labib, 1961:
112-116, pls. 13-14; Mazuel, 1937: 99-103); Pfullman
(2001: 176-178) cited this as “1830-33,” without expla-
nation; Anonymous (1834: 150) stated, without details, that
“La publication a eu lieu par livraisons successive de 1830 a
1833.”
The key to determining the publication dates is the
Bibliographie de la irae e, which Stafleu and Cowan
(1976: xxi) described as an invaluable resource, even
though, ironically, they did not check it for the dates of
Laborde’s V. oyage:
Bibliographie de la France . . . Important source for dating
French publications. The weekly issues contain lists of
new books, pamflets [sic] and of instalments of serial
publications. The date on which BF lists a book pub-
lished in Paris can usually be taken as that of actual
publication (Stafleu and Cowan, 1976: xxi).
Fortunately, the Bibliographie de la France allows partial
reconstruction of the publication history of Laborde’s
Voyage, because it specifies the plate(s) and “feuilles”
(leaves, each leaf corresponding to 2 text pages) for this
book. The “feuille” numbering is given at the lower right
corner of each odd- pampered page, so that aes 1 thes
feuille number 1; page 3 has feuille number 2, and so on
(however, the prefatory material and the list of plates do
not have feuille numbers).
Interestingly, only some of the plates were issued first in
1830, and the publisher's description as cited in the
Bibliographie de la France initially stated that a separate,
smaller octavo-size text volume would follow. Instead, the
THE NAUTILUS, Vol. 133, No. 2
text was ultimately published in 1833 in the same large
folio size as the plates. As set forth below, this source
confirms that 29 plates were published in 1830, while at
least 31 plates and text pages 1-88 were published in 1833:
Bibliographie
de la France Date Laborde, Voyage
19(4): 89 23 January Livraison 1, 4 pls.
1830
19(10): 157 6 March 1830 Livraison 2, 5 pls.
eek 2 Mil 17 April 1830 Livraison 3, 5 pls.
19(21): 333 22 May 1830 Livraison 4, 5 pls.
19(29): 490 17 July 1830 Livraison 5, 5 pls.
19(38): 626 18 September Livraison 6, 5 pls.
1830
22(32): 10 August Livraisons 7-9,
499-500 1833 feuilles 1-18
[= pp. 1-36]:
12 pls.
22(48): 757 30 November Livraisons 10-12,
1833 feuilles 19-44
[= pp. 37-88
and preface; list
of plates(?)]; 19 as.
No further n/a [9 pls.; title pages (?)]
citations
However, further confusing matters is that the plates were
not numbered when they were issued, nor were the plates
issued in the same sequence as they were finally num-
bered. Instead, the plates were not numbered until 1833,
with the issuance of the list of plates, which required the
purchasers to re-arrange the plates in the correct nu-
merical sequence, and (some) purchasers then hand-
numbered the plates before having them bound in the
correct sequence.
In an “Avis” (guide) to the reader, inserted in one of the
copies in the Library of Congress, the publisher provided
an explanation for the delay in publication and a listing of
the plates in Livraisons 10, 11, and 12 (Anonymous, 1833).
The publisher wrote: “M. Collin, graveur distingué ...
étant tombé malade fut force d’ interrompre ses irae aux; il
n/a pales reprendre que dupuis peu de temps et ne pourra
les terminer que vers la fin de lannée” [Mr. Collin, a
distinguished engraver, became sick and was obliged to
interrupt his lelboress he has not been able to resume 5 dhe
until recently, and cannot finish them until the end of the
year].
The “Avis” provided details on the contents of
Livraisons 10 to 12, with the caveats that the listing of
plates in the “Avis” does not include the plate number,
and the description of each plate in the “Avis” does not
always match the actual title of the plate in the final “list
of plates” — but the “Avis” does confirm that the three
natural history plates (63, 64, and 65) were issued in
1833:
A.R. Kabat and E.V. Coan, 2019
“Dédicace; Six
feuilles de texte”
[pp. 37-48]
6 plates (including
what are now
plates 5, 6, 46, 47,
either 57 or 58,
and 63).
§ plates (including
what are now
plates 1, 4, 5, 31,
and 64).
7 plates (including
what are now
plates 10, 11 or
12, 25, 53 or 55,
65, 66, and 68).
Livraison 10
“Douze feuilles
de texte”
[pp. 49-72]
Livraison 1]
“Préface et table:
Huit feuilles de
texte” [pp. 73-88]
Livraison 12
The listing of plates in the “Avis°—21 plates for Livraisons
10 to 12—is inconsistent with the Bibliographie de
France, which indicates only 19 plates for Those Livraisons,
suggesting that two plates listed in the “Avis” were not issued
until 1834.
Vicaire (1900: 758-759) cited the Bibliographie de la
France in noting that this was issued in 12 livraisons from
1830 to 1833, but did not notice that there had to be at
least one more Livraison (for nine of the plates), or that
the oversize map was dated 1834. Augé and de Bellefonds
(1994: 20-21) briefly discussed the publication history of
the Voyage, and stated that the last text section on botany
(pp. 81-87) was sent to subscribers in 1833, with the
oversize map (pl. 69) being sent in 1834: “la derniére
livraison, comprenant la Flore de l'Arabie Pétrée, par M.
Delille [sic], est envoyée aux souscripteurs en 1833, et un
grande Carte de l’Arabie vient méme s’y ajouter en
1834.”
In most copies of the work that we have seen, the three
natural history plates are in black and white, as are the
remaining archaeological and anthropological plates. We
have seen several copies (including the scanned copy from
the Bibliotheque Nationale de France, Paris) in which
plate 63, illustrating the rock hyrax, a desert mammal
[Procavia capensis (Pallas, 1766)], is colored. In addi-
tion, of the three copies currently for sale through
AbeBooks.com (last viewed on 10 May 2019: average
price $34,182 or €29,977), all have plate 63 colored, ae
none have plate 65 colored (B. El] Hage, in litt. 15 Jan.
2019; C. Frey, in litt. 16 Jan. 2019: S. Thompson, in litt. 15
Jan. 2019): the copy recorded for sale by Maggs in 1989
was similarly listed as having “1 [plate] edlemedl by hand”
(Navari, 1989: 196-197). The catalogue of Deshayes’ own
sizable library (which was auctioned shortly after his
death) indicates that he only had Livraisons 1-5 (ie., 24
plates), but not the text or the remaining plates, which
means that he did not even have a complete set of this
work (Anonymous, 1875: 16). We have learned of only two
copies (both in Washington, D.C., at Dumbarton Oaks
and the Library of C ongress) in which plate 65, of the
mollusks, is also colored. Yaron (1979: 248) also saw a copy
with plate 65 colored, but he did not indicate which library
owned it (presumably in Europe). Given the rarity and the
Page 33
scientific value of the colored version of plate 65 for
identifying the molluscan species, we have republished it
here (Figure 1) in roughly half its original size. Some of
the plates have bilingual English-French captions, and
others have titles in preach allorives plate 65 is labeled in
both languages, “C oquilles de la Mer Rouge / Schells [sic]
from ane Rede Sea.”
The narrative and archaeological sections of the
Voyage were soon translated mee English with addi-
tional Biblical commentary, but the translation did not
include the natural history sections (Anonymous, 1836,
1838).
“Coquilles de la Mer Rouge”
The list of the Red Sea mollusks by Deshayes, essentially a
plate explanation without formal descriptions, appears on
page 66, which is the second page of feuille 33 (published
by November 1833), and the specimens were then il-
lustrated on plate 65 (published in 1833, because plate 67
has an 1833 date). Seventeen species of marine mollusks,
thirteen of them new, all from the northern Red Sea, are
listed, the majority only by citation to the illustration, i-e.,
“name and figure only >” some with brief notes about
similar species Thm few if any with sufficient information to
provide availability absent the illustrations. Plate 65 does
not have any taxonomic names printed on the plate orona
separate plate caption sheet. Pursuant to ICZN C ode
Recommendation 51E (ICZN Code, 1999), these taxa
should be cited as “Deshayes, in Laborde, 1833.” Because
the map of the expedition indicates that Laborde and
Linant traveled along both eastern and western coasts of
the Sinai Peninsula, the type locality of these species
should be “northern Red Sea’ as it cannot be determined
whether the specimens were collected in the Gulf of
Aqaba, the Gulf of Suez, or both.
The shells on Plate 65 were drawn by Paul Louis Oudart
(1796-1860), a French artist best known for his dr: awings
of birds and flowers (Bellier de la Chauvignerie, 1885:
180: Nissen, 1953: 48, 194; 1967: 238; 1978: 573. Bischoff,
2015). Oudart also did some of the illustrations for
Deshayes’ first monograph on Cenozoic fossil mollusks
from outcrops near Paris, the Descriptions des coquilles
fossils des environs de Paris (Deshayes, 1824-1837)
(Nissen, 1966: 116).
Based on review of the malacological literature over the
past century, fourteen (14) publications have used “1830”
as the date for the new species, fifty-four (54) publications
have used “1834” as the date, and we have not found
any that used 1833 as the correct date. Houart and
Trondlé (2008: 86, 88) explained their determination that
1834 should be used, but later in the same paper used
“Deshayes, 1833” in a table (2008: 93), which while
correct as to the year was unintentional. There are also
numerous per mutations of the author ship: 25 publications
attributed this to Deshayes alone; 3 to Ua alone: 10
correctly to Deshayes, in Laborde; 2 to Laborde and
Deshey es: 1 to Deshayes, in Laborde a Deshayes; and
27 to Deshay: es, in Laborde and Linant.
Page 34 THE NAUTILUS, Vol. 133, No. 2
Figure 1. Reprint of Plate 65 from Laborde (1833), reduced from the original size (15 X 22 inches, circa 43 X 59 cm). Reproduction
© Dumbarton Oaks Research Library and Collection, Rare Book Collection, Washington, D.C.
A.R. Kabat and E.V. Coan, 2019
Three malacological publications have discussed this
publication and its new taxa as a whole. The first was Lamy
(1927), who used a manuscript by Félix Pierre Jous-
seaume (1835-1921), as well as the literature on the
mollusks of the Red Sea, as the basis for determining the
then-current identity of these taxa. Lamy erred by at-
tributing the species to Laborde alone and by dating them
all to 1830.
Shortly thereafter, Tomlin and Salisbury (1928), not
having seen or cited Lamy’s paper, independently rean-
aly ei the molluscan taxa. They noted that two of the
maps “bear dates 1833 and 1834, so that the book could
not have been published prior to this last date, in spite of
the date on the title page, which is given as 1830,” and
stated that “the names should be enraged to Laborde and
Deshayes” (Tomlin and Salisbury, 1928: 32). Meanwhile,
Sherborn (1925: cxxxviii; 1932: exl) attributed the species
to “Deshayes in Laborde, 1830.”
Subsequently, Yaron (1979) compared the results of
Lamy (1927) and Tomlin and Salisbury (1928), noting that
the two publications “arrived at practically identical
conclusions with regard to the identity of most of
Laborde’s species” enon 1979: 248). Yaron erred,
however, in listing Linant as a co-author, by using
“Deshayes in ber de and Linant, 1834” for the new
species. Yaron (1979: 252) concluded that at least three,
and possibly four of the new species were still accepted as
valid, with the remaining species either junior synonyms
or junior homonyms.
Based on our review of the subsequent literature, it
appears that of the 14 “new” species, one was actually first
described by Brocchi (1814); seven are currently ac-
cepted as vi alid, five are junior synonyms (one of which is
also a junior homonym), and one is a nomen dubium.
Although this was not the first description of mollusks
from ane Red Sea—that honor belongs to Peter Forsskal
and Carsten Niebuhr who collected there in the
1760—and several other expeditions also collected
mollusks in the Red Sea in the ensuing decades (Issel,
1869: 4-12: Yaron et al., 1986), it is a useful benchmark
for the northernmost Red Sea.
Interestingly, of the twelve currently available species
(or their senior synonyms), four are endemic to the Red
Sea and the adjacent Gulf of Aden, another four are
limited to the Red Sea and the western Indian Ocean, and
the remaining four have a broader Indo-Pacific distri-
bution. This is consistent with the high degree of ende-
micity of the marine molluscan (ana of the Red Sea.
Dekker and Orlin (2000: 4) concluded that “16% of
the Red Sea species are endemics,” not including the
poorly studied Triphoridae, Eulimidae, Epitoniidae, and
Pyramidellidae. Subsequently, Dekker estimated that the
endemicity of the Red Sea molluscan fauna is in the range
of 20% to 25% (H. Dekker in litt., 11 Feb. 2019).
We provide an annotated catalog of the molluscan taxa,
including references to the secondary literature, em-
phasizing the Red Sea literature (for several widespread
species that have been extensively cited, the citations are a
representative sampling). Syntypes or potential syntypes,
festiva (Deshayes, in Laborde,
Page 35
housed in the Muséum National d’Histoire Naturelle
(Paris), are known for only two of the species.
Bivalvia
pictus, Pecten — 1833: 66, pl. 65, figs. 1-2. 2. Junior hom-
onym of Pecten pictus da Costa, 17 78. Deshayes’ species is
now known as Gloripallium maculosum (Forsskal, 1775)
(Dekker and Orlin, 2000: 10; Dijkstra and Knudsen, 1998:
61-63, pl. 6, fig. 28), endemic to the Red Sea and Gulf of
Aden fimechattbe)|
rufa, Cardita — 1833: 66, pl. 65, figs. 3-4. Cardites rufa
(Deshayes, in Laborde, 1833) (Oliver, 1992: 115-116, pl.
94: Dekker and Orlin, 2000: 12; Mienis, 2000: 131;
Rusmore-Villaume, 2008: 224-225), with a female ending
on this male genus; or more correctly Cardites rufus
(Deshayes, in TLalzondle, 1833) (Huber, 2010: 654), en-
demic to the Red Sea [Carditidae].
Gastropoda
biangulatus, Fusus — 1833: 66, pl. 65, figs. 13-14. Syno-
nym of Fusinus undulatus (Gmelin, 1791). Possible
syntypes, MNHN-IM 2000-6469; however, “The two
alleged syntypes of [the] Deshayes species ... housed in
the “MNHN collection ... are probably not conspecific
with that species” (Snyder, 2006: 107), Indo-Pacific
[Fasciolariidae].
candidula, Bulla — 1833: 66, pl. 65, fig. 9-10. Synonym of
Atys cylindricus (Helbling, 1779) (Yaron, 1979: 249), now
Aliculastrum cylindricus (Helbling, 1779) (Too et al.,
9014: 357, 362), Indo-Pacific [H aminoeidae].
depexa, Mitra — 1833: 66, pl. 65, figs. 23-25. Vexillum
(Pusia) depexum (Deshayes, in Laborde, 1833) (Cerno-
horsky, 1970: 56; Mienis, 1985; 2004; Dekker and Orlin,
2000: 30: Turner, 2001: 28: Rusmore-Villaume, 2008:
120-121), endemic to Red Sea; introduced into eastern
Mediterranean. Dekker (1993: 3) determined that the
specimen illustrated by Sharabati (1984: 66, pl. 26, fig. 3)
of this species is instead Vexillum microzonias (Lerner
1811). [Costellariidae].
fasciolaris, Mitra — 1833: 66, pl. 65, figs. 18-19. Mitra
fasciolaris Deshayes, in Laborde, 1833 Cer
1970: 35; 1976: 320-321, pl. 255, figs. 12-13, pl. 2
Sharabati, 1984: 66, pl. 26, fig. 1; Bosch et al., 1995: we:
fig. 625; Dekker and Orlin, 2000: 29: Rusmore-Villaume,
2008: 116-117), now Strigatella fasciolaris (Deshayes, in
Laborde, 1833) inedecour et al., 2018: 301), endemic to
the Red Sea [Mitridae].
festiva, Columbella — 1833: 66, pl. 65, figs. 39-41. Euplica
1833) (Drivas and Jay,
1997: 28: Dekker and Orlin, 2000: 28: Rusmore-Villaume,
2008: 102-103), Red Sea and western Indian Ocean
[Columbellidae].
Page 36
THE NAUTILUS, Vol. 133, No. 2
incarnata, Turbinella — 1833: 66, pl. 65, figs. 20-22.
Engina_ incarnata (Deshayes, in Laborde, 1833)
(Cemohorsky, 1971: 160-162, 164, figs. 80-81; 1975:
183, fig. 17); ‘Clivipollia incarnata (Deshayes, in Laborde,
1833) (Dekker and Orlin, 2000: 27: Rusmore-Villaume,
2008: 100-101: Fraussen and Stahlschmidt, 2016: 33—
34, figs. 3F-H, 6G—L), restricted to Red Sea and west-
erm Indian Ocean. Syntype, MNHN-IM 2000-30244
(figured, Fraussen and Stahlschmidt, 2016: fig. 61-])
[ Pisaniidae ].
leucozonias, Mitra — 1833: 66, pl. 65, figs. 26-28. Mitra
(Costellaria) leucozonias (Deshayes, in Lebondle. 1833)
(Lamy, 1938: 47); or Vexillum (Costellaria) leucozonias
(Deshayes, in Laborde, 1833) (Cernohorsky, 1970: 55;
Ladd, 1977: 64, pl. 20, fig. 11; Kay, 1979: 321, fig. 107-C:
Tumer, 2001: 41; Severns, 2011: 302-303, pl. 134, fig. 3):
or Vexillum leucozonias (Deshayes, in Laborde, 1833)
(Sharabati, 1984: 68, pl. 27, fig. 5; Dekker and Orlin, 2000:
30: Rusmore-Villaume, 2008: 122-123), Indo-Pacific
[|Costellariidae ].
obvelatum, Buccinum — 1833: 66, pl. 65, fig. 5—6. Synonym
of Nassarius arcularia plicatus (Réding, 1798) (Gerno-
horsky, 1984: 57-58); or Nassarius oisalains (Deshayes,
in Laborde, 1833) (Dekker and Orlin, 2000: 28), Red Sea
and Persian Gulf to South Africa | Nassariidae].
pauperculus, Fusus — 1833: 66, pl. 65, figs. 15-17. Status
uncertain: Lamy (1927: 380) noted that previous authors
had doubtfully compared this with Fusus strigatus Phil-
ippi, 1850, and F. tuberculatus Lamarck, 1822: Tomlin
and Salisbury (1928: 33) stated that this was “A Murex, but
hardly determinable”, which Snyder (2006: 158 and 275)
followed. If this species is a “Murex” then it would be a
senior secondary homonym of Murex pauperculus C. B.
Adams, 1850 (a widely cited Caribbean species now
placed in Dermomurex). However, as Deshayes’ name has
not been used as an available species, it is here regarded as
a nomen dubium.
serriale, Buccinum — 1833: 66, pl. 65, figs. 32-34. Mac-
ulotriton serriale (Deshayes, in Laborde, 1833) (Cerno-
horsky, 1972: 129-130, pl. 36, fig. 11; Salvat and Rives,
1975: 317, fig. 217; Cemohorsky, 1982: 130-134, figs.
22-30; Tr éndlé and Houart, 1992: 90-91, figs. 51-56;
Wilson, 1994: 23; Bosch et al., 1995: 121, fig. 482: Houart,
1995: 263-264, fig. 87; Higo et al. 1999: 209: Dekker and
Orlin, 2000: 27; Houart, 2008: 200, pl. 395, fig. 1; Houart
and Trondlé, 2008: 86, 88: Rusmore- Willlanrme, 2008:
92-93; Spencer et al., 2009: 210; Tréndlé and Boutet,
2009: 28; Houart et al., 2010: 265; Severns, 2011:
280-281, pl. 123, fig. 8; Claremont et al., 2013: 21; Tan
and Low, 2014: 354; Okutani, 2017: 956, pl. 249, fig. 8)
(and numerous other publications), Indo-Pacific. Several
authors have erroneously emended the species name to
“serrialis” (e.g., Satyamurti, 1952: 161-162; Asakura et al.,
1993: 12; Apte, 1999: 346; Lee and Chao, 2003: 34, 42,
pl. 4, fig. 21; Smith, 2003: 260). Cernohorsky (1982: 132,
fig. 22) designated Deshayes’ illustration as the lectotype
of this species [Muricidae].
teniata, Turbinella — 1833: 66, pl. 65, fig. 7-8. Synonym of
Latirus turritus (Gmelin, 1791) G@roralin and. Salisbury,
1928: 33; Snyder, 2003: 204), now Turrilatirus turritus
(Gmelin, 1791) (Vermeij and Snyder, 2006: 419), Red Sea
and western Indian Ocean [Fasciolariidae].
vermicularis, Turritella “nob.” — Lamy (1927: 380) at-
tributed this species to “Laborde” (i.e., Deshayes, in
Laborde, 1833: 66, pl. 65, figs. 11-12), but Deshayes cited
Turbo vermicularis Brocchi (1814: pl. 6, fig. 13), ‘which is
now Helminthia vermicularis (Brocchi, 1814) (Landau
et al., 2013: 62-64, pl. 5, fig. 11), so Deshayes’ usage was a
new combination, not a new species. Deshayes stated that
this material was the Recent analog of Brocchi’s fossil
species [“lanalogue vivant de la coquille fossile figurée par
Brocchi’ |, so that Deshayes’ usage of “Nob.” leobas or
new] for this species would now be seen as inappropriate,
but he and others of his time often did this on the transfer
of an earlier author's species to a different genus. How-
ever, Brocchi’s species is limited to the iBevaby Miocene to
Early Pleistocene of the Mediterranean, and is not known
from the Recent (Landau et al., 2013: 64), so Deshayes’
material may be referrable to another species of (Recent)
Turritellidae from the Red Sea [Turritellidae].
“Flore de VArabie pétrée™
Alire Raffeneau-Delile (1778-1850) authored the bo-
tanical section of the Voyage, titled “Flore de lArabie
pétrée,” printed on pages 81-87 and accompanied by
plate 64. Delile, a professor at Montpellier, was best
known for his publications of the botany of Egypt, issued
as part of the Description de Egypte in two editions from
1812 to 1829 (Motte, 1971; Rioux, 2014: 334-336; Stafleu
and Cowan, 1976: 617). Delile was thus a logical choice to
describe the plants collected by Laborde and Linant, and
his text included a description of 85 species, of which eight
are new. Plate 64 has illustrations of six of the new species.
However, Delile must have realized that the folio size of
the Voyage and its focus on geography and archaeology
would limit the distribution of | his ‘botanical chapter, as Hie
had the entire botanical section reprinted and re-typeset
in a smaller quarto size, titled “Fragments dune Flore de
VArabie Pétrée: plantes recueillies par M. Léon de
Laborde” (Delile, 1833).
The botanical literature has used either 1830 or 1833 as
the date for Delile’s section. Pritzel (1872: 79) correctly
dated the “Fragments” version to 1833, but incorrectly
dated the original Laborde version as 1830. This error was
repeated by Stafleu and Cowan (1976: 617) who wrote:
“The original text, in folio, appeared in 1830, p. 81-87
(fide PR), : citing Pritzel. Jackson (1881: 379) dated the
“Fragments” version to 1833, but made no mention of the
original Laborde version. Rioux (2014: 338) dated
the Laborde version as 1833 and illustrated the title page
of the “Fragments” version, but did not further discuss
A.R. Kabat and E.V. Coan, 2019
Page 37
either version. Embacher (1882: 180), an encyclopedia of
travel books, dated the entire Voyage as 1830-33, and the
“Flore de lArabie pétrée” to 1833; the latter being
Delile’s reprint. Joly (1859: 95-96) correctly dated ine
“Fragments” to 1833, and noted that there was an un-
published second part, “Nouveaux fragments d'une Flore
de V’Arabie Pétrée: plantes recueillies aux environs du mont
Sinai par M. le baron Taylor’, prepared in April 1834 to
be presented to the Académie des Sciences, Toulouse. As
noted above, Augé and de Bellefonds (1994: 20-21)
explained that hel botany section of the Voyage was sent
to subscribers in 1833, which should be used as the
publication date for both the Voyage version and the
“Fragments” version. The botanical section thus confirms
the 1833 publication date for the molluscan taxa.
ACKNOWLEDGMENTS
One of us (Kabat) thanks the librarians who allowed him
to examine the set(s) of the Voyage at their institutions:
British Library; Houghton Library, Harvard University;
Library of Congress (Michael North and Marianna Stell);
and New Youk: Public Library. Anatole Tchikine (Dum-
barton Oaks Research Library and Collection, Harvard
University) kindly provided a scan of the color plate of
mollusks in the Dumbarton Oaks copy, which is reproduced
here by permission of the Dumbarton Oaks Research Li-
brary and Collection, Rare Book Collection, Washington,
D.C. We thank the antiquarian book dealers who pr ovided
information about the copies of the Voyage in their stock:
Badr El Hage (Folios Ltd., London); Christopher Frey
(Antiquariat INLIBRIS, Vienna); and Seth Thompson
(Sequitur Books, Boonsboro, Maryland). Henk Dekker
provided helpful information on the endemicity of Red Sea
marine mollusks. Leslie Overstreet (Natural History Rare
Books, Smithsonian Libraries), Riidiger Bieler (Field Mu-
seum) and Paula Mikkelsen (Paleontological Research In-
stitution) provided helpful reviews of the manuscript.
LITERATURE CITED
André, M. 1943. Edouard Lamy. Abbeville: F. Paillart, 21 pp.
Anonymous. 1830, 1833. Livres Frangais. Bibliographie de la
France, ou Journal Général de Imprimerie et de la Librairie,
19(4): 81-92: 19(10): 145-158; 19(16): 241-252: 19(21):
321-334: 19(29): 481-490: 19(38): 617-626 (1830): 22(32):
489-500; 22(48): 745-758 (1833).
Anonymous. 1834. Voyages [review of Laborde]. Bibliotheque
Universelle des Sciences, Belles-Lettres et Arts, 19(2),
Littérature, 56: 150-164.
Anonymous, ed. 1836. Journey through Arabia Petraea, to
Mount Sinai and the Excavated City of Petra, the Edom of
the Prophecies. London: John Murray, xxviii + 331 pp.,
26 pls. [partial translation and revision of Laborde,
1830-1834].
Anonymous, ed. 1838. Journey through Arabia Petraea, to
Mount Sinai and the Excavated City of Petra, the Edom of
the Prophecies (2"" ed.). London: John Murray, xxviii + 340
pp.. 26 pls. [revised from Anonymous (1836): partial
translation and revision of Laborde, 1830-1834].
Anonymous. 1875. Catalogue de livres histoire naturelle et
particulieérement de géologie et de conchyliologique com-
posant la bibliothéque de M. G. P. Deshayes. Paris: J.-B.
Bailliére et Fils, xii + 160 pp.
Apte, D. 1999. Maculotriton serrialis (Deshayes in Laborde and
Linnet [sic] 1834) from Okha, Gulf of Kutch: a new ere
Journal of the Bombay Natural History Society 96(2): 346.
Asakura, A., S. Nishihama, and Y. Kondo. 1993. ates on the
biology and ecology of the intertidal animals of Chichijima
Tela in the Ogasawar a (Bonin) Islands. 1. List of collected
species with comments on some species. Atoll Research
Bulletin 383: 1-17.
Augé, C. and P. Linant de Bellefonds. 1990. Deux Frangais a
Petra: Le Voyage de Laborde et Linant de Bellefonds en
1828. Archeologia 262: 48-59.
Augé, C. and P. Linant de Bellefonds. 1994. Pétra retrouvée:
Voyage de l’Arabie Pétrée, 1828. Paris: Pygmalion, 285 pp.
[reprinted 1998].
Bellier de la Chavignerie, E. (continued by L. Auvray). 1882-1887.
Dictionnaire général des artistes de l'école frangaise. Paris:
Renouard, 2 vols. and supplement [reprinted 1979].
Bischoff, W. 2015. Bemerkungen zum Umschlagbild: Paul Louis
Oudart (1796-1860) und seine “Planche unique” —
Anmerkungen zum Autorennamen von Lacerta galloti. Die
Eidechse (Magdeburg / Hamburg), 26 (Suppl.): 27-30.
Bosch, D.T., S.P. iDEN, R.G. Moolenbeek, and P.G. Oliver.
1995. Seashells of eastern Arabia. Dubai: Motivate Pub-
lishing, 296 pp.
Brocchi, G. 1814. Conchiologia fossile subapennina, con
osservazioni geologiche sugli Apennini e sul suolo adia-
cente. Milano: Stamperia Reale, 2 vols. [1-240 (vol. 1);
941-712 (vol. 2), 16 pls.].
Cemohorsky, W.O. 1970. Systematics of the families Mitridae &
Volutomitridae (Mollusca: Gastropoda). Bulletin of the Auck-
land Institute and Museum 8: iv + 1-190.
Cemohorsky, W.O. 1971. Indo-Pacific Pisaniinae (Mollusca:
Gastropoda) and related buccinid genera. Records of the
Auckland Institute and Museum 8: 137—167.
Cernohorsky, W.O. 1972. Marine Shells of the Pacific (vol. II).
Sydney: Pacific Publications, 411 pp.
Cernohorsky, W.O. 1975. Supplementary notes on the taxonomy
of buccinid species of the subfamily Pisaniinae (Mollusca:
Gastropoda). Records of the Auckland Institute and Mu-
seum 12: 175-211.
Cernohorsky, W.O. 1976. The Mitridae of the world, Part I. The
subfamily Mitrinae. Indo-Pacific Mollusca 3(17): 273-528.
Cernohorsky, W.O. 1982. The taxonomy of some Indo-Pacific
Mollusca. Part 10. Records of the Auckland Institute and
Museum 19: 125-147.
Cemohorsky, W.O. 1984. Systematics of the family Nassariidae
(Mollusca: Gastropoda). Bulletin of the Auckland Institute
and Museum 14: 1-356.
Claremont, M., R. Houart, $.T. Williams, and D.G. Reid. 2013.
A molecular phylogenetic framework for the Erga-
lataxinae (Neogastropoda: Muricidae). Journal of
Molluscan Studies 79(1): 19-29.
Dekker, H. 1993. Werklijst met correcties en aanvullingen op D.
Sharabati, “Red Sea Shells,” 1984. Winkel (Netherlands),
priv. publ., 5 pp.
Dekker, H. and Z. Orlin. 2000. Check-list of Red Sea Mollusca.
Spirula 47 (Supplement): 1-46.
Delile, A. 1833. Fragments dune Flore de lArabie Pétrée:
plantes recueillies par M. Léon de Laborde. Paris: Giard, 26
pp. 1 pl.
Page 38
Dijkstra, H.H. and J. Knudsen. 1998. Some Pectinoidea
(Mollusca: Bivalvia: Propeamussiidae, Pectinidae) of the
Red Sea. Molluscan Research 19(2): 43-104.
Drivas, J. and M. Jay. 1997. On a collection of Columbellidae
from the Red Sea. Apex 12(1): 27-30.
Embacher, F. 1882. Lexikon der Reisen und Entdeckungen.
Leipzig: Verlag des Bibliographischen Instituts, viii + 394 pp.
[reprinted (1988), Pforzheim: Antiquariat P. Kiefer].
Fedosov, A., N. Puillandre, M. Herrmann, Yu. Kantor, M.
Oliverio, P. Dgebuadze, M.V. Modica, and P. Bouchet.
2018. The collapse of Mitra: molecular systematics and
morphology of the Mitridae (Gastropoda: Neogastropoda).
Zoological Journal of the Linnean Society 183(2): 253-337
Fischer, P.-H. and E. Fischer. 1945. Nécrologie. Edouard Lamy
1866-1942. Journal de Conchyliologie 86(1): 6-28.
Fraussen, K. and P. Stahlschmidt. 2016. Revision of the Clivi-
pollia group (Gastropoda: Buccinidae: Pisaniinae) with
description of two new genera and three new species.
Novapex 17(2—3): 29-46.
Goby, A. 1981. Louis Linant (Pacha de Bellefonds) (1798-1883).
Publications de Académie des Sciences d’Outre-Mer:
Travaux et Mémoires: Hommes et Destins (Dictionnaire
biographique d’Outre-Mer) 4: 470-475.
Higo, S., P. Callomon, and Y. Goto. 1999. Catalogue and bib-
liography of the marine shell-bearing Mollusca of Japan.
Osaka: Elle Scientific Publications, 749 pp.
Houart, R. 1995 [“1994"]. The Ergalataxinae (Gastropoda,
Muricidae) from the New Caledonia region with some
comments on the subfamily and the description of thirteen
new species from the Indo-West Pacific. Bulletin du
Muséum National d'Histoire Naturelle, Paris (ser. 4)
(section A, Zoologie, Biologie et Ecologie Animales)
16(2-4): 245-297.
Houart, R. 2008. Muricidae. Pp. 132-221, in (G. T. Poppe, ed.),
Philippine marine mollusks, Volume II (Gastropoda — Part
2). Hackenheim: ConchBooks, $48 pp.
Houart, R. and J. Tréndlé. 2008. Update of Muricidae (excluding
Coralliophilinae) from French Polynesia with description of
ten new species. Novapex 9(2—3): 53-93.
Houart, R., R.N. Kilburn, and A.P. Marais. 2010. Muricidae. Pp.
176-270, in A. P. Marais and A. D. Seccombe, eds.,
Identification guide to the seashells of South Africa: Volume
1. Groenkloof: Centre for Molluscan Studies, 376 pp.
Huber, M. 2010. Compendium of Bivalves. Hackenheim:
ConchBooks, 901 pp. + CD with searchable version of
chapter 6 and Excel file with additional synonymies.
ICZN. 1999. International Code of Zoological Nomenclature
(4th ed.). London: International Trust for Zoological No-
menclature, xxix + 306 pp:
Issel, A. 1869. Malacologia del Mar Rosso: ricerche zoologiche e
paleontologiche. Pisa: Biblioteca Malacologica, 387 pp., 5
ds.
reson B.D. 1881. Guide to the literature of botany: being a
classified selection of botanical works, including nearly 6000
titles not given in Pritzel’s Thesaurus. London: Longmans,
Green (Index Society, Publications, vol. 8), xl + 626 pp.
[reprinted, New York: Hafner (1964)].
Joly, N. 1859. Eloge historique d’Alyre Raffeneau Delile, Pro-
fesseur de Botanique a la Faculté de Médecine de
Montpellier. Mémoires de Académie Impériale des Sci-
ences, Inscriptions et Belles-Lettres de Toulouse (ser. 5) 3:
63-98.
Kay, E.A. 1979. Hawaiian marine shells. Honolulu: Bishop
Museum Press, xviii + 653 pp.
THE NAUTILUS, Vol. 133, No. 2
Kormrumpf, H.-J. and J. Kormrumpf. 1998. Fremde im Osma-
nischen Reich 1826-1912/13: Bio-bibliographisches Regis-
ter. Stutensee: priv. publ., x + 445 pp.
Kurz, M. and P. Linant de Bellefonds. 1998. Linant de Belle-
fonds: Travels in Egypt, Sudan and Arabia Petraea,
1818-1828. Pp. 61-69, in (P. Starkey and J. Starkey, eds.),
Travellers in Egypt. London and New York: I. B. Tauris,
318 pp.
Labib, M. 1961. Pélerins et voyageurs au Mont Sinai.
Recherches d’archéologie, de philologie et dhistoire 25.
Caire: Institut Francais d’Archéologie Orientale, vi + 180
op., 24 pls.
Laborde, L. de. 1830-1834. Voyage de l’Arabie Pétrée, par Léon
de Laborde et Linant. Paris: Giard, [iv] + 87 + [ii] pp.
69 pls., online at: http://bibliotheque-numerique.inha.fr/
viewer/16377/?offset=#page = 1 &viewer=picture
Ladd, H.S. 1977. Cenozoic fossil mollusks from western Pacific
islands; gastropods (Eratoidae through Harpidae). U. S.
Geological Survey Professional Paper 533: iv + 84 pp., 23
ols.
ay E. 1927 (27 January) [“1926"]. Sur diverses coquilles de la
mer fares figurées en 1830 par Léon de Laborde. Bulletin
du Muséum Reto d Histoire Naturelle (ser. 1) 32(6):
375-383. [Published in 1927, although dated 1926 (André,
1943: 14; Fischer and Fischer, 1945: 17; P. Bouchet, in litt. 22
Aug. 1991)].
Lamy, E. 1938. Mission Robert Ph. Dollfus en Egypte: VIL,
Mollusca Testacea. Mémoires Preséntés a_ l|Institut
d' Egypte 37: 1-90, 1 pl.
Landau, B.M., M. Harzhauser, Y. Islamoglu, and C.M. da Silva.
2013. Systematics and palaeobiogeography of the gastro-
pods of the middle Miocene (Serravallian) Karaman ‘Basin,
Turkey. Cainozoic Research 11-13: 3-584.
Lee, S.-C. and S.-M. Chao. 2003. Shallow-water marine shells
from northeastern Taiwan. Collection and Research (Tai-
wan) 16: 29-59.
Mazuel, J. 1937. L’ouevre Bees aphique de Linant de Belle-
fonds: étude de géogr: apl rie historique. Cairo: Société
Royale de Géogr aphie ic Egypte, xxiv + 400 + xxiii pp., 6
maps, 9 pls.
Mienis, H.K. 1985. First record of the Erythraean species Vex-
illum depexum from the Mediterranean coast of Israel.
Levantina: a Journal of Malacology 58: 640.
Mienis, H.K. 2000. Some notes concerning Carditidae from the
Red Sea and a case of blister pearl Sareviton in Cardites
rufa. Of Sea and Shore 23(3): 131.
Mienis, H.K. 2004. Mariene mollusken uit het oostelijk deel van
de Middellandse Zee 22: Een tweede vondst van Vexillum
(Pusia) depexum in Israel. Spirula, Correspondentieblad
van de Nederlandse Malacologische Vereniging 339: 76-77.
Motte, J. 1971. Delile (or Raffeneau-Delile), Alire. Dictionary of
Scientific Biography 4: 21-22.
Navari, L. 1989. Greece and the Levant: the catalogue of the Henry
Myron Blackmer collection of books and manuscripts. Lon-
don: Maggs Bros. Ltd., xxiv + 446 pp., 16 pls.
Nissen, C. 1953. Die Illustrierten Vogelbiicher: ihre Geschichte
und Bibliographie. Stuttgart: Hiersemann, 222 pp.
[reprinted 1976].
Nissen, C. 1966-1969. Die Zoologische Buchillustration:
ihre Bibliographie und Geschichte. Band I: Bibliographie.
Stuttgart: Anton Hiersemann, 666 pp. [1(1): 1-80 (1966);
1(2): 81-160 (1966): 1(3): 161-240 (1967); 1(4):
241-320 (1967): 1(5): 321-400 (1968): 1(6): 401-480
(1969): 1(7): 481-560 (1969): 1(8): 561-666 (1969)].
A.R. Kabat and E.V. Coan, 2019
Nissen, C. 1971-1978. Die Zoologische Buchillustration: ihre
Bibliographie und Geschichte. Band IL: Geschichte der
Zoologischen Buchillustration. Stuttgart: Anton Hier-
semann, xvi + 604 pp., 48 pls. [2(9): 1-72, pls. 1-8 (1971);
2(10): 73-144, pls. 9-16 (1972); 2(11): 145-216, pls. 17-24
(1973); 2(12): 217-288, pls. 925-32 (1974): 2(13): 289-360,
pls. 33-40 (1975); 2(14): 8361-432, pls. 41-48 (1976): 2(15):
433-512 (1978): 2(16): 513-604 + xvi (1978)].
Okutani, T., ed. 2017. Nihon kinkaisan kairui zukan / Marine
Mollusks in Japan, 2°¢ ed. Kanagawa-ken Hiratsuka-shi:
Tokai Daigaku Shuppankai, 1379 pp., incl. 567 pls.
Oliver, P.G. 1992. Bivalved seashells of the Red Sea. Cardiff:
National Museum of Wales & Wiesbaden: Hemmen, 330
pp.. incl. 46 pls.
Pfullmann, U. 2001. Durch Wiiste und Steppe: Entdeck-
erlexikon arabische Halbinsel; Biographien und Berichte.
Berlin: Trafo, 560 pp.
Pritzel, G.A. 1871-1877. Thesaurus literaturae botanicae omnium
gentium, inde a rerum botanicarum initiis ad nostra usque
tempora, quindecim millia operum recensens. Leipzig: F.A.
Brockhaus, ii + 577 pp. [reprinted, Milano: Gorlich (1950);
Koenigstein: Otto Koeltz Antiquariat (1972)] [pp. 1-80
(1871-72); pp. 81-320 (1872); pp. 321-577 (1877), fide
Stafleu (1972: 126)].
Rioux, J.-A. 2014 [“2013”]. Alire Raffeneau-Delile (1778-1850),
de la campagne @’ Egypte a la direction du Jardin des plantes
de Montpellier. Bulletin de !Académie des Sciences et
Lettres de Montpellier (n.s.) 44: 331-340.
Rusmore-Villaume, M. L. 2008. Seashells of the Egyptian Red
Sea: the illustrated handbook. Cairo: American University
in Cairo Press, xii + 307 pp.
Salvat, B. and C. Rives. 1975. Coquillages de Polynésie. Papeete:
Editions du Pacifique, 391 pp.
Satyamurti, $.T. 1952. The Mollusca of Krusadai Island (in the
Gulf of Mannar), I. Amphineura and Gastropoda. Bulletin
of the Madras Government Museum, Natural History (n.s.)
1(2)[6]: 1-267, 34 pls.
Severns, M. 2011. Shells of the Hawaiian Islands: the sea shells.
Hackenheim: ConchBooks, 564 pp., incl. 225 + [11] pls.
Sharabati, D. 1984. Red Sea shells. London: KPI Limited, 128
op.
Shatsann C.D. 1922-1933. Index Animalium sive index
nominum quae ab A.D. MDCCLVIII generibus et
speciebus animalium imposita sunt. Sectio Secunda. A
kalendis Ianuariis, MDCCCI usque ad finem Decembris,
MDCCCL. London: British Museum (Natural History).
[Issued in 33 parts: 1: [i]-cxxxii, 1-128 (1922); 2: 129-384,
exxxiii-cxxxvi (1923): 3: 385-640 (1923): 4: 641-943 (1924):
5: 945-1196 (1924): 6: 1197-1452 (1925): 7: 1453-
1771, [exxxvii|-exxxix (1925); 8: 1773-2008 (1925); 9:
2009-2248 (1926): 10: 2249-2568 (1926): 11: 2569-2880
(1926): 12: 2881-3136 (1927); 13: 3137-3392 (1927); 14:
3393-3746 (1927): 15: 3747-3970 (1928): 16: 3971-4194
(1928): 17: 4195-4450 (1928): 18: 4451-4690 (1929): 19:
4691-4930 (1929): 20: 4931-5138 (1929); 21: 5139-5348
(1929): 22: 5349-5702 (1930); 23: 5703-5910 (1930); 24
5911-6118 (1930); 25: 6119-6358 (1931); 26: 6359-6582
‘ a 2C
Page 39
(1931); 27: 6583-6806 (1931); 28: 6807-7056 (1932); 29:
lil-vii, exxxiii-cxlviii, 1-208 (1932); 30: 209-416 ( eee 31:
417-654 (1932): 32: 655-878 (1933): 33: 879-1098 (1933)].
Smith, B.D. 2003. Prosobranch gastropods of Guam. Micro-
nesica 35-36: 244-270.
Snyder, M.A. 2003. Catalogue of the marine gastropod family
Fasciolariidae. Academy of Natural Sciences of Phila-
delphia, Special Publication 21: iv + 1-431.
Snyder, M.A. 2006. A new species of Fusinus ( sastropoda:
Fasciolariidae) from the Red Sea and the identity of Fusinus
undulatus (Gmelin, 1791). Gloria Maris 45(5): 104-114.
Spencer, H.G., B.A. Marshall, and R.C. Willan. 2009. Checklist
of New Anat living Mollusca. Pp. 196-219, in (D. P.
Gordon, ed.) New Ze: sitar inventory of biodiversity. Volume
One. Kingdom Animalia: Radiata, Lophotrochozoa, Deuter-
ostomia. Cirasidnndye Canterbury University Press, 566 pp..
16 pls.
Stafleu, F.A. 1973. Pritzel and his Thesaurus. Taxon 22(1):
119-126.
Stafleu, F.A. and R.S. Cowan. 1976. Taxonomic Literature: a
selective guide to botanical publications and collections
with dates, commentaries and types (2°4 ed.). Volume 1:
A-G. Utrecht: Bohn, Scheltema & Holkema, xl + 1,136 pp.
Tan, S.K. and M.E.Y. Low. 2014. Checklist of the Mollusca of
Cocos (Keeling) / Christmas Island ecoregion. Raffles
Bulletin of Zoology. Supplement 30: 313— 375.
Tomlin, J.R. le B. and A.E. Salisbury. 1928. Laborde’s “Voyage”
and the Mollusca therein described by Deshayes. Pro-
Soci of the Malacological Society of London 18(1):
B95) ‘pl. il,
Too, C.C., C. Carlson, P.J. Hoff, and M.A. E. Malaquias. 2014.
Diversity and systematics of Haminoeidae gastropods
(Heterobr. anchia: Cephalaspidea) i in the tropical West Pacific
Ocean: new data on the genera Aliculastrum, Atys, Diniatys
and Liloa. Zootaxa 3794(3): 355-392.
Trondlé, J. and M. Boutet. 2009. Inventory of marine molluscs of
French Polynesia. Atoll Research Bulletin 570: 1-90.
Trondlé, J. and R. Houart. 1992. Les Muricidae de Polynésie
Francaise. Apex 7(3-4): 67-149.
Tumer, H. 2001. Katalog der Familie Costellariidae MacDonald
1860 (Gastropoda: Prosobranchia: Muricoidea). Hack-
Paes ConchBooks, 100 pp.
Vermeij, G.J. and M.A. Snyder, 2006. Shell characters and
taxonomy of Latirus and related fasciolariid groups. Journal
of Molluscan Studies 72(4): 413-424.
Vicaire, G. 1900 [in 1894-1920]. Manuel de lamateur de livres du
XIX siécle, 1801-1893 [vol. 4]. Paris: A. Rouquette, 8 vols.
Wilson B.R. 1994. Australian Marine Shells: Prosobranch
Gastropods, Part Two (Neogastropods). Kallaroo: Odyssey
Publishing, 370 pp.
Yaron, I. 1979. The Red Sea Mollusca collected by Laborde in
1828. Levantina: a Malacological Newsletter 22: 248-253.
Yaron, I., T. Schigtte, and G. Wium-Andersen. 1986. A review of
molluscan taxa described by P. Forsskal and C. Niebuhr,
with citation of original descriptions, discussion of type-
material available and selection of some _ lectotypes.
Steenstrupia 12(10): 157-203.
THE NAUTILUS 133(2):40-47, 2019
Page 40
Crepidula fornicata (Linnaeus, 1758) (Gastropoda: Calyptraeidae)
as a hermit crab commensal in the North Sea
J.G.M. Raven
Naturalis Biodiversity Center
Leiden, The Netherlands
han.raven@naturalis.nl
ABSTRACT
The association of Crepidula fornicata (Linnaeus, 1758) with
hermit crabs is discussed, based on recent observations from the
coast of Zuid-Holland (The Netherlands). In the Netherlands,
the species is a generalist that occupies a wide range of sub-
stratum types, ee at the investigated localities it is most fre >quent
on hermit-crab occupied gastropod shells, in particular those of
Euspira catena (Da Costa, 1778). For the first time a number of
specimens is reported that has C. fornicata not only on the
outside, but also on the inside of the aperture, to which it adapts
its shell shape.
INTRODUCTION
In the late 19" century, Crepidula fornicata (Linnaeus, 1758)
was introduced, via oysters, from the East coast of North
America to the United Kingdom, from where it spread
throughout the North Sea ‘(living specimens were first
recor ad from the Netherlands in 1929: Wolff, 2005). In the
Netherlands it became abundant in the delta and in the
Wadden Sea, where it mainly grows on oysters and mussels
(De Bruyne et al., 2013). Thieltges et al. (2002) studied a part
of the German Wadden Sea where they made the same
observation, but also found that in subtidal areas it relies on
other substrates, mainly provided by Buccinum undatum
(Linnaeus, 1758). In that area, about 70% of C. fornicata
occur on B.undatum shells occupied by hermit crabs, 14% on
dead B.undatum, 2% on living B. undatum, and the remaining
14% on crabs (Cancer pagurus Linnaeus, 1758). In its aativs
range, Karlson and Shenk (1983: table 3) even found a greater
preference for shells occupied by hermit crabs (compared to
empty shells). The preference to settle on shells occupied by
hermit crabs is explained by the longer time to grow and
develop as larger species of hermit crabs prevent their shells
from being covered by sediment or buried in the substratum,
provide aeneawell food supply due to mobility of the crab and
water currents set up by its feeding, and provide protection
from predators (Conover, 1975, 1979).
Along the coast of the province Zuid Holland, an area
with a camnly sea bed that does not provide a suitable substrate
for Crepidula fornicata, only small numbers of beached
specimens were observed until recently. These were always
attached to shells (own observations since mid-1970s). After
storms in the winters of 2017 and 2019, large numbers of
marine organisms washed up (e.g., Raven, 2017) including the
bivalve Mytilus edulis (Linnaeus, 1758), of which dozens with
C, fornicata and hundreds of specimens of the gastropod
Euspira catena (Da Costa, 1778). The latter were partly alive,
partly fresh-dead shells occupied by hermit crabs and with
Crepidula fornicata. Previously, only old (often blue-tinted)
Holocene specimens of this gastropod washed up, which
indicates that (like several other species) it now lives closer to
the coast (Raven, 2017). Their typical collar-shaped egg
capsules also frequently wash up on the same beaches (onan
observations). As Euspira catena lives buried in the sand, the
smooth shells of the living specimens have no epibionts. Upon
death of the mollusk, the shell quickly becomes covered by all
kinds of organisms, especially when a hermit crab (Pagurus
bernha ig (Linnaeus, 1758)) occupies the shell: hese are
mostly barnacles, sea mat Hydractinia echinata (Fleming,
1828), and the subject of this paper: C. fornicata.
In the Netherlands, Crepidula fornicata has been ob-
served to occupy a wide range of substratum types: on large
stones, on living bivalves (especially the Mussel Mytilus
edulis (Linnaeus, 1758), and the Pacific oyster Magellana
gigas (Thonberg, 1793)), inside empty biv: alvesh (e.g.,
Acanthocardia echinata (Linnaeus, 1758), Figure 4), on
shells inhabited by hermit crabs, etc. It thus is a faculta-
tive commensal of hermit crabs, living on the outside of
the shell (also noted by Conover, 1976; Williams and
McDermott, 2004). This agrees to the general behavior
reported for this species (Anonymous, 2019). Vermeij
(1989) reported Grandicrepidula ¢ grandis (Middendorff,
1949), which occupies the full range of substratum types
that in other regions are occupied iby three or four more
specialized calyptraeids, including the interior surfaces of
gastropod shells occupied by her mit crabs.
The compelling reason that drove me to embark on this
research is that several specimens not only had Crepidula
fornicata on the outside, but also inside the aperture. Only
a single unpublished record of this behavior has been
found, although the literature about hermit crabs and
their commensals is vast and other observations could be
buried in a more generic paper on this subject.
].G.M. Raven, 2019
MATERIALS AND METHODS
This work is based on observations by the author at various
localities in the Netherlands over a period of more than 40
years, but key input is from material that washed up along
the coast of Zuid-Holland after storms during the winters of
2017 and 2019. During each visit to Scheveningen and
Hoek van Holland (Figure 1) about 3 km of shor kine were
inspected for 2 hours. All gastropods with Crepidula for-
nicata inside the shell were collected, as well as some
specimens with C. fornicata only on the outside. Numbers
of shells observed were estimated. The collected shells
were photographed, cleaned and re- -assembled. All material
is kept in the author's collection. Abbreviations: L = length,
H = height, W = width. These were measured using a
digital caliper. For shell height, the protruding septum
oO
(Figures llc, 17c) was included in the measurement.
RESULTS AND DISCUSSION
All shells inhabited by hermit crabs are Euspira catena,
in addition to a single, gerontic Buccinwm undatum
Linnaeus, 1758. Both species have large shells (compared
to others in the local fauna) with wide apertures. Several
shells were found with Crepidula but no hermit crab; the
crab may have died before, have washed out during
tr ansport to the beach, or abandoned the shell. Hermit
crabs often change shell as they grow or if epibionts make
the shells too heavy or unstable e (e.g. Conover, 1979).
Men. in
{ Ouddorps
me
Figure 1.
Maps.
Location map. 1. Hoek van Holland. 2
. Northem Beach,
Page 41
All shells found with hermit crabs had C. fornicata on
them, which indicates how successful its larvae are in
finding shells occupied by hermit crabs. Crepidula for-
nicata grows fast: two months after settling they can be
4mm long and sexually mature as females (Walne, 1956).
This could be an advantage as it allows the animal to grow
and reproduce before the hermit crab abandons the shell
(Conover, 1979). Once a single C. fornicata is present,
more will follow; either they settle on the first specimen or
next to it. As more specimens settle, the Crepidula for-
nicata form stacks (see Figure 3 for rather long stacks). In
such stacks, the oldest specimens are female, ne youngest
are male, those in between are protandrous hermaph-
rodites; as the stack grows specimens will change sex
(e.g., Fretter and Graham, 1981; Collin, 2006).
As the oldest specimen in a stack dies it becomes dis-
connected from its substratum but the stack will survive.
Frequently such stacks are found with an overgrown and/or
eroded specimen at its base. As the individual Cr epidula
fornicata and the stacks grow, their weight or weight dis-
tribution can become a serious burden “or the Nett (e.g.
Figure 3) and the crab may abandon the shell (Conover.
1978). In several of the shells with larger c Wea no
hermit crab was present (e.g., Figures 3, 7, 10), but it is
assumed the stacks started on a hermit crab- inetteed shell.
On a few female shells two or three males were present
(e.g., Figures, 7, 8). Only in one case a female had multiple
sealsonn. ©) epidula fornicata observed left a clear mark on
the host shell. Some host shells were found with such marks
indicating that a C. fornicata (or stack of such shells) had
Womuideny ye
IAN
Spiikehisse
Scheveningen. 3. Wassenaarse Slag, Wassenaar. Source: Google
Page 42 THE NAUTILUS, Vol. 133, No. 2
Figures 2-10. Hermit crab-inhabited shells with Crepidula fornicata. 2. Euspira catena (H ~35 mm) with hermit crab and with C.
fornicata on outside, Scheveningen, Jan. 2017. 3. Euspira catena with two long and complex stacks of Crepidula fornicata on outside,
some with sea mat or barnacles, Wassenaarse Slag, Wassenaar, March 2019 (total W ~10 cm). 4. Acanthocardia echinata (1. = 40 mm)
with two stacks and three individual Crepidula fornicata inside, Hoek van Holland, Feb. 2019. Note the “flatness” of the female in the
valve on the right. 5-6. Euspira catena with C. fornicata on outside and inside aperture, Scheveningen, Jan. 2017. 5. Euspira catena (H ~
35 mm) (Table 2: 1). 6. Euspira catena (H = 33 mm) (Table 2: 3). 7. Euspira catena (H = 38 mm) with C. fornicata stacks on outside and
inside the aperture, Hoek van Holland, Feb. 2017 (Table 2: 4). 8. Euspira catena (H = 35 mm) with several stacks of Crepidula fornicata
on the outside and another stack inside the aperture, Hoek van Holland, Feb. 2019 (Table 2: 9). 9. Euspira catena (H = 38 mm) with C.
fornicata stacks on outside and inside the aperture, also in adapical position, Hoek van Holland, Feb. 2019 (Table 2: 11). 10. Buccinum
undatum (H = 62 mm) with large Crepidula fornicata on outside and inside the aperture, also in the adapical position, Hoek van
Holland, Feb. 2019 (Table 2: 14). Note the profuse presence of boring sponges, sea mat, barnacles and bryozoans that died some time
ago.
J.G.M. Raven, 2019
Page 43
been present but dropped off (e.g. Figure 16). As the
specimens studied were transported during a stormn it is likely
the C. fornicata were lost as result of the transport to shore,
rather than by death in deeper water. The marks are in-
formative in the re-assembly of host shells and their com-
mensals (in addition to the photos taken before cleaning), but
the marks appear not to be preserv red in fossils (Walker,
1992). Crepidula fornicata fit precisely onto the host shell or
underlying C. fornicata, but in stacks with overlapping shells
a narrow slit may be left open. It was noted that the com-
mensals are generally (not always) oriented with their head
toward the aperture of the host shell: this suggests that they
could benefit most from the water currents produced by the
host and are closest to their food leftovers.
In January 2017, at Scheveningen and Hoek van Holland
several specimens were found “with C repidula fornicata
inside the aperture—the first ones recorded (Tables 1 and 2).
Where specimens on the outside are convex and rather dark
colored due to the presence of numerous brown spots, the
females on the inside are quite flat and predominantly white
or pale colored (compare Figures 11-15 with Figure 10).
Their outline reflects the inside of the last whorl ae the host
shells, and being flat most likely minimizes hindrance
resulting from movements of the host. In many specimens
on the inside a short stack formed with one to two specimens
on top of the female. Those are typically more globose
(Figures 5, 7, 10, 11) and may be darker colored, not only as
juveniles (Figure 7), but also as adults (e.g., Figure 10). As
the eonnmencell grow there soon will be no space left for the
host. Some shells were found with such large C. fornicata
Bees that indeed no space was available for a host (Figures
7, 10). It remains uncertain whether growth of the com-
Cae eventually killed the host, cone her the host died by
natural cause, but more likely it just moved to another shell.
In any case, the commensals on the outside and inside
survived. These were always shells that have been occupied,
apparently for some time, by a hermit crab, as evidenced by
the large size of Crepidula fornicata, the presence of a large
number of boreholes from boring SPOMEES, and remains aii
barnacles and hydrozoans (see also Table 2).
When living inside gastropod shells occupied by hermit
crabs, Crepidula fornicata has a flat shell (H max. =
6.3 mm: H/L ratio = 0.21 to 0.31 or 0.24 + 0.032 (n= 11))
having its head oriented towards the opening.
Some specimens also had a small specimen in a position
not noted before: on the adapical side of the aperture of
the host shell (e.g. Figures 5, 9, 10), in one case a stack of
two. These specimens are always convex and dark colored.
They occur on shells with or without Crepidula fornicata
on the inside.
As mentioned above, C. fornicata is a generalist, but
unlike Grandicrepidula grandis it has thus far not been
reported from the interior surfaces of ¢ gastropod shells
occupied by hermit crabs (Vermeij, 1989). The only ev-
idence that was found of C. fornicata living inside a shell is
a photograph on a website (Krisberg, 2011) of both C.
fornicata and C. plana Say, 1822 inside the aperture of
Sinistrofulgur sinistrum (Hollister, 1958) from Fort
Pierce Inlet (Florida). Although there is no mention of a
hermit crab being present, the presence of very flat
Crepidula for nicata inside the aperture suggests that the
shell must have been occupied by a hermit cual (Walker,
1989), but the overgrowth indicates the shell had been
abandoned for some time. Interestingly, these Crepidula
fornicata are flat, despite the aperture being very large.
Vermeij (1989) concluded his paper stating: “It “wil Ibe
interesting to determine whether C. fornicata in Europe
occupies the interior surfaces of hermit-crab shells, a habit
rarely if ever occupied by C. fornicata in its native
American range.
This question can now be answered: yes, it does.
Specimens of Crepidula fornicata living inside the ap-
erture are very flat and white or pale- <ealonedl and thus are
similar to Crepidula maculosa Conrad, 1846. However,
the sinuate septum and the protoconch of one whorl
(when preserved) demonstrate we are not dealing with a
new American immigrant (a good comparison between
the two species is provided by Krisberg (2009). Numerous
other calyptraeids that live mainly “anettle hermit-crab
occupied gastropods are flat and white, such as the slip-
per limpet Ergaea walshi (Reeve, 1859) (own observa-
tions in northwestern Bomeo; Figures 17-18) and the
species of the Crepidula plana species complex in North
America (Collin, 2000).
Ergaea walshi benefits from its association with hermit
crabs through increased food supply, reduced predation,
and (in the northern part of its range) extended breeding
season due to host migration to more suitable environ-
ments during winter (Yipp, 1980). Yoshikawa et al. (2018)
describe how the commensal prefers hermit-crab occu-
pied shells with wide aperture, which it finds by selecting
larger hermit crabs. Even so, it has to limit its thickness to a
maximum of about 5 mm to not be hindered by the
movements of the hermit crab (note Yoshikawa et al. (2018)
exclude the septum from the measurement of shell height).
It thus has the flattest shell of 23 species of Calyptraeidae
they investigated. The shell of C. fornicata living inside the
aperture is “flatter than that of Grandicrepidula g grandis
living inside the aperture (H/L ratio of 0.30 = 0.025
(r= i10) Vermeij, 1989), but not as flat as that of E. walshi
(H/W ratio of 0.11 to 0.18 or 0.14 + 0.027 (n=6)). Also, itis
elongate whereas E. walshi typically has a wide. shell
(compare Figures 11-15 with 17-18) as it has its head
towards the adapical side of the aperture (own observations;
this is why above H/W ratio is used for E. walshi).
Species with flat white shells have evolved multiple
times within Crepidula and as eco- phenotypes within
Crepidula species (Collin, 2019). Although in our case the
flatness of the shells has a logical explanation, no specific
cause has thus far been ‘dlentited for these shells being
much lighter colored than specimens on the antisite.
Interestingly, the flatness and pale coloring only applies to
C. fornic ue directly attached to the host diel Specimens
that settle on these to form a stack have a more convex
shape and (slightly) darker color (Figures 5, 7, 8, 10-12).
Two specimens on the photogr aph. from lecteh (men-
tioned above) form a stack, with the specimen on top
being more convex.
Page 44
Whereas Yoshikawa et al. (2018) found E. walshi not
showing host specificity for particular species of hermit
crabs or snail shells, in Zuid-Holland C. fornicata occurs
with the hermit crab Pagurus bernhardus in shells of E.
catena, which is probably opportunistic as in this area
these are currently the only large hermit crab and most
frequent large gastropod. No E. catena have been found
with shells of C fornicata only on the inside.
That leaves the question why C. fornicata started oc-
cupying the narrow space inside the aperture. Yoshikawa
et al. (2018) state that for E. walshi this position gives
protection from physical disturbances and predation.
Water currents set up by the feeding of the hermit crab
are beneficial to epibionts (Conover, 1979), from which
calyptraeids inside the aperture will benefit more. Spe-
cifically for C. fornicata in Europe, competition with other
epibionts of hermit crabs may be the main reason for
choosing this position (competition between epibionts is
discussed by Karlson and Shenk, 1983). Shells with C
fornicata on the inside typically have numerous Crepidula
fornicata and other commensals (e.g. barnacles, Figures 5,
6, 10) on the outside (Table 2). No shells have been found
with C. fornicata on the inside, but not on the outside.
This is an indication that the inside is a less preferred place
that most likely is chosen at a later stage, which would
make sense as growth is greatly abted and the animal
has to build more shell for a small increase in volume. The
position on the adapical side of the aperture appears to
rank below that on the inside, although a single shell was
found with Crepidula fornicata on the owistéle and in this
position, but none on the inside of the aperture (and no
indication one has been present). In its native range, this
competition likely plays a lesser role as large gastropod
shells are plentiful ( (Conover, 1979). The shells themselves
are also much larger (for example the Sinistrofulgur
sinistrum maeatiionedl above can be up to 45 cm long) and
thus each shell occupied by a hermit crab provides space
for many more epibionts than the Euspira catena shells
of <4 cm long in the study area. Karlson and Shenk (1983)
reported 96 G fornicata from the outside of a single
Busycon carica (Gmelin, 1791) shell.
The shells with numerous and/or large Crepidula for-
nicata may have been abandoned by the hermit crab that
occupied them. Hermit crabs change shells as they grow
and carefully select the shell to live in ‘(own observations and
Yoshikawa et al, 2018). The presence of symbionts in-
fluences shell selection, Conover (1976) describes how all
Table 1.
Scheveningen Jan. 2017
Number of Euspira catena alive
Number of Euspira catena dead
Number of Euspira catena with hermit crabs
and Crepidula fornicata on outside
Number of Euspira catena with hermit crabs and
Crepidula fornicata inside and on outside
hundreds
THE NAUTILUS, Vol. 133, No. 2
hermit crabs he studied rejected shells containing barnacles.
Some rejected shells had C. fornicata, which, depending on
their weight and position, may impact the center of gravity
of the system. Shells with C. fornicata on the outside were
more often tolerated than those with C. fornicata on the
inside. On the other hand, the presence of hydroids posi-
tively influenced selection (Conover, 1976). The skeleton of
hydroids often expands the total shell volume, allowing the
crab to continue using it (own observations).
The geographic patterns of habitat generalists and
specialists in calyptraeids are not yet properly understood
(Vermeij, 1989; Collin, 2019). Only for part of the species
(and therefore genera), their life habits as generalist or
specialist have ree documented. Very few photographs
are available of calyptraeid shells inside the aperture of
hermit crab occupied shells (e.g., Collin, 2019: fig. 1A).
Such information is required as basis for a theoretical
framework that can predict shifts as described in this
paper, which could be caused by migration, changes in
number of type of available shells, competition sorta other
symbionts (including of the same species), etc. According
to Low and Tan (2014) and Yoshikawa et al. (2018).
Ergaea walshi is also found on other substrates, e.g
underneath horseshoe crabs or on dead bivalves. How-
ever, it is never found on the outside of gastropods,
whether occupied by a hermit crab or not (own obser-
vations). The species of the C. plana complex can also be
found on the outside and other substrates (Collin, 2000).
In northwestern Borneo, shells occupied by hermit crabs
frequently host another calyptraeid, Desmaulus. extinc-
torium (Lamarck, 1822), which lives only on external shell
surfaces and is never found on the inside or on another
substratum (own observations).
The understanding of the behavior of calyptraeids as
habitat generalist or specialist is also relevant for the
recognition of hermit crab-occupied shells in the fossil
recor ral Walker (1992) specifically mentions that external
shell-inhabiting species of Crepidula are not good in-
dicators of Theses crab occupancy, but Crepidula that
inhabit gastropod apertures are excellent indicators. In
the current study it was found that, specifically for C.
fornicata shells with C. fornicata on the outside were
likely, but not certainly inhabited by a hermit crab, as the
species lives on various substrates. If a flat specimen of
C. fornicata is present on the inside, it is likely that the
shell has been inhabited by a hermit crab: C. fornicata will
not limit its growth if there is no crab. Other indications
Overview of Euspira catena found at the localities discussed.
Hoek van Holland Hoek van Holland
Jan. 2017 Feb. 2019
tens tens
hundreds hundreds
tens tens
3 7
Page 45
J].G.M. Raven, 2019
“SUBOZOAI PUR sopoRULtE ‘FU vas
‘apuods SuLtog ‘aut Buoy] juasaid uooq
DARL] “OBILT ST[OYS taddys soy 10F Yo] aords on
JJO [J2} Opisyno uo suautoadg “Ayjenprarpul
poyorye uotttoods xoauoo [[PUIS UO sey] OpIsUy
sapeureg ‘osuods suri0q
S(QJo] SyLeUL OAY) JJO [[O}F OpIsyNO uo suouttoads
*SoTPUl OMY UpIM o[Pulot Quo 9pIts}no ud
SUBOZOAIG, “SO[OVUIRG
“yEUL BIS ‘opuods BULIOG OR UL Toy,OUuR pue
SOLS OM} UI o[Pulot 9uoO OpIS}Lo ud
JUL BIS “Se poRuteq “SoTBUL 1OYyO
OM} pue yoruys B qq o[PUlot 9uO OpIts}Ho ud
suvozodArq ‘asuods SuLtog qeut vas
‘SOPOBUIBG “SO[LUE OA) UPI o[LULo} v OPISUL UOC
suvozodiq ‘asuods Surt0q
JWUL vas ‘sapovuUE ‘soy Io} Yor oovds ON
suvozodrq ‘osuods Surto0q ‘soporuseq sno1owuNn Ny
osuods BULIO “SOORUIRG STLO.TOULTL AY
SyIBUOY
) [ I o V4 uyDpUN UNUIIING =~— BLOG YIN = puvjjoH uva yao FI
(Joy Syreut .M0F)
atnO) 0 I G Vv puayvo paudsny GLO PW — PUeTOH Uea YooH CT
+P (0) G [ d (6 puajpo vudsny LOS YW puRTJOH{[ URA YOR, ZI
JUL Bas ‘SoPRUIe
9 | % I 6 [ puajpo badsny LOS YW puRT[OH{ ura xooy] [I
A, 0 I I I puajpo vadsny GLOS YIW —_ purT[OH{ UeA YOoH] OT
OT 0 G I € L puaywo vudsny 610% YIN PUR[[OH UkA YOOH 6
JUL BOS ‘soprUIEg G 0) I I p I puajpo vadsng LOS YW pur[OH{ ura yoor] 8
yeut vas ‘asuods Suriog Pp 0 I I 6 b puajpo pudsngy GLOS YW purR[OP{ ura yoox]
asuods SuLIOg yRUl vas (6 0 S I 0) () puayo virdsny L[0G ‘uef[ puryjoyy ura yoox] 9
Pp () 6 I [ (6 puaypo virdsny LOG ‘uef[ puryjoyy uea Yoox] G
b 0 fp I ¢ [ puaypo vudsng 2.70% “uel puR]OF{ ua yooH] fp
6 0 I I I I puaywo vudsny LTO% ‘uel UdBUIUDAIIPG —-&
sopoRuleg g 0) [ I I G nua vardsny LT0% ‘url UIBULUDADYG
L T S I ¢ % puawo vudsny 110% ‘uel UasUTUdAIYYS |
suowutoods # — yoy toddn uo yor}s Ul Sopeuloy # yorys Ul so[RUloy # [Joys ISOH awd AyRoOT
[ROL uouttoads suowutoads suowutoads
JO # XP] JO # Xe
WUOL YY opisuy apismoO
De ee ee
‘So[PUlof oq O} peounsse ole [[P Us }SOU out OF peyprye A}}9.11p sy[eys he JON “Optsul out uo DVIUALOf 1) UI s]peys [[P jo MOIALIAQ) °G 9[4PRL
Page 46 THE NAUTILUS, Vol. 133, No. 2
Figures 11-18. Calyptraeidae. 11-15. Crepidula fornicata found inside the aperture of hermit crab-inhabited Euspira catena,
Scheveningen, Jan. 2017. 11-12. Slipper stack (Table 2: 1). 11. Female (L = 17.6 mm, H = 4.6 mm). 11a. Ventral view. 11b. Dorsal
view. 11e. Lateral view. Note the protruding septum. 12. Male (L = 9.7 mm, H = 3.7 mm). 12a. Later view, “stretched” to same
length as female to show difference in shape and resulting space available. 12b. Ve a view. 13. Shell from E. catena in Figure 5 (Table
(L 14.4 mm, H 3.3 mm. 13a. Ventral view. 13b. Dorsal view. 14. Dorsal view (Table 2: 2) (L = 19.9 mm, H = 4.7 mm). 15. Shell
from Euspira catena in Figure 7 (Table 2: 4). Base of a stack of four (L = 26.0 mm, H = 5.5 mm). 15a. Ventral view. 15b. Dorsal view.
16. Euspira catena (H = 32.6 mm) with imprints of C. fornicata on the outside, Hoek van Holland, Feb. 2019 (Table 2: 13). Note in this
(rare) case not only the foot left an imprint, but also the tentacles and lips. The animals were oriented toward the aperture. This specimen
also had a C repidula fornicata inside the aperture. 17-18. Ergaea walshi found inside the aperture of hermit-crab inhabited gastropods,
Piasau Beach, Miri, Sarawak, Malaysia, June-Sept. 1992. 17. Adult female (W = 33.2mm, H = 5.6 mm). Note the laterally elongate,
concave shape (optimally conforming to the inside surface of the host shell), the se ptum indicating the dnidlane »ss of the living specimen,
the barnacle, and the imprint of another barnacle. 17a. Ventral view. 17b. Dorsal view. 17e. Lateral view. Note the protruding septum.
18. Juvenile male (W = 6.6 mm, H = 1.3 mm). Note rounded shape. 18a. Ventral view. 18b. Dorsal view.
J.G.M. Raven, 2019
that a shell has been inhabited by a hermit crab are
specific wear-and-tear patterns and the presence of other
symbionts: barnacles, sea mat, etc. (own observations:
Walker, 1992 gives many examples). The recognition of
shells that ees been occ upied by hermit crabs is relevant
both for recent communities and fossil thanatocoenoses,
as hermit crabs frequently displace shells to different
depositional environments than those where the mollusks
lived (own observations in northwestern Borneo and
Walker, 1989). It would be useful to construct a broader
framework including all calyptraeid species.
ACKNOWLEDGEMENTS
Gregory Herbert (University of South Florida) and Geerat
Vermeij (University of California) are thanked for critically
reviewing this work. Their constructive comments have
helped placing several observations into a wider context.
LITERATURE CITED
Anonymous. 2019. Global invasive species database. Species
profile: Crepidula fornicata. _http:/Avww.iucngisd.org/gisd/
species.php?sc=600 [consulted 6-2-2019].
De Bruyne, R.H., S. van Leeuwen, A. Gmelig Meyling and R.
Daan (eds.). 2013. Schelpdieren van het Neds landse
Noordzeegebied. Ecologische atlas van de mariene week-
dieren (Mollusca). Tirion Natuur en Stichting ANEMOON,
Leiden. j
Collin, R. 2000. Phylogeny of the Crepidula plana (Gastropoda:
Calyptraeidae) cryptic species complex in North America
Canadian Journal of Zoology, 78: 1500-1514.
Collin, R. 2006. Sex ratio, life-history invariants, and patterns of
sex change in a family of protandrous gastropods. Evolu-
tion, 60(4):735-745.
Collin, R. 2019. Calyptraeidae from the northeast Pacific (Gas-
tropoda: Caenogastropoda). Zoosymposia, 13: 107-130.
Conover, M.R. 1975. Prevention of shell burial as a benefit hermit
crabs provide to thei symbionts (Decapoda, Paguridae).
Crustaceana, 29(3): 311-313.
Conover, M.R. 1976. The influence of some symbionts on the shell-
selection behaviour of the hermit crabs, Pagurus pollicarus
and Pagurus longicarpus. Animal Bahamian 24: 191-194.
Conover, M. R., 1979. Effect of gastropod shell characteristics
and hermit crabs on shell epifauna. Journal of Experimental
Marine Biology and Ecology, 40: 81-94.
Fretter, V. and A. Ge aham. 1981. The prosobranch molluscs of
Britain and Denmark. Part 6. Journal of Molluscan Studies,
Supplement 9: 309-313.
Page 47
Karlson, R.H. and M.A. Shenk. 1983. Epifaunal abundance,
association, and overgrowth patterns on large hermit crabs.
Journal of Experimental Marine Biology ma Ecology 70:
55-64.
Krisberg, M., 2009 and 2011. ee fornicata (Linnaeus,
1758). In: Let’s Talk Seashells. https:/Avww.tapatalk.com/
groups/conchologisttorum/crepidula- -fornicata-linnaeus-1758-
t1816.html [consulted 7-5-2019].
Low, M. and S.K. Tan. 2014. Ergaea H. Adams & A. Adams,
1854, the correct genus for Crepidula walshi Reeve, 1859,
with nomenclatural notes on Syphopatella Lesson, 1831,
Siphonipatella L. Agassiz, 1846, and the incorrect sub-
sequent spelling “Siphopatella” of authors (Gastropoda:
Calyptraeoidea: Calyptraeidae). Occasional Molluscan Pa-
pers, Publishing on Malacology in the Sundaland Region 3:
11-14.
Raven, J.G.M. 2017. Mooie vondsten na de storm van 13 januari
2017 en reflecties op veranderingen in de kustfauna van
Zuid-Holland. Spirula, 411: 97-34.
Thieltges, D.W., M. Strasser and K. Reise. 2002. The American
slipper limpet Crepidula fornicata (L.) in the northern
Wadden Sea 70 years after its introduction. Helgoland
Marine Research (2003) 57: 27-33.
Vermeij, G.J. 1989. Habitat and form of Crepidula grandis in
Japan, with comments on habitat specialization in calyptraeid
gastropods. The Nautilus 103: 89- 91.
Walker, S.E. 1989. Hermit crabs as taphonomic agents. Palaios
4(5): 439-452
Walker, S.E. 1992. Criteria for recognizing marine hermit crabs
in the fossil record using gastropod shells. Journal of Pa-
leontology 66(4): 535-558.
Walne, P.R. 1956. The biology and distribution of the
slipper limpet Crepidula | fis nicata in Essex rivers with
notes on the distribution of the larger epibenthic in-
vertebrates. Fisheries Investigations London, Series 2,
20(6): 1-50.
Williams, J.D. and J.J. McDermott. 2004. Hermit crab bio-
coenoses: a worldwide review of the diversity and natural
history of hermit crab associates. Journal of Experimental
Marine Biology and Ecology 305: 1-128.
Wolff, W.]. 2005. None indigenous marine and estuarine species
in the Netherlands. Zoologische Medelingen Leiden 79(1):
1-116.
Yipp, M.W. 1980. The functional morphology of the organs of
feeding and digestion in Crepidula walshi (Prosobranchia:
Calyptraeidae). In: Morton, B. (ed.). Proceedings of the
first international workshop on the malacofauna of Hong
Kong and Southern China. Hong Kong University Press,
Hong Kong: 221-252.
odie A., R. Goto and A. Asakura. 2018. Morphology and
habitats of the hermit-crab-associated calyptraeid gastropod
Ergaea walshi. Zoological Science 35: 494-504
THE NAUTILUS 133(2):48-56, 2019
Page 48
The Miocene to Recent biogeographic history of vesicomyid
bivalves in Japan, with two new records of the family
Kazutaka Amano
Department of Geoscience
Joetsu University of
Education
Joetsu 943-8512, Japan
amano@juen.ac.jp
Yusuke Miyajima
Geochemical Research Center
Graduate School of Science
University of Tokyo
Bunkyo-ku, Tokyo 113-0033, Japan
yusukemiya@eqchem.s.u-tokyo.ac.jp
Steffen Kiel
Swedish Museum of
Natural History
Department of
Palaeobiology
Box 50007, "10405
Stockholm, Sweden
steffen.kiel@nrm.se
Robert G. Jenkins
School of Geosciences and
Civil Engineering
College of Science and
Engineering
Kanazawa University
Kanazawa City, Ishikawa
920-1192, Japan
robertgj@staff. kanazawa-u.ac.jp
ABSTRACT
We report on two fossil species of the chemosymbiotic bivalve
family Vesicomyidae that were recently collecte d from Cenozoic
strata in Japan. The new species Ple surophopsis matsumotoi is
described from the upper Oligocene to lower Miocene Hioki
Complex in Kochi Prefecture, and the extant species Calyp-
togena pacifica Dall, 1891 is reported from the upper Miocene
Onnagawa Formation in Akita Prefecture. With these new re-
cords, vesicomyid bivalves in Japanese strata show the following
distribution pattern through the latter half of the Cenozoic Era:
during the early to wattilalle Miocene Plewrophopsis was the
dlonatiacrat genus and is found in sediments of both the Japan Sea
and the Baxettiie Ocean. From the late Miocene through to today
Archivesica and Calyptogena are the most common genera. Of
these, Calyptogena species including the extant C. pacifica, are
the most common vesicomyids in the cold waters of the Japan
Sea, whereas Archivesica species are mostly found in the warmer
waters of the Pacific side of Japan. Since the late Miocene, the
diversity of vesicomyids rapidly increased in the Japan Sea,
prob yably because of the semi-enclosed geographic situation with
opened northern straits.
Additional Keywords: Plewrophopsis, Calyptogena, _ fossil,
paleobiogeography
INTRODUCTION
Vesicomyids are peculiar bivalves that derive their nu-
trition from symbiotic, sulfur-oxidizing bacteria (Fisher,
1990). They include the iconic Gane White Clam
“Calyptogena” magnifica that was discovered at deep-sea
hydrothermal vents in the late 1970s (Boss and Turner,
1980). To date, more than 100 extant species have been
found at hydrothermal vents, hydrocarbon seeps, and
whale falls (e.g g., Taylor and Glover, 2010; Krylova et al.,
2010). Voriconnads also have a good fossil record, in-
(o>)
cluding more than 30 named fossil species, the oldest
being from the middle Eocene (Amano and Kiel, 2007;
Kiel and Taviani, 2017: and references therein).
Japan has a rich Cenozoic fossil record of hydrocarbon
seeps, W hale falls, and organic-rich shales, from which
species belonging to five vesicomyid genera have been
reported: Adulomya Kuroda, 1931 (= Pleurophopsis Van
Winkle, 1919, see discussion below), Archivesica Dall,
1908, Calyptogena Dall, 1891, Hubertschenckia Takeda,
1953, and Pliocardia Woodring, 1925 (Kanno et al., 1989;
Amano and Kiel, 2007, 2010, 2011, 2012; Amano and
Suzuki, 2010; Amano, 2014). Of these genera, Hubert-
schenckia is an exclusively fossil genus lenovo from upper
Eocene to lower Oligocene strata. Only two extant species
are known from the fossil record: Calyptogena pacifica
Dall, 1891 and Archivesica kawamurai (Kuroda, 1943)
(Amano and Jenkins, 2011). Although numerous new
vesicomyid species have been discovered and described in
the last two decades, the history of this family in Japan has
never been thoroughly reviewed from a biogeographic
point of view.
Here we review and discuss the biogeographic history
of vesicomyid bivalves in the Miocene—Recent of Japan,
describe a new species of Pleurophopsis from the “Muroto
Formation” in Kochi Prefecture, Shikoku, and report the
new fossil record of Calyptogena pacifica from the
Onnagawa Formation in Akita Prefecture, Honshu.
MATERIALS AND METHODS
We carried out a comprehensive review of the litera-
ture on fossil vesicomyids in Japan to assess their paleo-
biogeographic distribution, including the two records
reported herein. Excluded from the review are the ves-
icomyids from the lower to middle Miocene Taishu Group
(Ninomiy a et al., 2014) in Tsushima Island at the western
entrance of the Japan Sea. This is so because most
specimens described by Ninomiya (2011) are deformed
kK. Amano et al., 2019
and internal shell features were not illustrated, hence their
generic assignments remain uncertain.
The eam specimens described here as Plewrophopsis
matsumotoi new species were collected from a limestone
nodule from mudstone of the Hioki Complex of the Nabae
Group. The nodule included also some specimens of a
thyasirid bivalve of the genus Conchocele. It was found about
1,500 m north of Mitsu Maruyama, Muroto City, in Kochi
Prefecture, Shikoku, Japan ( at 33°18'15” N, 134°11’23” E;
Figure 1, Loc. 1). The vesicomyid specimens were pre-
viously reported as “Akebiconcha uchimuraensis” by Mat-
sumoto and Hirata (1972). The age of the Hioki Complex
was considered as late Oligocene based on planktonic fo-
raminifera (Taira et al., 1980) and as late Oligocene to early
Miocene based on radiolarian fossils (Suyari et al., 1989).
Seven specimens identified as the extant species
Calyptogena pacifica Dall were recovered from a cal-
careous concretion derived from shales of the Onnagawa
Formation, found as float under the bridge at Ashigafuchi,
Chokai-cho, Yurthonjo City in Akita Prefecture, Honshu,
Japan (at 39°11'15” N, 140°11'39.8" E; Figure 1, Loc. 2).
The age of the Onnagawa Formation ar ound this locality is
considered as late Miocene (around 8.4 Ma) based on
radiolarian fossils (Tsuji et al., 1991).
All specimens described and illustrated here are de-
posited in National Museum of Nature and Science,
Tsukuba, Japan (NMNS PM for fossils and NSMT-Mo for
Recent specimens).
SYSTEMATICS
Family Vesicomyidae Dall and Simpson, 1901
Genus Pleurophopsis Van Winkle, 1919
130°E (40° (\ Okhotsk
ZANs Sea
—\'
Y ) Hokkaido_
| SAG
\
f) ‘(7 mee
Ko
40.N _ of —— ed et
A oe ‘) Honshu
C Ro) fe2 J
C Ka / f
z > or
\ y ¢
é oo &
\ nf
(¢ al aa CN D Bs ae a Bad
A pe? y = (he { ° o
nw” p SJ 2) ae
dn pe 7 as Oy Y
—-) A} J A
r POI Shine Aiel
be
Bs te) 200km
of
30 .—— — = -- = = —————— = = ——
Figure 1. Localities of the new vesicomyid fossils described in
this paper.
Page 49
Type Species: Pleu rophopsis unioides Van Winkle, 1919
(monotypy), middle Miocene, Trinidad (Van Winkle,
1919: Kiel, 2007).
Remarks: The genus Plewrophopsis was introduced and
used for elongated, fossil vesicomyids from the Caribbean
region, Peru and Ecuador (Van Winkle, 1919: Olsson
1931, 1942). Its shell characters are virtually identical to
those of Adulomya Kuroda, 1931, including an elongate
shell, two cardinal teeth in the right welve. the anterior
original point of the pallial line at the posterior side of
arntreretor adductor muscle scar, and the lack of a pallial
sinus and a subumbonal pit (Kiel, 2007; Krylova et al.,
2010). Kiel (2007) considered the genus doubtful because
the type material of Plewrophopsis unioides is lost and the
then available material lacked critical hinge features to
fully characterize the species; he suggested using Adu-
lomya instead. Krylova et al. (2010) essentially followed
this view, but Krylova and Sahling (2010) included
Pleurophopsis in their list of wesioonanial genera. Ongoing
work on new material of Pleui “ophopsis unioides from the
type locality (SK and CTS Little) indicates that Pleuro-
phopsis is a valid genus and that Adulomya should be
synonymized with it. The name Aduloma ya has long been
used for elongated fossil vesicomyids in Japan (Kanno
et al. 1998; Majima et al. 2005; Amano and Kiel, 2007,
2011; Isaji, 2013; Miyajima et al., 2017) and virtually all
species previously assigned to Adulomya match the
characteristics of Plewrophopsis. Only two species show a
slight deviation: in Adulomya chitanii Kanehara, 1937, the
pallial line bends backwards before reaching the posterior
adductor scar; this is unlike other Adulomya/Pleuro-
phopsis species but we consider this not to represent an
actual pallial sinus. Adulomya hokkaidoensis Amano and
Kiel, 2007 has a very small subumbonal pit, which,
however, is very unlike the lar ge and elongate-triangular
subumbonal pit of Ectenagena elongata (Dall, 1916)
which is otherwise very similar.
Abyssogena Krylova, Sahling, and Janssen, 2010 is an
extant vesicomyid genus with very elongated shells. In
particular, the deep-sea living species Abyssogena pha-
seoliformis (Métivier, Olienterat, and Ohta, 1986) from the
Japan. Kurile, and Aleutian Trenches, resembles the type
species of Adulomya, A. uchimuraensis (Kuroda, 1931) in
having a large and very elongated shell (Figure 9).
Horecee a9 fnillowa et all (2010) observed, Abyssoge na
can be easily distinguished from Pleurophopsis in having
an anterior original point of the pallial line located at
ventral part of aie anterior adductor muscle scar and
possessing an indistinct and irregular shaped pallial sinus
(Figures 8, 11, 12).
Like Pleurophopsis and Abyssogena, Ectenagena
Woodring, 1938 also has an elongate shell and two cat ‘dinal
teeth in the right valve. Ectenagena also shares with Pleu-
rophopsis the anterior point of origin of the pallial ne located
at the posterior part of anterior adlaineion muscle scar (Krylova
et al., 2010). However, Ectenagena has a small (up to 50 mm;
Coan et al., 2000), very itn compressed shell with a
short nymph. Further, unlike Abyssogena and Pleurophopsis
Page 50 THE NAUTILUS, Vol. 133, No. 2
Figures 2, 3, 5-7, 10, 11, 13, 14. Pleurophopsis matsumotoi new species. 2, 3. Right-valve hinges, NUNS PM13233. 5. Paes
(NMNS PM13227), left valve showing anterior adductor muscle scar (AAS) and anterior original point of pallial line (glitters arrow). 6, 11,
13. poeies (NMNS PM13228). 6. Dorsal view. 11. Left valve, showing anterior adductor muscle scar (AAM) and anterior original
point of pallial line Se arrow). 13. Right valve, showing posterior adductor muscle scar (PAS) and posteriorly backward bent pallial
line (white arrow). 7. Dorsal view, NUNS PM28254. 10. Left valve. showing posterior adductor muscle scar (PAS) and posteriorly
backward bent alia line (white arrow), NMNS PM28255. 14. Right valve, showing posterior adductor muscle scar (PAS) and
posteriorly backward bent pallial line ethitie arrow), NMNS PM13256. Figures 4, 9,12. Pleurophopsis uchimuraensis (Kuroda). 4. Left
valve hinge, NMNS PM28257. 9. Left valve, NMNS PM28258. 12. Right valve, showing anterior adductor muscle scar (AAS) and the
original point of pallial line (white arrow), NMNS PM28259. Figure 8. Abyssogena phaseoliformis (Métivier, Okutani and Ohta), Holotype,
NSMT-Mo 64164, showing anterior adductor muscle scar (AAS) and the original point of a pallial line (white arrow). Scar bars = 10 mm.
kK. Amano et al., 2019
(except for P. hokkaidoensis), the hinge of Ectenagena has a
deep subumbonal pit.
Pleurophopsis matsumotoi new species
(Higures 2,3; 5=7, 10; 11) 13, 14)
Akebiconcha uchimuraensis (Kuroda). Matsumoto and
Hirata, 1972: 755-757, pl. 1, figs. 1-8, pl. 2,
figs. 1-2
Adulomya? sp. Amano and Kiel, 2011: figs. 30-31.
Diagnosis: A large-sized, well-inflated and moderately
elongate Plewrophopsis; antero-dorsal margin short,
posterior margin subtruncated, ventral margin concave;
narrow hinge plate with thin anterior Gardin tooth (1)
and rather ‘Ghialk posterior cardinal tooth (3b); posterior
end of pallial line turning toward to anterior before
reaching posterior adductor muscle scar.
Holotype: Internal mold of articulated specimen (NMNS
PM 13228), length 112.5 mm, height 39.3 mm.
Baratypes Internal mold of articulated specimen (NMNS
PM 13227), length 93.1 mm-+, height 36.9 mm (from the
type locality).
Type locality: 1.5 km north of Mitsu, Muroto City in
Shikoku, Japan (Matsumoto and Hirata, 1972).
Material examined Four specimens from the type
locality.
Description: Shell up to 112.5 mm long, elongate
(height/length-ratio = 0.34), equiva alve and inequilateral,
well inflated (width/height-ratio = 0.67-0.73). Surface
sculptured only by rough, irregular growth lines. Beak
situated anteriorly at owt one- fifth of shell length.
Antero-dorsal margin short and nearly straight, gr aduating
into rounded arate margin; ventral margin concave;
postero-dorsal margin str aight, gently sloping, posterior
margin galinammentedl Deanizdheor and lunule absent.
Hinge plate narrow, with two cardinal teeth in right valve:
or cardinal tooth (1) very thin and inclined sa verstor ly:
posterior cardinal tooth (3b) rather thick and oblique
posteriorly; no subumbonal pit. Ligament occupying two-
fifth of the postero-dorsal margin. Nateesétor adductor muscle
scar ovate, bordered posteriorly by thick ridge; posterior
adductor muscle scar ovate, less distinct than the anterior
scar; distinct ridge running from umbonal area to ventral side
of posterior muscle scar. Original point of pallial line located
at posterior part of anterior adductor muscle scar, and pallial
line at posterior end bent toward to anterior before reaching
posterior adductor muscle scar.
Remarks: Pleurophopsis matsumotoi new _ species
represents the as-yet oldest record of Pleurophopsis (=
Adulomya) in Japan.
Comparison: Pleurophopsis matsumotoi differs from
P. uchimuraensis (Kuroda) (Figure 4, 9, 12) in having a
much more higher shell (height/length-ratio = 0.34
compared to 0.17-0.24 in P. uchimuraesnis; Kanno
Page 51
et al., 1998, Amano and Kiel, 2011), an _ inflated
concave ventral margin, and the pallial line turning
backward before reaching the posterior adductor
muscle scar. Pleurophopsis pa sornanatb also shares the
posteriorly backward bent pallial line with P. chitanii
(Kuroda). However, P. matsumotoi has a larger shell
than P. chitanii (up to 70.4 mm in length) and a
concave ventral margin. The present new species
differs from other Pleurophosis such as P. akanudaensis
(Tanaka), P. hamuroi Amano and Kiel, P. kuroiwaensis
Amano and Kiel from Japan in having larger, more inflated
shell and a concave ventral margin.
Distribution: Only from the type locality.
Etymology: For Dr. Eiji Matsumoto who collected the
type material of this new species.
Genus Calyptogena Dall, 1891
Type Species: Calyptogena pacifica (
Recent, southeastern Alaska.
Dall) (monotypy),
Calyptogena pacifica Dall, 1891
(Figures 15-24, Table 1)
Calyptogena pacifica Dall, 1891: 190; Dall, 1895: 713, pl.
25, fig.4; Grant and Gale, 1931: 278-279, pl. 13, fig.
13a, Be Otuka, 1937: text-fig.; Woodring, 1938: fig. ah:
Otatume, 1942: 435-437, pl.16, figs. 1- 12: Olean,
1966: 301, pl.27, figs. 1, 3; Boss, 1968: 8: figs.16, 17, 19,
20: Keen, 1969: N664, fig. E138, 11a, be Tiba, 1972
155, pl. 19, figs.6, 6a; Amano and Kanno, 1991: as
4.18, ‘4.19. Blorleosth and Hashimoto, ee yall, IL, fig.
da, pl. 2, fig. 4b; Okutani et al., 1993: fig. 7 - Okutani,
2000: 997, pl. 496, fig. 7; Amano, 2002: OT, as 3.4, 3.9;
Amano, 2003: figs. 3-14; Amano and Kanno, 2005:
204-207, fig. 3; Krylova and Sahling, 2006: 362-368,
figs. 3-6; Sonal, 2007: figs. IH-3-4.5, III-3-4.6; Huber,
2010: 354; Amano and Jenkins, 2011: 166-169, figs.
3-16; Nevesskaja et al., 2013: fig. 150-9; Okutani, 2017:
1233, pl. 528, fig. 8.
Unio moraiensis Saad, 1941: 55-56, pl. 4, figs. 2-5.
Calyptogena (Calyptogena) pacifica Dall. Berard, 1974:
11, figs. 1A, 2A, 3A, 4A-D; Boss and Tumer, 1980:
188— 189, figs. 10B, C; Kanno et al., 1989: figs. 1.7-1.15.
Calyptogena sp. Tsuji et al., 1991: fig. 25, 26.
Vesicomya (Calyptogena) pacifica ( Dall). Coan et al.,
2000: 341, pl. 70.
Material Examined: Seven specimens. NMNS PM28260—
PM28266.
Remarks: The shells are rather small (less than 30.0 mm
in length) and ovate in outline (height/length = 0.68,
(0.71). Their beaks are located at anterior one-fifth to two-
fifths of shell length (umbo% = 21, 41). In the right valve,
a posterior tooth (3b) is large and triangular with a small
thin anterior tooth (3a), showing U- shaped connection,
and surrounding a small middle eon ( 1). The pallial line
is entire. These shell and hinge characters are identical to
Page 52 THE NAUTILUS, Vol. 133, No. 2
Figures 15-24. Calyptogena pacifica Dall from the upper Miocene Onnagawa Formation. 15. Left valve hinge, silicone rubber cast,
NMNS PM28263. 16, 17, 19. 16. Silicone rubber cast of inner part of left valve. 17. Left valve. 19. Silicone rubber cast of inner part of
right valve; NMNS PM28260. 18. Rubber cast of right valve hinge, NMNS PM28261. 20, 23, 24. 20. Dorsal view. 23. Left valve. 24.
Right valve; NMNS PM28262. 21. Inner part of left valve, showing pallial line (pl) and posterior adductor muscle scar (PAS), NMNS
PM28264. 22. Inner part of right valve, showing anterior adductor muscle scar (AAS), NMNS PM28265. Scar bars in 15-19 = 5 mm and
20-24 = 10 mm.
those of Recent Calyptogena pacifica specimens. Tsuji end and a long anterior cardinal tooth (3a) in the right
et al. (1991: fig. 25, 26) identified and illustrated a valve.
specimen as Calyptogena sp. collected from the Onna-
: te : : Distribution: Upper Miocene: Morai Formation (Ota-
gawa Formation, near the fossil locality discussed here.
tume, 1942; Amano, 2003) and Toyama Formation
(Suzuki, 2007) from Hokkaido, Akaishi Formation
(Amano and Jenkins, 2011) from Aomori Prefecture,
Onnagawa Formation (this study) from Akita Prefecture,
Nodani Formation (Kanno et al., 1989; Amano, 2002)
Although no pallial line and hinge features were shown,
the outline of their specimen is similar to that of
Calyptogena pacifica illustrated here. (See Table 1 for
morphometric data. )
Comparison: The oldest species of Calyptogena, C. from Niigata Prefecture; Pliocene: Kurokura, Kawazume
I I Z yplog §
katallaensis Kiel and Amano, 2010 from the Oligocene and Nadachi Formations (Kanno et al., 1989: Amano and
o
Kulthieth Formation in Alaska is most similar to C. Kanno, 1991; Amano and Kanno, 2005) from Niigata
pacifica. However, C. pacifica differs slightly from the Prefecture; middle Pleistocene: Wakimoto Formation
Oligocene species by having a nymph with a rather abrupt (Otuka, 1937) from Akita Prefecture: Recent: Sea of
g I , g anymy I
Table 1. Morphometry of Calyptogena pacifica studied.
Measurements (mm):
length (L) height (H) H/L Umbo (%)* Valve
NMNS PM28260 18.0 12.3 0.68 4] left
NMNS PM28262 30.0 21.3 0.71 21 left
NMNS PM28263 29.6 +18.9 - = left
* Ratio of anterior length (distance from umbo to anterior margin) to shell length.
k. Amano et al., 2019
Okhotsk (Tiba, 1972), Dixon Strait, Alaska to Monterey
Bay, California (Coan et al., 2000).
BIOGEOGRAPHIC DISTRIBUTIONS OF
MIOCENE TO RECENT VESICOMYIDS IN JAPAN
In this section, we review the geographic distribution of
vesicomyid species and genera in Japan from the Miocene
to the present (Table 5 2) in relation to tectonics, paleo-
geography, and climate. Following the appearance of
Pleurophopsis matsumotoi new species around the early
Miocene, Pleurophopsis was the dominant vesicomyid
genus in Japan until the middle Miocene. The Japan Sea
Page 53
was formed in the early Miocene and, initially, had deep-
water connections with the Pacific Ocean through the
central part of Honshu, an area called the Fossa Magna
Region (Iijima and Tada, 1990; Ogasawara, 1994). The
first vesicomyids to colonize the Japan Sea were Pleuro-
phopsis chitanii and Pliocardia kawadai, with the oldest
record from the lower Miocene Kurosedani Formation in
Toyama Prefecture (Amano et al., 2019). Through the
early Miocene to early middle Miocene, Plewrophopsis
chitanii, along with Pleurophopsis uchimuraensis and
Pliocardia kawadai, occurred both in the Japan Sea and
the Pacific side of Japan, while a remarkable number of
endemic species evolved in the Japan Sea, namely
Table 2. Age and distribution of the fossil vesicomyids without the Paleogene species in Japan. * Species from the Sea of Okhotsk.
** Bessho Formation in Nagano Prefecture is treated as the Japan Sea scl
Species Age Pacific Japan Sea Formation
Archivesica sp. Pliocene tf Horinouchi F.
Archivesica ? bosoensis (Kanie and Pliocene + Shiramazu F.
Kuramoch, 2001)
Archivesica shikamai Amano and Kiel, Pliocene ale Ikego F.
2010
Archivesica kawamurai (Kuroda, 1943) Pliocene-early + Tomioka F., Hitachi F., Na-arai
Pleistoscene F., Kurotaki F., Ikego F.,
Imaizumi F., Hijikata F, Tamari
Siltstone, Shinzato F.
Archivesica kannoi Amano and early Pliocene + Kurokura F.
Kiel, 2010
Calyptogena veneriformis Amano and early Pliocene + Kurokura F.
Kiel, 2012
Pleurophopsis sp. early Pliocene + Kurokura F.
Archivesica shiretokensis (Uozumi, 1967)* late Miocene Rusha F.
Archivesica nipponica (Oinomikado and late Miocene-early + Kubiki F., Araya F.
Kanehara, 1938) Pliocene
Calyptogena pacifica Dall, 1891 late Miocene—middle P Onnagawa F., Morai F., Toyama F.,
Pleistocene Akaishi F., Nodani F.,
Kawazume F., Nadachi F.,
Kurokura F., Wakimoto F.
Pleurophopsis kuroiwaensis (Amano latest middle Miocene + Ogaya F.
and Kiel, 2011)
Pleurophopsis akanudaensis (Tanaka, middle Miocene + Bessho F.**
1959)
Pliocardia ? tanakai Miyajima, middle Miocene 4 Bessho F.**
Nobuhara and Koike, 2017
Pleurophopsis hokkaidoensis early middle Miocene + Chikubetsu F.
(Amano and Kiel, 2007)
Pleurophopsis hamuroi (Amano early—early middle + Higashibessho F.
and Kiel, 2011) Miocene
Pliocardia kawadai (Aoki, 1954) early—middle Miocene ap + Honya F., Kurosedani F.,
Higashibessho F., Nupinai F.
Pleurophopsis uchimuraensis early—middle Miocene + + Bessho F.**, Takinoue F.,
(Kuroda, 1931) Shikiya F.
Pleurophopsis chitanti (Kanehara, 1937) early—middle Miocene + Sf Mizunoya F., Kamenoo F., Taira
F., Morozaki G., Nupinai F.,
Kurosedani F.
Pleurophosis sp. middle Miocene +f Aokiyama F.
Archivesica sakoi Amano, Jenkins, early Miocene dP Shikiya F.
Ohara and Kiel, 2014
Pleurophopsis matsumotoi n. sp late Oligocene-early + Hioki Complex
Miocene
Page 54
Pleurophopsis hamuroi, P. hokkaidoensis, P. akanu-
daensis, and P. kuroiwaensis (Table 2).
The vesicomyid faunas both sides of Japan became more
distinct from the late Miocene onward, when Archivesica
and C alyptogena became the dominant vesicomyid genera.
Several species of Archivesica have been documented from
the Japan Sea and the Pacific side of Japan, but none of those
species occurs on both sides of Japan (Table 2). However,
there is no late Miocene record of Archivesica in Japan,
partly because the late Miocene on the Pacific side is
characterized by strata barren of molluscan fossils in
northeastern Japan and by a hiatus in southwestern Japan
(Chinzei, 1986). Fossils of C alyptogena are only known from
the Japan Sea, including the extant C. pacifica. Remarkably,
the timing of the al change from the dominance of
Pleurophopsis to that of Archivesica/Calyptogena coincides
with the tectonic inversion from tension to compr ession in
Northeast Japan (Sato, 1994). However, potential reasons
for this coincidence remain unclear. Vesicomyids became
locally extinct in the Japan Sea due to deep- water anoxia
during the Pleistocene glacial period; their absence from the
present- -day Japan Sea. might be because they have been
unable to pass through es shallow straits from the Pacific
Ocean (Amano, 2001, 2007; Amano and Jenkins, 2011).
Based on the areas of origin and the ecology of the two
extant vesicomyid species that have fossil records in Japan
(Calyptogena pacifica Dall and Archivesica kawamurai:
Amano and Jenkins, 2011), and their relatives, we pos-
tulate that the observed paleobiogeographic patterns
might be broadly related to temperature preferences
among Calyptogena and Archivesica. Present-day
C alyptogena pacifica occurs mainly in the cold waters
of the North Pacific Ocean (Coan et al. 2000). The genus
Calyptogena first appeared in the Oligocene of Alege
(Kiel and Amano, 2010) and migrated southward to the
Japan Sea in the late Miocene, potentially related to the
late Miocene climate cooling (Amano and Jenkins, 2011).
The Japan Sea at that time was connected to the Pacific
Ocean mainly through its northern straits (Iijima and
Tada, 1990; Ogasawara, 1994), and was probably blocked
from influx of warmer waters from the south, thus pro-
viding a suitable habitat for Calyptogena species.
The oldest species of Archivesica, A. sakoi Amano, Jenkins,
Ohara, and Kiel, 2014, is from the lower Miocene Shikiya
Formation in Wakayama Prefecture on the Pacific side of
southern Japan. From the late Miocene onward Archivesica
also occurs in the Japan Sea, but during the same time it is
more diverse and more widely distributed on the Pacific side,
with extant Archivesica kawamurai and three further
Archivesica species distributed from central Honshu to
southwestern Japan (Amano and Jenkins, 2011; Table 2). The
Pacific side of central Honshu to southwestemm Japan has
been influenced by warm-water currents since early Miocene
(Ogasawara, 1994) and has been situated at a subduction
zone, resulting in constant methane seepage (Amano and
Jenkins, 2011). Today, Archivesica includes at least ten
species living in the warm-water area from central Honshu to
southwestern Japan on the Pacific side (Okutani, 2017) and is
the most diversified genus among the vesicomyids.
THE NAUTILUS, Vol. 133, No. 2
ACKNOWLEDGMENTS
We thank Takuma Haga (National Museum of Nature and
Science) for his help examining the Akebiconcha uchi-
muraensis specimens, collected by Matsumoto and Hirata
(1972). We also thank Crispin Little (University of Leeds)
and Krzysztof Hryniewicz (Paleobiology Institute of
Polish Academy of Sciences) for their critical reviews that
improved the manuscript. This study was supported by a
Grant-in-aid for Scientific Research from the Japan So-
ciety for Promotion of Science (C, 17K05691, 2017-2019)
to KA and RG].
LITERATURE CITED
Amano, K. 2001. Pliocene molluscan fauna of Japan Sea bor-
derland and the ee ca as conditions. Biological
Science (Tokyo) 53: 178-184. (in Japanese)
Amano, K. 2002. Molluscan fauna from the Neogene Nodani
Formation of the western mountains of Joetsu City and its
Pe one eay Study of History of Joetsu City 8: 20-30.
(in Japanese, title translated)
Amano, K. 2003. Predatory gastropod drill holes in Upper
Miocene cold seep bivalves, Hokkaido, Japan. The \ eliger
46: 90-96.
Amano, K. 2007. The Omma-Manganji fauna and its temporal
change. Fossils (Palaeontological Society of Japan), 82:
6-12. (in Japanese with English abstract)
Amano, K. 2014. Fossil records erat evolution of chemosynthetic
bivalves. Fossils (Palaeontological Society of Japan), 96:
5-14. (in Japanese with English abstract)
Amano, kK. and R.G. Jenkins. 2011. Fossil records of extant
vesicomyid species from Japan. Venus (Journal of the Mal-
acological Society of Japan) 69: 163-176.
Amano, K. and S. Kanno. 1991. Composition and structure of
Pliocene associations in the western part of Joetsu City,
Niigata Prefecture. Fossils (The Palaeontological Society of
Japan) 51: 1-14. (in Japanese with English abstract)
Amano, k. and S. Kanno. 2005. Calyptogena (Bivalvia: Ves-
icomyidae) from Neogene strata in the Joetsu District,
Niigata Prefecture, oonimll Japan. The Veliger 47: 202-212.
Amano, K. and S. Kiel. 2007. Fossil vesicomyid bivalves from the
North Pacific region. The Veliger 49: 270-293.
Amano, K. and S. Kiel. 2010. Taononnyy and distribution of fossil
Archivesica (Bivalvia: Vesicomyidae) in Japan. The Nautilus
124: 155-165.
Amano, K. and S. Kiel. 2011. Fossil Adulomya (Vesicomyidae,
Bivalvia) from Japan. The Veliger 51: 76-90.
Amano, K. and S. Kiel. 2012. Two "Neogene vesicomyid species
(Bivalvia) from Japan. The Nautilus 126: 79-85.
Amano, K., Y. Miyajima, K. Nakagawa, M. Hamuro and T.
Hamuro. 2019. @hemosymbioue bivalves from the lower
Miocene Kurosedani Formation in Toyama Prefecture,
central Honshu, Japan. Paleontological Research 23: 178-189.
Amano, K. and A. Suzuki. 2010. Redescription of “Calyptogena”
shiretokensis Uozumi (Bivalvia: Vesicomyidae) from the
Miocene Rusha Formation on the Shiretoko Peninsula,
eastem Hokkaido, Japan. Venus (Journal of the Malaco-
logical Society of Japan) 68: 165-171.
Bernard, F.R. 1974. The genus Calyptogena in British Columbia
with a description a a new species. Venus 33: 11-22.
kK. Amano et al., 2019
Boss, K.]. 1968. New species of Vesicomyidae from the Gulf of
Darien, Caribbean Sea (Bivalvia: Mollusca). Bulletin of
Marine Science 18: 731-748.
Boss, K.J. and R.D. Tuner. 1980. The giant white clam from the
Galapagos Rift, Calyptogena magnifica species novum.
Malacologia 20: 161-194.
Chinzei, K. 1986. Faunal succession and geographical distri-
bution of Neogene molluscan faunas in Japan. Palae-
ontological Society of Japan, Special Papers 29: 17-32.
Coan, E.V., P.V. Scott, and F.R. Bemard. 2000. Bivalve Sea-
shells of Western North America. Santa Barbara Museum
of Natural History, Santa Barbara, 764 pp.
Dall, W.H. 1891. Scientific results of explorations by the U.S.
Fish Commission Steamer Albatross. XX. On some new or
interesting West American shells obtained from dredgings
of the U.S. fish commission steamer Albatross in 1888.
Proceedings of the U.S. National Museum 14: 174-191.
Dall, W.H. 1895. Scientific results of explorations by the U.S.
Fish Commission Steamer “Albatross”. XXXIV. Report
on Mollusca and Brachiopoda dredged in deep water,
chiefly near the Hawaiian Islands, with hitherto unfig-
ured species from northwest America. Proceedings of
the U.S. National Museum 17: 675-733.
Dall, W.H. 1908. Reports on the dredging operations off the
west coast of Central America to the Galapagos, to the west
coast of Mexico and in the Gulf of California, in charge of
Alexander Agassiz, carried on by the U. S. Fish Commission
steamer “Albatros” during 1891, Lieut.-Commander Z. L.
Turner, U. S. N. commanding XXXVIT. Reports on the
scientific results of the expedition to the eastern tropical
Pacific, in charge of Alexander Agassiz, carried on by the U.
S. Fish Commission steamer “Albatros” from October, 1904
to March, 1906, Lieut.-Commander L. M. Garrett, U.S. N.
commanding XIV. The Mollusca and the Brachiopoda.
Bulletin of the Museum of Comparative Zoology at Harvard
University 43: 205-487.
Dall, W.H. 1916. Diagnoses of new species of marine bivalve
mollusks from the Northwest coast of America in the
collection of the United States National Museum. Pro-
ceedings of the United States National Museum 45:
587-597.
Dall, W.H. and C.T. Simpson. 1901. The Mollusca of Porto Rico.
Bulletin of the United States, Fish and Fisheries Com-
mission 20: 351-524.
Decker, C., K. Olu, R. L. Cunha, and S$. Amaud-Haond. 2012.
Phylogeny and diversification patterns among vesicomyid bi-
valves. PLoS ONE 7: €33359. doi:10. 1371journal. pone.
0033359.
Fisher, C.R. 1990. Chemoautotrophic and methanotrophic
symbioses in marine invertebrates. Reviews in Aquatic
Sciences 2:399-436.
Grant, U.S. and H.R. Gale. 1931. Catalogue of marine Pliocene
and Pleistocene Mollusca of California and adjacent regions.
Memoir of San Diego Society of Natural History 1: 1- 1036.
Horikoshi, M. and J. Hashimoto. 1992. Two distinct growth stages
of a deep-sea, giant white clam, “Calyptogena” soyoae, and its
allied species. La mer 30: 73-82.
Huber, M. 2010. Compendium of Bivalves. A full-color guide to
the World’ marine bivalves. A status on Bivalvia after 250
years of research. Conchbooks, Hackenheim, 901 pp.
lijima, A. and R. Tada. 1990. Evolution of Tertiary sedimentary
basins of Japan in reference to opening of the Japan Sea.
Journal of the Faculty of Science, the University of Tokyo,
Section 2 22: 121-171.
Page 55
Isaji, S. 2013. Adulomya from the Miocene Aokiyama Forma-
tion, Hota Group, in the Boso Peninsula, Japan. Paleon-
tological Research 17: 196-199.
Johnson, S.B., E.M. Krykova, A. Audzijonyte, H. Sahling, and
R.C. Vrijenhoek. 2017. Phylogeny and origins of chemo-
synthetic vesicomyid clams. Sy stematics anit Biodiversity
15: 346-360.
Kanehara, K. 1937. Miocene shells from the Joban coal-field.
Bulletin of the Imperial Geological Survey of Japan 27:
1-12.
Kanno, S., K. Amano, and H. Ban. 1989. Calyptogena
(Calyptogena) pacifica Dall (Bivalvia) from the Neogene
system in the Joetsu district, Niigata prefecture. Transactions
and Proceedings of the Palaeontological Society of Japan, New
Series 153: 95-35,
Kanno, S., K. Tanaka, H. Koike, K. Narita, and T. Endo. 1998.
Adulomya uchimuraensis Kuroda (Bivalvia) from the
Miocene Bessho Formation in Shiga- mura, Nagano Pre-
fecture, Japan. Research Report of the Sivialhrnel timers
Fossil Museum 1: 17-28.
Keen, M. 1969. Family Vesicomyidae. In: C. Moore (ed.),
Treatise on Invertebrate Paleontology, Part N, vol. 2 (of 3).
University of Kansas and Geological Society of America,
Lawrence: N663-664.
Kiel, S. 2007. Status of the enigmatic fossil vesicomyid bivalve
Pleurophopsis. Acta Palacontologica Polonica 52: 639-642.
Kiel, S. and K. Amano. 2010. Oligocene and Miocene vesicomyid
bivalves from the Katalla district in southern Alaska, USA.
The Veliger 51: 76-84.
Kiel, S. and B.T. Hansen. 2015. Cenozoic methane-seep faunas
of the Caribbean Region. PLoS ONE 10: e0140788. doi:
10.1371 /journal. pone.01407S8.
Kiel, S., and M. Taviani. 2017. Chemosymbiotic bivalves from
Miocene methane-seep carbonates in Italy. Journal of
Paleontology 91:444—466.
Krylova, E.M. and H. Sahling. 2006. Recent bivalve molluscs of
the genus Calyptogena (Vesicomyidae). Journal of Mol-
luscan Studies 72: 359-395.
Krylova, E.M., H. Sahling, and R. Janssen. 2010. Abyssogena: A
new genus of the Family Vesicomyidae (Bivalvia) from
deep- water vents and seeps. Journal of Molluscan Studies
76: 107-132.
Kuroda, T. 1931. Fossil Mollusca. In: Homma, F. (ed.), Geology
of the central part of Shinano, part 4. Kokon Shoin, Toky 0:
1-90 (in Japanese)
Kuroda, T. 1943. Akebiconcha, a new pelecypod genus. Venus
13: 14-18. (in Japanese with English description)
Matsumoto, E. and M. Hirata. 1972. Akebiconcha uchimur-
aensis (Kuroda) from the Oligocene formations of the
Shimanto terrain. Bulletin of the National Science Mu-
seum, Tokyo 15: 753-760.
Métivier, B., T. Okutani, and S. Ohta. 1986. Calyptogena
(Ectenagena) phaseoliformis n. sp., an unusual vesicomyid
bivalve ailected by the submersible Nautile from aby: ssal
depths of the Japan and Kurile Trenches. Venus 45:
161-168.
Miyajima, Y., T. Nobuhara, and H. Koike. 2017. Taxonomic
reexamination of three vesicomyid species (Bivalvia) from
the middle Miocene Bessho Formation in Nagano Pre-
fecture, central Japan, with notes on vesicomyid’ diversity.
The Nautilus 131: 51-56.
Nevesskaja, L.A., S.V. Popov, LA. Goncharova, A.V. Guzhov,
B.T.Janin, 1.V. Plubotko, A.S. Biakov, and V.A. Gavrilova.
2013. Phanerozoic Bivalvia of Russia and surrounding
Page 56
countries. Transactions of the Paleontological Institute 294:
1-524. (in Russian)
Ninomiya, T. 2011. Chemosynthetic fossil molluscan faunas
from the Neogene Taishu Group, distributed in Tsushima
Islands, Nagasaki Prefecture, the southwest Japan. Mem-
oirs of the Faculty of Science, Kyushu University, Series D
(Earth and Planetary Science) 32: 11-26.
Ninomiya, T., S. Shimoyama, kK.A. Watanabe, K. Horie, D.
J. Dunkley, and K. Shiraishi. 2014. Age of the Taishu
Group, southwestern Japan, and implications for the origin
and evolution of the Japan Sea. Island Arc 23: 206-220.
Ogasawara, K. 1994. Neogene paleogeography and marine cli-
mate of the Japanese Islands based on shallow-marine
molluscs. Palaeogeography, Palaeoclimatology, Palae-
oecology 108: 335-351,
Okutani, T. 1966. Identity of Calyptogena and Akebiconcha
(Bivalvia, Cyprinidae). Venus 24: 297-303. (in Japanese
with English summary)
Okutani, T. 2000. Vesicomyidae. In: Okutani, T. (ed.), Marine
Mollusks in Japan. Tokai Univ ersity Press, Tokyo: 996-999.
(in Japanese and English)
Okautani, T. 2017. Vesicomyidae. In: Okutani, T. (ed.), Marine
Mollusks in Japan, 2°4 edition. Tokai Univ ersity Press,
Tokyo: 571-574 (Atlas), 1232-1234 (Text). (in Japanese and
English)
Okutani, T., K. Fujikura, and J. Hashimoto. 1993. Another new
species of Calyptogena (Bivalvia: Vesicomyidae) from
bathyal depth in Suruga Bay, Japan. Venus 52: 121-126.
Olsson, A.A. 1931. Contabnttons: to the Tertiary paleontology of
northern Peru: Part 4, The Peruvian Oligocene. Bulletins of
American Paleontology 17:97-264.
Olsson, A. A. 1942. Tertiary deposits of northwestern South
America and Panama. Pages 231-287 in 8th American
Science Congress held in Washington, 1940, vol. 4, Geo-
logical Sciences. Department of State.
Otatume, K. 1942. On the occurrence of fossil Calyptogena from
the Ishikari Oil-Field, Hokkaido. Journal of the Geological
Society of Japan 49: 435-437.
Otuka, Y. 1937. Occurrence of Calyptogena pacifica Dall from
Wakimoto on Oga Peninsula. Journal of the Geological
Society of Japan 44: 231. (in Japanese)
THE NAUTILUS, Vol. 133, No. 2
Sato, H. 1994. The relationship between late Cenozoic tectonic
events and stress field and basin development in northeast
Japan. Journal of Geophysical Research 99: 22261-22274.
Suyari, K., Y. Kuwano, and T. Yamazaki. 1989. Distribution of
lithofacies and geological ages in the Shimanto South
Subbelt in eastern ee Journal of Science, University
of Tokushima 22: 33-57. (in Japanese with English abstract)
Suzuki, A. 2007. III-3 Molluscan Fossils. In: Center of Sapporo
City Museum (ed.), Synthetic report on megafossils from
Sapporo City. —Elucidation of Sapporo Sea Cars and its age-.
Sapporo City: 69-75. (in Japanese, title translated)
Suzuki, K. 1941. Three new species of non-marine shells from
the Tertiary formations of Hokkaido and Karahuto. Japanese
Journal of Geology and Geography 18: 53-58.
Taira, A., M. Tachi, M. Olkaranraa, and J. Katto. 1980. The
Geology of the Shimanto Belt in Kochi Prefecture, Shikoku,
Japan. Selected papers in Honor of Prof. Jiro Katto, Ge-
ology and Paleontology of the Shimanto Belt. 319-389. (in
Japanese with English Abstract)
Takeda, H. 1953. The ‘Pononet Formation (Oligocene Tertiary)
of Hokkaido and South Sakhalin and its fossil fauna. Studies
on Coal Geology, the Hokkaido Association of Coal Mining
Technologists 3: 1-103.
Taylor, J.D. and E.A. Glover, 2010. Chemosymbiotic bivalves.
In: S. Kiel (ed.), The Vent and Seep Biota. Topics in
Geobiology 33: 107-136.
Tiba, R. 1972. Some interesting shells from the Sea of Okhotsk and
Bering Sea. Venus 30: 153-156. (in Japanese with English
abstract)
Tsuji, T. Y. Masui, A. Waseda, Y. Inoue, K. Kurita, and K. Kai.
1991. The Onnagawa Formation in the vicinity of the
Yashima Town, Akita Prefecture, northern Japan—with
special reference to the lithologic units, the depositional
environments and their relation to source rock character-
istics. Research report of the Japan Petroleum Exploration
Technology Research Institute 7: 45-99. (in Japanese with
English Alster’)
Van W alld, K. 1919. Remarks on some new species from
Trinidad. Bulletins of American Paleontology 8: 19-27.
Woodring, W.P. 1938. Lower Pliocene mollusks aratil echinoids from
the Los Angeles basin, California, and their inferred enyi-
ronment. U.S. Geological Survey Professional Paper 190: 1-67.
THE NAUTILUS 133(2):57-66, 2019
Page 57
Two warm-water species of Trochoidea (Gastropoda) from Pliocene
deposits on the Japan Sea side of Honshu, Japan, with remarks on
the influence of the onset of Northern Hemisphere glaciation
Kazutaka Amano
Department of Geoscience
Joetsu University of Education
Joetsu 943-8512, JAPAN
amano@juen.ac.jp
ABSTRACT
Two warm-water trochoidean gastropods are studied. One of
them, Monodonta joetsuensis new species, is the first Pliocene
record of this genus in Japan. Another, Pomaulax omorii (Shibata,
1957), is distributed widely along the Japan Sea side of Honshu.
In the Japan Sea borderland, both species are confined to late
Pliocene deposits and became extinct as a result of cooling at the
onset of Northern Hemisphere glaciation near the end of the
Pliocene (2.75 Ma). It has become clear that thirteen shallow-
water suspension-feeding bivalves, including Miocene relict
forms, and eleven grazing or predatory/scavenging gastropods
that mostly lived in warm shallow-water disappeared from the
Japan Sea during this cooling event.
INTRODUCTION
Many species of trochoidean gastropods live on rocky
bottoms (Hickman and McLean, 1990). Fossils of these
species are not plentiful, and are usually collected from
sandy sediments or turbidites, deeper than their original
living habitats. Two extinct species of Trochoidea have
been recovered from the upper Pliocene desposits on the
Japan Sea side of Honshu. One of them, a small trochid
gastropod, is a new species of Monodonta, and the other,
is a large turbinid gastropod Pomaulax omorii (Shibata,
1957). Pomaulax omorii was originally described by
Shibata (1957) as Astraea (Pachypoma) omorii from
the lower Pliocene Ochiai Formation in Kanagawa Pre-
fecture, on the Pacific side of Honshu. On the Japan Sea
side of Honshu, this species was illustrated as Astralium
(Distellifer) aff. rhodostoma (Lamarck, 1822) by Amano
et al. (2000b) from the upper Pliocene Tentokuji For-
mation. Fortunately, I collected some well-preserved
specimens of Pomaulax omorii from the upper Pliocene
formations in Niigata.
I describe the new species of Monodonta, redescribe
the features of Pomaulax omorii from the Japan Sea
borderland and discuss their paleogeographic significance.
Accordingly, I also discuss the influence of the onset of
major Northem Hemisphere glaciation to the molluscan
fauna in the Japan Sea borderland.
MATERIALS AND METHODS
Monodonta joetsuensis new species was recovered from
an alternation of fine-graind sandstone, yielding pebbles
and plant fragments, and dark gray mudstone of the
Nadachi Formation, 220 m upstream from a tributary
0.8 km upstream from the Fujikake-dani River in Joetsu
City, Niigata Prefecture (Figure 1, Loc. 4). From this
locality, 18 species of gastropods and 22 bivalves co-
occurred, crowded together (Table 1). Many shallow-
water species were carried into deep water by turbidity
currents—most bivalves, even the deep-sea bivalve
Calyptogena pacifica Dall, 1891 are disarticulated at this
site. Two warm-water species, Thais clavigera (Kiister,
1860) and Veremolpa micra (Pilsbry, 1904), are associated
with many cold-water or endemic extinct species. Cal-
careous nannofossils from this locality were identified by
Tokiyuki Sato (Akita University), who indicated that their
ages fall within the lower to middle part of zone NN16
(Martini, 1971; 3.66-2.75 Ma).
Seventeen specimens of Pomaulax omorii were ob-
tained from pebble-bearing sandstones of the lower part
of the Tentokuji Formation at the large cliff along the
Sannai River in Kami-Sannai, Akita City, Akita Prefecture
(Figure 1, Loc. 1). This locality corresponds to the fossil
locality described by Shimamoto and Koike (1986) and the
Loc. 2 of Amano et al. (2000b). The age of the Tentokuji
Formation was assigned to the lower to middle zone
NN16 (3.85-2.75Ma) via calcareous nannofossils (Amano
et al., 2000). Some warm-water species such as Erronea
sp. and Oliva mustelina Lamarck, 1811 were collected
with many cold-water mollusks (Amano et al., 2000b:;
Table 2). One specimen of Pomaulax omorii was also
collected from a calcareous concretion yielding plant
debris included in siltstone of the upper part of the Araya
Page 58
THE NAUTILUS, Vol. 133, No. 2
130° E 140° (\ Okhotsk
vo Sea
\
= cc
Vy Hokkaido_
} i} SU WY
— Ww - Pasa
{oO
vo ‘ ~
Vara
i Bae
40°N = re ee Cet Fy) — See
( a 11 Bonshu
52 fe ot
ee LS
e2
< U4 &
7
—— Ret a O? a ay
ce Se ¢
6 GEE fb —~ Qa?
ee AON =
yD) a
by
i
Wy 0 200km
920
30° = —— --—--——- OF ~ - a + ———
Figure 1. Localities of fossils.
Formation on the bank of the Maekawa River, 600 m
south of Kiyamazawa, Nagaoka City, Niigata precomme
(Figure 1, Loc. 2). The age of the upper part of this
formation was assigned to the late Pliocene based on
foraminifers by Kobayashi et al. (1991). Other than
Nemocardium samarangae (Makiyama, 1934), the asso-
ciated fauna consists of cold-water or extinct endemic
species (Table 1). One more well-preserved specimen of
Pomalulax omorii was collected from sandy siltstone of the
Shitoka Formation, 550m upstream in the Shitoka River
in Minami Uonuma City, Niigata Prefecture (Figure 1,
Loc. 3). The age of the Shitoka Formation was assioned to
the late Pliocene based on calcareous nerimnoftnacl by
Amano et al. (2009). Many cold-water species and one
warm-water species, Nemocardium samarangae (Makiyama,
1934), are associated with this specimen (Table 2)
I have followed the arrangement in Bouchet et al.
(2017). All pictures were t neler with the specimens cov-
ered with ammonium chloride. All specimens are de-
posited at the National Museum of Nature and Science,
Tsukuba (NMNS PM).
SYSTEMATIC PALEONTOLOGY
Class Gastropoda Cuvier, 1797
Subclass Vetigastropoda Salvini-Plawen, 1980
Order Trochida Cox and Knight, 1960
Superfamily Trochoidea Rafinesque, 1815
Family Trochidae Rafinesque, 1815
Subfamily Monodontinae Gray, 1857
Genus Monodonta Lamarck, 1799
Type Species: Trochus labio Linnaeus,
monotypy.
1758 by
Remarks: According to Williams et al. (2010), the subfamily
Monodontinae includes three genera: Monodonta Lamar ck,
1799, Austrocochlea P. msde 1885 and Diloma Philippi,
1845. Based on molecular and morphological data, Aus-
trocochlea seems to be very close to Monodonta (Donald
et al., 2005; Willams et al., 2010). However, Austrocochlea
has narrower spiral cords without any axial grooves, a weak
tooth on the inner lip, and very weak ‘crenulations inside the
outer lip. Diloma differs from Monodonta in having no tooth
on the inner lip and no crenulations inside the nattnere thin
outer lip.
Monodonta joetsuensis new species
(Figures 2-5)
Diagnosis: Small Monodonta with 4.25 whorls and rather
low spire, sculpture of about 20 spiral cords on last whorl,
separated by nearly equal interspaces. Inner lip with
prominent basal tooth and eight denticles inside outer lip.
Description: Shell small (9.5 mm in height, 10.4 mm in
diameter), rather thick, turbiniform nit relatively low
spire; protoconch decolated: teleoconch with 4.25 whorls:
suture deep. Surface sculptured with growth lines and
spiral cords; growth lines distinct, particularly near ap-
erture, and oblique posteriorly; spiral cords separated by
nearly equal interspaces, seven on penultimate whorl,
twenty on last whorl, cords color alternating light and dark
gray; area below substural wide cord concave, sculptured
with three fine cords. Aperture circular; columella arched,
with prominent basal tooth, deeply notched below; in-
terior of outer lip crenulated by eight denticles.
Holotype: NMNS PM 28267 (9.5 mm in height, 10.4 mm
in diameter).
Figures 2-5.
Monodonta joetsuensis new species, holotype,
NMNS PM 28267) from the Nadachi Formation. 2. Apertural
view. 3. Abapertural view. 4. Apical view. 5. Basal view. Scale
bar = 5 mm.
K. Amano, 2019
Table 1. Molluscan fossils associated with Monodonta
joetsuensis new species from the upper Pliocene Nadachi
Formation (Loc. 4 in Figure 1). * warm-water species.
Species
Notoacmaea sp.
Cocculina japonica Dall
Homalopoma noctrum (Gould)
Monodonta joetsuensis new species
Umbinium (Suchium) akitanum Suzuki
Omphalius aff. rusticus (Gmelin)
Bittium sp.
Littorina brevicula (Philippi)
Turritella (Naohaustator) saishuensis motidukii Otuka
Euspira pallida (Broderip and Sowerby 1)
Neverita (Glosaulax) vesicalis (Philippi)*
Cryptonatica clausa (Boroderip and Sowerby 1)
C. janthostoma (Deshayes)
Thais clavigera (Kiister)*
Lirabuccinum fuscolabiatum (Smith)
Buccinum sp.
Mitrella bicincta (Gould)
Reticunassa acutidentata (Smith)
Acila (Truncacila) nakazimai Otuka
Leionucula niponica (Smith)
Portlandia (Portlandella) toyamaensis (Kuroda)
Nuculana (Nuculana) onoyamai Otuka
Agpadlanea (Scapharca) ommaensis Otuka
Porterius dalli (Smith)
Glycymeris (Glycymeris) yessoensis (Sowerby III)
Chlamys (Chlamys) cosibensis (Yokoyama)
Anomia chinensis (Philippi)
Felaniella usta (Gould)
Astarte hakodatensis Yokoyama
Tridonta borealis (Schumacher)
“Dinocardium” angustum (Y okoyama)
Macoma (Marconna) calcarea (Gmelin)
Calyptogena pacifica Dall
Pseudamiantis tauyensis (Conrad)
Humilaria perlaminosa (Conrad)
Securella sp.
Veremolpa micra (Pilsbry)*
Protothaca tateiwai (Makiy. ama)
Anisocorbula venusta Gould
Myadora fluctuosa Gould
Type Locality: Small outcrop at 220 m upstream from a
tributary 0.8 km upstream from the Fujikake-dani River in
Joetsu City, Niigata Prefecture, Japan.
Remarks: This new species can be safely assigned to
Monodonta, not to Austrocochlea nor Diloma, Tresemee it
has a strong basal tooth and strong denticles inside the
outer lip. The present species is most util to the recent
Monodonta australis Lamarck, 1822, from around the
Ogasawara Islands, Okinawa Islands, Mariana Islands,
iSlexcvett, and Australia (Higo et al., 1999) in having a similar
number of cords (6-12 on the penultimate whorl and
15-22 on the last whorl) and a similar number of cren-
ulations inside the outer lip (6-13). Monodonta joetsuensis
new species, however, has narrow spiral cords, as in some
species of Austrocochlea. In contrast, M. australis has flat
Page 59
spiral cords separated by deep grooves and crossed by
vertical grooves near the aperture.
Another recent species, Monodonta canalifera
Lamarck, 1822, differs from the present species in having
fewer (14-18), lower spiral cords.
Etymology: Named after the city name of locality.
Distribution: Late Pliocene, Nadachi Formation in
Niigata Prefecture.
Family Turbinidae Rafinesque, 1815
Subfamily Turbininae Rafinesque, 1815
Genus Pomaulax Gray, 1850
Type Species: Trochus japonicus Dunker, 1844 by
subsequent designation.
Remarks: Astraea Roding, 1798 previously encompassed
species now included in distinct genera such as Astralium
Link, 1807, Lithopoma Gray, 1850, Megastraea MacLean,
1970, Pomaulax Gray, 1850, and others (Gee Alf and Kreipl,
2011). This usage is now considered outdated because
many paraphy latte clades were included (based on mo-
lecular data by Williams (2007, 2012) and Williams et al
(2008)). The type species, Astraea heliotropium (Martyn,
1784) is the only living species left in this genus. It is re-
stricted to New TeslenGl Astraea can coal be distin-
guished from Pomaulax by presence of well- inflated whorls,
a very wide umbilicus and many prominent, wide spines at
the periphery. Pachypoma was established by Gray (1850).
based on Trochus caelatus Gmelin, 1791 (see Bouchet,
2011). Thus, Pachypoma is a junior synonym of Lithopoma.
Megastraea, based on Astraea undosa (Wood, 1828),
most “lossy resembles Pomaulax. Megastraea differs
from Pomaulax by having an operculum aida three spiny
ridges, and one or two wavy spiral ridges on and above the
periphery (see also Alf and Kreipl, 2011). However, recent
molecular work shows a close relationship between
Pomaulax and Megastraea (Willams, 2007: 2012).
Lithopoma, from the western Atlantic, is another genus
similar to Pomaulax. Lithopoma can be Giaerttatnattedl | from
Pomaulax in generally having a smaller shell and a periphery
with strong otal ridges or atin hollow spines (see also Alf
and Kreipl, 2011). Mioleonllar data shows a close relationship
with Lithopoma and Pomaulax (Willams, 2007, 2012).
Pomaulax omorii (Shibata, 1957)
(Figures 6-13, 15, 17)
Astraea (Pachypoma) omorii Shibata, 1957, p. 24, pl. 4,
figs. 2a—c.
? Omphalius pfeifferi cf. carpenteri (Dunker), Ogasawara
et al., 1986, pl. 26, figs. 7a—c.
Astralium (Distellifer) ) aff. rhodostoma (Lamarck), Amano
et al., 2000b, pl. 1, figs.17a, b.
Astraea omorii Shibata, Meartsmalstiine et al.,
1, 2; Shiba et al., 2013, figs. 4.3, 4.4.
?non Pomaulax omorii (Shibata), Amano et al., 2011, figs.
Doll, Bods,
2003, pl. 4, fig
Page 60
Table 2
(Shibata) from Loc. 1 (Tentokuji For nee a)) 2
3 (Shitoka Formation) in Figure 1.
mation), <
species.
Species name
Puncturella nobilis A. Adams
Lepeta cf. lima Dall
Niveotectura pallida (Gould)
Minolia sp.
Turcica sp.
Littorina sp.
Turritella (Neohaustator) saishuensis
saishuensis Yokoyama
T. (N.) nipponica (Yokoyama)
Erronea sp.*
Neverita (Glossaulax) cf. vesicalis
Philippi)
Cryptonatica janthostomoides
(Kuroda and Habe)
C. sp.
Fusitriton izumozakiensis Amano
F. aff. oregonensis (Redfield)
Mohnia yanamii (Yokoyama)
Neptunea (Neptunea) eos (Kuroda)
N. (N.) insularis (Dall)
N. sp.
Buccinum cf. unuscarinatum Tiba
B. sp.
Oliva mustelina Lamarck*
Fulgoraria prevostiana (Crosse)
F. masudae Hayasaka
Propebela sp.
Antiplanes contraria (Yokoyama)
Rectiplanes sanctiioannis (Smith)
Conidae gen. et sp. indet.*
Acila (Acila) divaricata (Hinds)
A. (Truncacila) insignis (Gould)
A. (T.) aff. castrensis (Hinds)
Leionucula cf. niponica (Smith)
Malletia inermis Yokoyama
Nuculana (Nuculana) pernula (Miller)
Yoldia (Cnesterium) notabilis Yokoyama
Y. sp.
Portlandia (Portlandella) japonica
(Adams and Reeve)
P. (P.) toyamaensis (Kuroda)
Arca boucardi Jousseaume
Anadara (Anadara) amicula (Yokoyama)
Porterius dalli (Smith)
Glycymeris (Glycymeris)
albolineata (Lischke)
G. (G.) nipponica (Yokoyama)
Limopsis tajimae Sowerby
L. cf. tajimae Sowerby
L. oblonga (A. Adams)
Verwouline laevigatus (Gray)
Megacrenella ealbiadbiainn (Dall)
Chlamys (Chlamys) cosibensis (Yokoyama)
C. (C.) tamurae Masuda and Sawada
Mizuhopecten yokoyamae (Masuda)
M. sp.
Molluscan fossils associated with Pomaulax omorii
(Araya For-
++++4¢4¢4]
++
++4
+ + t++++4]4
+++4
+++
* warm-water
+
+
+
+
+
oe
+
+
+
+
+
oe
+
fk
+
+
+
a
+
+
+
(Continued)
THE NAUTILUS, Vol. 133, No. 2
Table 2. (Continued)
bo
(oe)
Species name Loc.
Yabepecten tokunagai (Yokoyama)
Cycladicama cumingii (Hanley)*
Felaniella usta (Gould)
Cardita leana Dunker
C. cf. leana Dunker
Megacardita sp.
Cyclocardia myogadaniensis (Itoigawa)
Tridonta borealis Schumacher
Nemocardium samarangae (Makiyama)*
Clinocardium (Glenn
ciliatum (Fabricius)
Serripes groenlandicus (Bruguiére) oF
Spisula (Mactromeris) voyi Gabb
S. (M.) grayana (Schrenck)
Cadella lubrica (Gould)
Macoma (Macoma) calcarea (Gmelin)
M. (M.) nipponica (Tokunaga)
Securella cf. stimpsoni ( (Gould)
S. chitaniana (Yokoyama)
Phacosoma tomikawensis (Takagi)
Pseudamiantis cf. tauyensis (Yokoy. ama)
Pandora wardiana (A. Adams) -
Entodesma ane Yokoyama
*
t+ttt+t+++¢4+44+
+
+
++t+++4+444
+
Type Locality: Bank of Hayato River near Ochiai,
Kiyokawa Village, Kanagawa Prefecture: Ochiai Forma-
tion, Pacific side of central Honshu, Japan.
Original Description: “Shell trochoid, depressed, large,
conic-pyramidal, solid, thick, broader than high: surface of
whorls flat, declined, sculptured by oblique axial plicae,
the plicae weaker on the upper but stronger on the lower
where they become somewhat crossed: peripheral margin
angulated: on the lower surface sculptured by nine spiral
cords, with alternating weak knotted and smooth cords:
the columella base somewhat elevated, umbilical area
somewhat concave: aperture squarely rounded in outline,
outer lip sharply inclined, basal lip thin: columella
somewhat concave, smooth, tooth-like swelling on lower
part.” (Shibata, 1957, p. 24).
Material Examined: Two well-preserved specimens
(NMNS PM 28268, 28269) from the Tentokuji Forma-
tion, one specimen (NMNS PM 28270) from the Araya
Formation, and one well-preserved specimen (NMNS
PM 28271) from the Shitoka Formation are examined.
Description of the Specimens from the Japan Sea
Side of Honshu: Shell medium in size, attaining
51.8 mm-+ in height, 56.8 mm in diameter, rather thick,
trochiform with high spire; protoconch decollated; tele-
oconch of more Shera five whorls; suture overlapped by
periphery of previous whorl. Surface of each whorl rather
flat, sculptured by growth lines, axial ribs, and spiral cords;
growth lines distinct and very oblique; axial ribs narrow,
low, oblique in opposite direction to growth lines, 37 to 42
on penultimate whorl, 35 to 37 on last whorl, becoming
more distinct near upper suture; spiral cords 6 to 13 on last
K. Amano, 2019 Page 6]
Figures 6-17. Fossil Pomaulax species from Japan. 6-13, 15, 17. Pamaulax omorii (Shibata). 6, 9. NMNS PM 28268. 6. Abapertural
view. 9. Apical view; Tentokuji Formation. 7, 11. NMNS PM 28270. 7. Apertural view. 11. Apical view; Araya Formation. 8, 10, 12.
NMNS PM 28271. 8. Apertural view. 10. Basal view. 12. Abapertural view; Shitoka Formation. 13, 15, 17. NMNS PM 28269. 13. Basal
view. 15. Apertural view. 17. Abapertural view. 14, 16. Pomaulax tyosiensis (Ozaki), syntype NMNS PM 4257. 14. Apertural view. 16.
Basal view; Na-Arai Formation. Scale bar = 10 mm.
Page 62
THE NAUTILUS, Vol. 133, No. 2
whorl, forming granulations at intersection with axial ribs.
pany with undulating carina bearing 14-15 nodes.
Base flat, sculptured contin five to nine spiral cords. Ap-
erture oblique and squarely rounded in outline; umblical
region covered by rather thick callus.
Remarks: The holotype specimen of Pomaulax omorii
(Shibata, 1957) attains ca. 61 mm in height and 65mm
in diameter. The topotype specimens illustrated by
Matsushima et al. (2003) differ slightly from the holotype
by their taller trochiform shape. Matsushima et al. (2003)
also pointed out two forms having higher and lower spires.
Amano et al. (2000b) illustrated Asti valium (Distellifer)
aff. rhodostoma (Lamarck, 1822) from the upper Pliocene
Tentokuji Formation at Kami-Sannai in Akita City, Akita
Prefecture. However, from its size (max. 51.8 mm in shell
height, 56.8 mm in diameter) and shell sculpture (37 to 42
axial ribs on the penultimate whorl, 14 to 15 nodes at the
periphery, and five to nine sprial cords on the base), this
species can be identified as Pomaulax omorii.
From the same locality as Amano et al. (2000b), Ogasawara
et al. (1986) illustrated a poorly preserved specimen as
Omphalius pfeifferi_ cf. carpenteri (Dunker, 1859). Judging
from its size, shape and seven sprial cords on the base, this
specimen probably can be identified with Pomaulax omorii.
Amano et al. (2011) illustrated two specimens as
Pomaulax omorii from the lowermost Sasaoka Formation
at the upstream of Ogurosawagawa River. However, one
specimen in their fig. *. 14 Thexs a noded spiral cord above
the suture as in species of Bolma Risso, 1826.
Pomaulax omorii is closely allied to P. tyosiensis (Ozaki,
1954) from the Pliocene Na-arai Formation at Inuwaka in
Choshi City, Chiba Prefecture. In the description of
P. tyosiensis, Ozaki (1954) assigned Nat. Sci. Mus., coll.
cat. no. 4257 as the holotype. However, as three speci-
mens are included in the container under no. 4257, they
should be considered as syntypes. I have examined all
three specimens. Pomaulax tyosiensis (Figures 14, 16)
differs from P. omorii in its two distinct rows of peripheral
nodes like those of Megastraea turbanica (Dall, 1910),
more fine and numerous axial ribs (132 on the last whorl),
and more numerous peripheral nodes (13-16).
The Recent species Pomaulax japonicus (Dunker in
Philippi, 1844) can be easily distinguished from P. omorii
easily by having a larger and lower Shell as well as no spiral
cords on the last Ww srrovdl
Distribution: On the Pacific side of Honshu, upper
Miocene Osozawa Sandstone Member of litomi Forma-
tion in Yamanashi Prefecture (Shiba et al., 2014) and
lower Pliocene Ochiai Formation in Kanagawa Prefecture.
1 . - S .
On the Japan Sea side of Honshu, upper Pliocene Ten-
tokuji Formation in Akita Prefecture, Araya and Shitoka
formations in Niigata Prefecture.
DISCUSSION
Fossils of two species of Monodonta have been known
from Japan: M. minuta Itoigawa and Nishikawa, 1976 and
M. labio (Linnaeus, 1758). The former species has been
proposed from the lower Miocene Lower Member of the
Bihoku Group in Okayama Prefecture (Itoigawa and
Nishikawa, 1976). Nakagawa (2009) reported this species
as M. kanzakii from the lower Miocene Kohnoura Shale
Member of Shimo Formation in Fukui Prefecture. Later,
Nakagawa (2018) synonymized M. kanzakii with M.
minuta. The Recent species M. labio has been recorded
from the middle Pleistocene Sakishima Formation in
Mie Prefecture (Itoigawa and Ogawa, 1973), an upper
Pleistocene deposit at Akaura in Ishikawa Prefecture
(Matsuura, 1977), the Holocene Numa Coral Bed in Chiba
Prefecture (Yokoyama, 1924), Holocene deposits at
Hachigasaki and Onogi in Ishikawa Prefecture (Matsuura,
1985) ‘andl the Holocene Takahama shell bed in Fukui
(Nakagawa et al., 1993). All these fossils are associated with
warm-water faunas. In conclusion, Monodonta joetsuensis
new species is the first record of Monodonta from the
Pliocene in Japan. As mentioned above, two warm-water
species are associated with Monodonta joetsuensis new
species. The warm-water Tsushima current has flowed into
the semi-enclosed Japan Sea for the past 4 Ma (Amano
et al., 2008; Gallagher et al., 2015). Moreover, the tem-
perature of this thin, in-flowing current in the late Pliocene
was 3 to 4 °C higher than that oe today (Amano et al., 2008;
Amano, 2019). Monodonta joetsuensis new species related
to M. australis possibly appeared along with the inflow of
the warm-water current during the late Pliocene.
On the other hand, Darinenlen omorii appeared in the
late Miocene to early Pliocene warm-water “Zushi-Ochiai
type fauna” (Chinzei and Matsushima, 1987) or the “Zushi
fauna” (Ozawa and Tomida, 1992) on the Pacific side of
central Honshu. In the late Pliocene, this species invaded
the Japan Sea to what is now the Akita Prefecture via the
warm-water current. The occurrence of this species from
the Araya and Shitoka formations seems to be nearly
authochtonous. Judging from the habitat depth of living
specimens of the associated recent species, the paleo-
depth of both formations can be estimated as 100-200 m,
which is slightly deeper than the Recent Japanese species,
Pomaulax japonicus (0-100 m; Higo et al., 1999). This
estimated paleodepth of Pomaulax omorii explains its co-
occurrence with some bathyal mollucs from conglomer-
ates of the Ochiai, Iitomi, and Tentokuji formations.
On the Atlantic side of North America and Europe,
severe extinction of mollucan species resulted from the
cooling event at the onset of major Northern Hemisphere
Glaciation (NHG) around the end of the Pliocene (e.g.
Stanley and Campbell, 1981; Stanley, 1986; Vermeij,
1991; Vermeij et al., 2008). The end- Pliocene extinction
event was also recognized in corals and vertebrates
(Woesik et al., 2012: iEmento et al., 2017). When Sato
and Kameo (1996) noticed the drastic change of nan-
nofossils and increase of ice rafted debris (IRD) in the
core from North Atlantic Ocean, they named this datum
plane as Datum A (2.78 Ma, changed to 2.75 Ma by Sato
et al., 2002) indicating the onset alk glaciation and traced it
in the land deposits in Akita Prettoalinne, Bailey et al.
(2013) considered the onset of NHG at 2.72 Ma (MIS
K. Amano, 2019
Table 3.
Miocene; * warm-water species.
Page 63
Molluscan species extinct by the end of the Pliocene in the Japan Sea borderland. ~ temperate relict species from the
Species
Formation
Reference
Chlamys ingeniosa tanakai (Akiy. ama)?
C. insolita (Yokoyama)
C. lioica shigaramiensis Amano and Karasawa
Mizuhopecten yamasakii ( (Yokoyama)°
M. tryblium (Yokoyama)~
M. naganoensis (Masuda)
Nec sgenella hokkaidoensis (Nomura)°
Rarpetonatia ausiensis (Ilyina)~
Ogikubo F.
Ogikubo F.
Ogikubo F.
Joshita F.
Joshita F.
Protothaca tateiwai (Makiyama)
Humilaria perlaminosa (Conrad)
Meretrix spp.
Thracia higashinodonoensis Oinomikado”
Pholadomya kawadai Omori?
Monodonta joetsuensis new species*
Pomaulax omorii ( Shibata)*
Vermeijia japonica Amano
Glossaulax didyma coticazae (Makiyama)~
Tentokuji F.
Nadachi F.
Joshita F.
Ranella yasumurai Amano Kuwae F.
Chicoreus totomiensis (Makiyama)* Kuwae F.
Babylonia elata (Yokoyama)* Mita F.
Buccinum sinanoense Makiyama
B. shibatense Amano and Watanabe
Cole satoi Amano*
Scalptia kurodai (Makiyama)*
Joshita F.
Kuwae F.
Tentokuji F.
Tentokuji F.
Arakurayama F.,
Ogikubo F., Mita F.
Joshita F., Mita F., Nadachi F.
Joshita F., Tentokuji F., Mita F.,
Nadachi F.
Kuwae F., Nadachi F.
Takafu F. , Sarumaru F.
Ogikubo F., Sasaoka F.
Tentokuji F., Araya F., Shitoka F.
Sasaoka F., Kuwae F.
Ogikubo F. Amano (1994)
Amano (1994)
Amano and Karasawa (1986)
Amano (2001)
Amano (2001), Amano et al.
Amano and Sato (1995)
Amano and Sato (1995)
Amano and Sato (1995), Amano et al.
(2008), this study
Amano and Sato (1995), Amano et al.
(2000b, 2008), this tudy
Amano et al. (2000a), this study
Nagamori and Yoshikawa (2019)
Amano (1995), Amano et al. (2011)
Amano et al. (2011)
this study
this study
Amano (2019)
Amano and Sato (1995)
Amano (1997)
Amano et al. (2000a)
Amano et al. (2008)
Amano and Watanabe (2001)
Amano and Watanabe (2001)
Amano (2019)
Amano et al. (2000b)
(2008)
G6), based on the abundant IRD deposition in the Nordic
Seas and subpolar North Atlantic Ocean. Many cold-water
species living now in Hokkaido migrated southward to
central Fone and some Mites: -type relict mollusks
sulteres from extinction by the cooling event around
2.75Ma (Amano, 2001, 2007, 2019: Aeaert et al., 2011).
Ako, Pacific-type deep-water radiolarians disappeared
from the Japan Sea around 2.75 Ma because of the in-
termittent development of low oxygen conditions in deep
water (Itaki, 2016).
The grazers Monodonta joetsuensis new species and
Pomaulax omorii also disappeared at the end of the Pli-
ocene, probably as a result of the cooling event of the
NHG. The species that probably became extinct at Datum
A in the Japan Sea are shallow-water dwellers (Table 3).
They include eight relict bivalves and the naticid Neverita
(Glossaurax) didyma coticazae (Makiyama, 1926) which
are temperate-water species that survived from the
Miocene, and many warm-water gastropods. Also, it is
interesting to note that the extinct species consist of
suspension-feeding bivalves and grazing and predatory/
scavenging gastropods. When Todd et al. (2002) examined
the endl Bikowene extinction of mollusks in the Carribean
region, they found that predatory gastropods and sus-
pension feeding bivalves declined in abundance. This
suggests that the nutrient supply changed by the cooling
event of the NHG caused the faunal change even in the
semi-enclosed Japan Sea.
ACKNOWLEDGMENTS
T acknowledge Alan Beu (GNS Science, New Zealand) for
reviewing the manuscript before submission to The
Nautilus. I also thank two reviewers, Claude Vilvens
(Muséum national d’ Histoire naturelle, Paris) and Sven N.
Nielsen (Universidad Austral de Chile) for their review
and useful comments. I also express many thanks to
Hiroshi Saito, a? Hasegawa, and Takuma Haga
(National Museum of Nature and Science, Tsukuba, Ja-
pan) who kindly showed me the Recent and fossil spec-
imens, and Tokiuki Sato (Akita Univ ersity) for information
on the age of Loc. 4 using calcareous nannofossils. This
study was supported by a Grant-in-aid for Scientific Re-
search from the Japan Society for Promotion of Science
(C, 17K05691, 2017-2019).
LITERATURE CITED
Alf, A. and K. Kreipl. 2011. A Conchological Iconography. The
family Turbinidae. Subfamily Turbininae Rafinesque,
1815 & Prisogasterinae Hickman & McLean, 1990.
Conchbooks, Hackenheim, 82 pp.
Amano, K. 1994. Diversity of Chlamys (Bivalvia) from the Plio-
cene Ogikubo Formation, Central Japan. Transactions and
Proceedings of the Palaeontological Society of Japan, New
Series 171: 237-248.
Page 64
THE NAUTILUS, Vol. 133, No. 2
Amano, K. 1995. Two relict species of Thracia (Bivalvia) from
the Pliocene Ogikubo Formation in Nagano Prefecture,
Central Japan. Venus 54: 143-151.
Amano, K. 1997. Taxonomy and distribution of Cymatiidae
(Gastropoda) from the Pliocene and Lower Pleistocene in
the Japan Sea Borderland. Venus 56: 121-129.
Amano, K. 2001. Pliocene molluscan fauna of Japan
Sea borderland and the Ws ue conditions.
Biological Science (Tokyo) 53: 178-184. (in Japanese)
Amano, K. 2007. The Omma- Manganji fauna and its temporal
change. Fossils (Palaeontological Society of Japan) 82:
6—12. (in Japanese with English abstract)
Amano, K. 2019. Two new gastr ‘opods from the late Pliocene
Omma-Manganji fauna in the Japan Sea borderland of
Honshu, Japan. Paleontological Research 23: 85—94.
Amano, K., M. Hamuro, and T. Sato. 2008. Influx of warm-water
curr ait to Japan Sea during the Pliocene - based on analysis of
molluscan fauna from the Mita Formation in Yatsuo-machi of
Toyama City. Journal of the Geological Society of Japan 114:
516-531. (in Japanese with English abstract)
Amano, K. and S. Karasawa. 1986. A new subspecies of Chlamys
lioica (Dall) from the Pliocene Shigarami Formation in the
environs of Nagano City, central Japan. Monograph of the
Mizunami Fossil Museum 6: 41-47.
Amano, K., K. Nagata, T. Sato, Y. Yanagisawa, and Y. Kurita.
2009. Influence of a warm-water current on the northem
Fossa Magna region during the Pliocene, based on analyses
of sialllnseam fauna from the Shitoka formation and the
lowermost part of the Uonuma Group along the Kama-
kurazawa River in Minami-Uonuma City, Niigata Pre-
fecture. Journal of the Geological Society of Japan 115:
597-609 (in Japanese with English abstract)
Amano, K. and H. Sato. 1995. Relationship between embay-
mental associations and relict species. — Molluscan fauna
from the Pliocene Joshita Formation in the northern part of
Nagano Prefecture. Fossils (Palaeontological Society of Ja-
pan) 59: 1-13 (in Japanese with English abstract)
Amano, K., T. Sato, and T. Koike. 2000a. Paleoceanographic
conditions during the middle Pliocene in the central part of
Japan Sea Borderland. Molluscan fauna from the Kuwae
Formation in Shibata City, Niigata Prefecture, central Ja-
pan. Journal of the Geological Society of Japan 106:
$83—894 (in Japanese with English 1 abstract)
Amano, k., M. Suzuki, and T. Sato. 2000b. Warm-water influx
into Japan Sea in the middle Pliocene —Molluscan fauna
from the Tentokuji Formation around Mt. Taihei in Akita
Prefecture. Journal of the Geological Society of Japan 106:
299-306 (in Japanese with English abstr act)
Amano, K. and M. Watanabe. 2001. Taxonomy and distribution
of Plio-Pleistocene Buccinum (Gastropoda: Buccinidae) in
Northeast Japan. Paleontological Research 5: 215-226.
Amano, K., A. Yoshida, and T. Sato. 2011. Influence of cooling
event at 2.75 Ma on the molluscan fauna of the Japan Sea
borderland: Molluscan fauna from central and northern
parts of Akita Prefecture. Journal of the Geological Society
of Japan, 117: 508-522. (in Japanese with English abstract)
Bailey, I, G.M. Hole, G.L. Foster, P.A. Wilson, C.D. Storey,
C.N. Trueman, and M.E. Raymo. 2013. An alternative
suggestion for the Pliocene onset of major northern
hemisphere glaciation based on the geochemical prove-
nance of North Atlantic Ocean ice-rafted debris. Quater-
nary Science Reviews 75: 181-194.
Bouchet, P. 2011. Pachypoma. World Register of Marine Species.
http:/Avww.marinespecies.org/aphia.php?p=taxdetails&id=
579277
Bouchet, P., J.-P. Rocroi, B. Hausdorf, A. Kaim, Y. Kano, A.
Niitzel, P. Parkhaev, M. Schrédl, and E.E. Strong. 2017.
Revised Classification, Nomenclator and Typification of
Gastropod and Monoplacophoran Families. Malacologia 61:
1-526.
Chinzei, K. and Y. Matsushima. 1987. Molluscan faunas in the
South Fossa Magna region. Fossils (Palaeontological So-
ciety of Japan) 43: 15-17 (in Japanese with English abstract)
Dall, W.H. 1891. Scientific results of explorations by the U.S.
Fish Commission Steamer Albatross. XX. On some new
or interesting West American shells obtained from
dredgings of the U.S. fish commission steamer Albatross
in 1888. Proceedings of the U.S. National Museum 14:
174-191.
Dall, W.H. 1910. New species of West American shells. The
Nautilus 23: 133-136.
Dillwyn, L.W. 1817. A descriptive catalogue of Recent shells,
arranged according to the Linnean mao: with particular
attention to the synonymy. Vol. 2. J. & A. Arch, London, pp.
581-1092.
Donald, K.M., M. Kennedy, and H.G. Spencer. 2005. The
phylogeny and taxonomy of austral monodontine topshells
iroliaees Gastropoda: Trochidae), inferred from DNA
sequences. Molecular Phylogenetics and Evolution 37:
474483.
Fischer, P. 1885. Manuel de conchology et de paléontologie
conchyliologique ou Histoire des mollusqueés vivantes et
fossils ah d'un appendice sur les Brachiopodes par D. P.
Oehlert) (9). F. Savy, Paris, pp. 785-896.
Gallagher, S. 1. Teun A., Iryu, Y., Itaki, T., Koizumi, I. and
Hoiles, P. W., 2015: The Pliocene to recent history of the
Kuroshio and Tsushima Currents: a multi-proxy approach.
Progress in Earth and Planetary Science 2: 17, DOI
10.1186/s40645-015-0045-6.
Gmelin, J.F. 1791. Caroli a Linné Systema naturae per
regna tria naturae. Editio decima tertia. Leipzig,
Germany, vol. 1, pt. 6, cl. 6, Vermes, 3021-3910.
Gray, J.E. 1850. Explanation of plates and list of genera. In M.E.
Gray, Figures of molluscous animals, selected from various
authors. val 4. Longman, Brown, Green and Longmans,
London, 124 pp.
Gray, J.E. 1857. Guide to the systematic distribution of Mollusca
in the British Museum. Part I. London, British Museum
(Natural History), 230 pp.
Hickman, C.S. and J.H. McLean. 1990. Systematic revision
and suprageneric classification of trochacean gastropods.
Science Series of the Natural History Museum of Los
Angeles County 35: 1-169.
Higo, Si} P. €: allomon, and Y. Goto. 1999. Catalogue and bibli-
ography of the marine shell-bearing Mollusca o Japan. Elle
Scemene Publications, Yao, 749 pp.
Itaki, T. 2016. Transitional changes in microfossil assemblages in
the Japan Sea from the Date Phoceneite Early Pleistocene
related to global climatic and local tectonic events. Earth
and Planetary Science 3:11 DOI 10.1186/s40645-0087-4.
Itoigawa, J. and I. Nishikawa 1976. A few problems on the
Miocene Setouchi Series in the northern part of Okayama-
Hiroshima Prefectures, southwest Japan. Bulletin of the
Mizunami Fossil Museum 3: 127-150. (in Japanese with
English abstract)
kK. Amano, 2019
Itoigawa, J. and H. Ogawa 1973. Pleistocene Molluscan Fauna of
“dhe 6 Sakishima Formation, Shima Peninsula, Central Japan.
The Science Reports of the Tohoku University, Second
series, Geology, Special Volume 6: 69-80.
Keen, M. 1971. Sea shells of tropical west America. Stanford
University Press, Stanford, 1064 pp:
Kobayashi, I., M. Tateishi, T. Yoshioka, and M. Shimazu. 1991.
Geology of the Nagaoka District. Geological Survey of Japan,
selena 132 pp. (in Japanese with English abstract)
Kiister, H.C. 1858-1960. Die Gattungen Bnaaitpunin, Purpura,
Concholepas und Monoceros. In “Kister, H. C. ed., Sys-
temetisches Cponchylien-Cabinet von Martini und
Chemnitz, vol. 3: 1-299.
Lamarck, J.B.P.A. de M. 1799. Prodrome dune nouvelle
Classification des Coquilles. Mémoires de la Société
d Histoire Naturelle de Paris 8: 63-91.
Lamarck, ].B.P.A. de M. 1822. Histoire naturelle des Animaux
sans Vertébres. Vol. 7. Paris, 711 pp:
Link, H.F. 1807. VII Mollusken. In Beschreibung der
Naturalien-Sammlung der Universitit zu Rostock. Rostock:
Adlers Erben. Section 3: 101—160.
Linnaeus, C. 1758. Systema Naturae (tenth edition). Holmiae,
Laurentii Salvii, $24 pp.
Makiyama, J. 1926. Tertiary fossils from north Kankyo-do,
Korea. Memoirs of the ‘College of Science, Kyoto Impe-
rial University, series B, 2: 143- 160.
Makiyama, J. 1934. The Asagaian Mollusca of Yotsukura and
Machgar. Memoirs of five College of Science, Kyoto Im-
perial University, series B, 3: 1- 147.
Martini, E. 1971. Standard Tertiary and Quaternary calcareous
nannoplankton zonation. In Farinacci, A., ed., Proceedings
of the Second Planktonic Conference, 739-785, Tecno-
scienza, Roma.
Matsushima, Y., K. Taguchi, and K. Chinzei. 2003. Molluscan
fossils from the Ochiai Formation, the Tanzawa Mountains,
Central Japan. Bulletin of the Kanagawa Prefectural Mu-
seum, Natural Science 32: 27-68.
Matsuura, N. 1977. Molluscan fossils from the Late Pleistocene
marine terrace deposits of Hokuriku Region, Japan Sea side
ee Central Japan. Science Reports of Kanazawa Univ ersity
22: 117-162.
Matsuura, N. 1985. Successive change of the marine molluscan
faunas from Pliocene to Holocene in Hokuriku Region,
Central Japan. Bulletin of the Mizunami Fossil Museum 12:
71-158. (in Japanese with English abstract)
McLean, J.H. 1970. New eastern Teacine subgenera of Turbo
Linnaeus, 1758 and Astraea Réding, 1798. The Veliger 13:
=.
Nagamori, H. and H. Yoshikawa. 2019. The distribution and
morphology of the genus Meretrix from the Pliocene of
Japan. Abstracts with Pr ograms of the 168" regular meeting
of the Palaeontological Society of Japan: 30. (in Japanese)
Nakagawa, T. 2009. Miocene tropical sandy and rocky bottom
molluscan assemblages of the Shimo Formation, Uchiura
Group, Fukui Prefecture, Central Japan. Bulletin of the
Mizunami Fossil Museum 35: 127-151. (in Japanese with
English abstract)
Nakagawa, T. 2018. Fossils from Ogrui in Takahama Town, Oi-
gun, Fukui Prefecture. Fukui City Museum of Natural
History, 78 pp. (in Japanese, title translated)
Nakagawa, T., O. Fukuoka, S. Fujii, and M. Chiji. 1993. Fossil
dralll assemblages in the Holocene Takahama shell bed
discovered at Atanarnee Cho, western part of Fukui Pre-
fecture, Central Japan. Monograph of the Fukui City
Page 65
Museum of Natural History 1: 1-113. (in Japanese with
English abstract)
Ogasawara, K., K. Masuda, and Y. Matoba, eds. 1986. Neogene
and Quatemary Molluscs from the Akita Oil-field, Japan,
310 p. Commemorative Association of Professor Taisuke
Takayasu’s Retirement and Supporters’ Foundation of
Mineral Industry ee Mining College, Akita Uni-
versity, Akita, 310 pp. (in Japanese)
Ozaki, H. 1954. On the paleontology of the basal conglomerate
of Pliocene in Tyosi City, Kanto Region. Bulletin of the
National Science Muesum 34: 9-20.
Ozawa, T. and S. Tomida. 1992. The Zushi fauna — Late Miocene-
Early Pliocene warm marine water molluscan fauna of Japan.
Bulletin of the Mizunami Fossil Museum 19: 427-440. (in
Japanese with English abstract)
Philippi, R.A. 1845. Abbildungen und Beschreibungen neuer
oder wenig gekannter Condhsliion, unter Mithiilfe mehrer
Deutscher Conchyliologen. Vol. 1: 187-190.
Pilsbry, H.A. 1904. New Japanese marine Mollusca; Pelecypoda.
Pr oceedings of the Academy of Natural Sciences of Phil-
adelphia 56: 550-561.
Pimiento, C., J.N. Griffin, C. F. Clements, D. Sivestro, S. Varela,
M. D. Uhen, and C. Jaramillo. 2017. The Pliocene marine
megafauna extinction and its impact on functional diversity.
Nature Ecology & Evolution: 1: 1100-1106.
Rafinesque, C.S. 1815. Analyse de la nature, ou Tableau de
lUnivers et des Corps Organisés. Privately published,
Palermo, 224 pp.
Réding, P.F. 1798. Museum Boltenianum, sive catalogus
cimeliorum e tribus regnis naturae quae olim collegerat Joa.
Fried. Bolten, M.D. p. d. per XL. annos Proto physicus
Hamburgensis. Pars secunda continens conchylia sive
Testacea univalvia, bivalvia & multivalvia. J. C. Trappii,
Hamburg, 199 pp.
Sato, T. and K. Kameo. 1996. Pliocene to Quaternary calcareous
nannofossil biostratigraphy of the Arctic Ocean, with ref-
erence to Late Shean glaciation. In: Thiede, J., A. M.
Myhre, J. V. Firth and W. F. Ruddiman (eds.) Proceedings
of the ODP, Science Results 151: 39-59. Texas A & M
University, College Station.
Sato, T., T. Saito, S. Yuguchi, H. Nakagawa, K. Kameo, and T.
Takayama. 2002. Late Pliocene * elleareons nannofossil
paleobiogeography of the Pacific Ocean: evidence for
glaciation at 2.75 Ma. Revista Mexicana de Ciencias
Geolégica 19: 175-189.
Shiba, M., Y. Hirose, T. Nobuhara, K. Takagi, M. Yasuda, Kk. Fuji
and M. Nakamura. 2013. The stratigraphy and fossil molluscan
assemblages of the so-called Shizukawa Group, the Neogene
series in the Fuji River Valley, central Japan. Earth Science
(Chikyu Kagaku) 67: 1-19. (in Japanese with English abstract)
Shibata, M. 1957. Some molluscan fossils from het eastern part
of the Tanzawa Mountainland. Transactions and Pro-
ceedings of the Palaeontological Society of Japan, New
Series 25: 21-25.
Shimamoto, M. and T. Koike. 1986. The molluscan assemblage
from the Tentokuji Formation, southwest of Mt. Taihei,
Akita Prefecture. Saito Ho-on Kai Museum Research
Bulletin 54: 27—49.
Stanley, S.M. 1986. Anatomy of a regional mass extinction: Plio-
Pleistocene decimation of the western Atlantic bivalve
fauna. Palaios 1: 17-36.
Stanley, S.M. and L.D. Campbell. 1981. Neogene mass ex-
tinction of western Atlantic molluscs. Nature 293: 457-459.
Page 66
Todd, J.A., ].B.C. Jackson, K.G. Johnson, H.M. Fortunato, A.
Heitz, M. Alvarez, and P. Jung. 2002. The ecology of ex-
tinction: molluscan feeding and faunal turnover in the
Caribbean Neogene. Proceedings of the Royal Society B
269: 571-577.
Vermeij, G.J. 1991. Anatomy of an invasion: the trans-Arctic
interchange. Paleobiology 17: 281-307.
Vermeij, G.]., GP. Dietl, and D. G. Reid 2008. The trans-Atlantic
history of diversity and body size in ecological guilds. Ecology
89 (Supplement): S39-S52.
Williams, S.T. 2007. Origins and diversiflcation of Indo-West
Pacifle marine fauna: evolutionary history and_bio-
geography of turban shells (Gastropoda, Turbinidae).
Biological Journal of the Linnean Society 92: 573-592.
Williams, S.T. 2012. Advances in molecular systematics of the
vetigastropod superfamily Trochoidea. Zoologica Scripta
doi:10.1111/).1463-6409.2012.00552.x
THE NAUTILUS, Vol. 133, No. 2
Williams, S.T., S. Karube, and T. Ozawa. 2008. Molecular
systematics of Vetigastropoda: Trochidae, Turbinidae and
Trochoidea redefined. Zoologica Scripta 37: 483-506.
Williams, S.T., K.M. Donald, H.G. Spencer, and T. Nakano.
2010. Molecular systematics of the marine gastr opod
families Trochidae and Calliostomatidae avollinces Su-
perfamily Trochoidea). Molecular Phylogenetics and
Evolution 54: 783-809.
Woesik, R. van, E.C. Franklin, J. O'Leary, T.R. McClanahan, J.S.
Klaus and A.F. Budd. 2012. Hosts of the Plio-Pleistocene
past reflect modem-day coral vulnerability. Proceedings of
the Royal Society B 279: 2448-2456.
Wood, W. 1828. A Supplement to the Index testaceologicus, or a
catalogue of shells, British and foreign, arranged according
to alive Linnean system. W. Wood, London, 59 pp:
Yokoyama, M. 1924. Mollusca from Coral-bed of Awa. Journal of
the College of Science, Imperial University of Tokyo 45, pt.
]: 1-62.
Sponseiee 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
™
INSTRUCTIONS TO AUTHORS
The Nautilus publishes articles on all aspects of the bi-
ology, paleontology, and systematics of mollusks. Manu-
scripts describing original, unpublished research and
review articles will be considered. Brief articles, not ex-
ceeding 1000 words, will be published as Research Notes
and do not require an abstract.
Manuscripts: Each original manuscript and accompanying
illustrations should be submitted to the editor via e-mail.
Authors should follow the general recommendations of
Scientific Style and Format—The CSE Manual for Au-
thors, Editors, and Publishers, available from the Council
of Science Editors at http://www.scientificstyleandformat.
org/Home.html.
The first mention of a scientific name in the text should
be accompanied by the taxonomic authority, including
year. Metric, not English, units are to be used. The se-
quence of sections should be Title, Author(s) and Affili-
ations, Abstract, Additional Keywords, Introduction,
Materials and Methods, Results, Discussion, Conclusions,
Acknowledgments, Literature Cited, Tables, Figure
Captions, Figures. If the author for correspondence is not
the senior author, please indicate in a footnote. The ab-
stract should summarize in 250 words or less the scope,
main results, and conclusions of the article. Abstracts
should be followed by a list of additional keywords. All
references cited in the text must appear in the Literature
Cited section and vice-versa. Please follow a recent issue
of The Nautilus for bibliographic style, noting that journal
titles must be unabbreviated. Information on plates and
figures should be cited only if not included within the
pagination of cited work. Tables must be numbered and
each placed on a separate page. If in doubt, please follow
a recent issue of the journal for sequence of sections and
other style requirements.
Illustrations: Illustrations are rendered either at full-
page width (maximum width 17 cm) or column width
(maximum width 8.2 cm). Please take these dimensions
into consideration when preparing illustrations. Page-
width illustrations ideally should span the entire width of
printed page (17 cm). “Tall” page-width illustrations
should be avoided, square or “landscape” formats work
better. Please design plates accordingly, such that there
will be enough space left at the bottom of printed page for
plate caption. (Digital technology has made this task much
easier.)
All line drawings must be in black, clearly detailed, and
completely labeled. Abbreviation definitions must be
included in the caption. Line drawings must be high
resolution fi les at least 600 dpi (dots per inch) resolution
at actual size. Standard digital formats for line drawings
include .tif, .bmp, .psd, .eps, and .pdf.
Photographs may be submitted in black-and-white or
color, preferably in RGB mode if in color. Standard digital
formats for photographs include .tif, .psd, .jpg. or .pdf.
Photographs must be high resolution files at least 300 dpi
resolution at actual (printed) size.
If more than one figure is included in an illustration,
all figures are to be consecutively numbered (Figures
1h Be obo. INGMP Toisas IVAN, NBS. IC, O34 NOR Plate 1,
Figure 1, . . .). In illustrations with more than one
figure, make sure that blank areas between figures
should be kept to a minimum, thereby allowing for
more area for each individual figure.
Compressed (e.g., .jpg) or other low-resolution file
formats may be used to facilitate original submission and
the review process, but may not be acceptable at final
submission (see below).
Types and Voucher Specimens: Deposition of the
holotype in a recognized institutional, public collection is
a requirement for publication of articles in which new
species-level taxa are described. Deposition of paratypes
in institutional collections is strongly recommended, as is
the deposition of representative voucher specimens for all
other types of research work.
The Editorial Process: Upon receipt, all manuscripts
are assigned a number and acknowledged. The editor
reserves the right to return manuscripts that are sub-
standard or not appropriate in scope for journal. Manu-
scripts deemed appropriate for the journal will be sent for
critical review to at least two reviewers. The reviewers
recommendations will serve as basis for rejection or
continuation of the editorial process. Reviewed manu-
scripts will be sent back to authors for consideration of the
reviewers comments. The revised version of the manu-
script may at this point be considered accepted for
publication by the journal.
Final Submission: Authors of accepted manuscripts are
required to submit a final version to the editor at jleal@
shellmuseum.org. High-resolution image files may be sent
to the editor at this stage.
Proofs: After typesetting, proofs will be sent to the author.
Author should read proofs carefully and send corrections to
the editor within 48 hours. Changes other than typesetting
errors will be charged to the author at cost.
Offprints: An order form for offprints will accompany the
proofs. Offprints will be ordered directly from the editor.
Authors with institutional, grant, or other research sup-
port will be asked to pay for page charges at the rate of $60
per page.
More information at http://shellmuseum.org/learn/the-
nautilus.
This paper meets the requirements of ANSI/NISO Z39.48-1992 (Permanence of Paper)
IOWA ANNU
3 9088 02022 2485
Live Music Archive
Librivox Free Audio
Metropolitan Museum
Cleveland Museum of Art
Internet Arcade
Console Living Room
Books to Borrow
Open Library
TV News
Understanding 9/11