The Nautilus

^^ u JiJi

THE NAUTILUS

Volume 113, Number 1
March 29, 7.9.99
ISSN 0028-1344

A quarterly devoted
to malacolooij.

EDITOR-IN-CHIEF

Dr. Jose H. Leal

The Bailey-Matthews Shell Museum
3075 Sanibel-Captiva Road
Sanibel, FL 33957

EDITOR EMERITUS

Dr. M. G. Harasewych
Department of Invertebrate Zoology
National Museum of
Natural History
Smithsonian Institution
Washington, DC 20560

CONSULTING EDITORS

Dr Riidiger Bieler
Department of Invertebrates
Field Museum of
Natural History
Chicago, IL 60605

Dr. Arthur E. Bogan

North Carohna State Museum of

Natural Sciences

Raleigh, NC 27626

Dr Phihppe Bouchet
Laboratoire de Biologie des
Invertebres Marins et Malacologie
Museum National d'Histoire Naturelle
55, rue Buffon
Paris, 75005 France

Dr Robert T Dillon, Jr.
Department of Biology
College of Charleston
Charleston, SC 29424

Dr. William K. Emerson

Department of Living Invertebrates

The American Museum of Natural

History

New York, NY 10024

Dr. Eileen H. Joldnen
Institute of Water Resources
Universit\' of Connecticut
Storrs, Ct 06269-4018

Mr. Richard I. Johnson
Department of MoUusks
Museum of Comparative Zoology
Harvard University
Cambridge, MA 02138

Dr. Douglas S. Jones
Florida Museum of Natural History
University of Florida
Gainesville, FL 32611-2035

Dr James H. McLean

Department of Malacology

Los Angeles County Museum of

Natural History

900 Exposition Boulevard

Los Angeles, CA 90007

Dr Arthur S. Merrill
% Department of Mollusks
Museum of Comparative Zoolog)'
Harvard University
Cambridge, MA 02138

Dr Paula M. Mikkelsen

Department of Living Invertebrates

The American Museum of Natural

History

New York, NY 10024

Dr. Gustav Paulay
Marine Laboratory
University of Guam
Mangilao' Guam 96923

Mr Richard E. Petit

PO. Bo.x 30

North Myrtle Beach, SC 29582

Dr. Edward J. Petuch
Department of Geology
Florida Atlantic University
Boca Raton, FL 33431

Dr. Gary Rosenberg
Department of Mollusks
The Academy of Natural Sciences
1900 Benjamin Franklin Parkway
Philadelphia, PA 19103

Dr Ruth D. Turner
Department of Mollusks
Museum of Comparative Zoology
Harvard University
Cambridge, MA 02138

Dr. Geerat J. Vermeij
Department of Geology
University of Cahfomia at Da\ds
Davis, CA 95616

Dr. G. Thomas Watters
Aquatic Ecology Laboratory
1314 Kinnear Road
Columbus, OH 43212-1194

Dr. John B. Wise

Houston Museum of Natural Science

Houston, TX 77030-1799

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TH E€9NAUTI LUS

Volume 113, Number 1
March 2.9. 1999
ISSN 0028-1344

CONTENTS

Philippe Bouchet

A new Lijria (Gastropoda: Volutidae) from soudieastem
Madagascar

1

Bruce A. Marshall

A rexision of the Recent Solariellinae (Gastropoda:

Trochoidea) of the New Zealand rej^ion

4

V'/c .

1

Liorary

APR 1 6 1999

THE NAUTILUS 113(l):l-3, 1999

Page 1

A New Lijria (Gastropoda: Volutidae) from Soudieastern
Madagascar

Philippe Bouchet

Museum uational d'Histoire naturelle

55 rue BufTon

75005 Paris

FRANCE

bouchet@mnhn.fr

ABSTRACT

Lijria patbaili new species is described based on crabbed shells
taken in lobster pots off Fort-Dauphin, SE Madagascar, a re-
gion with cooler hydroclimatic conditions that demarcate it
from the tropical conditions elsewhere in Madagascar The new
species differs from the two other species of Lijria lixang off
Madagascar, L. delesscrtiana and L tulearensis, bv its much
larger protoconch, narrower teleoconch, with fewer and lower
axial ribs, less elaborate columellar and parietal plaiting, and a
more comple.x color pattern.

Keij words: Indian Ocean, \'olutidae, endemisni; cold water

INTRODUCTION

Only two species of volutes (Gastropoda, Volutidae) are
presently known from Madagascar, both belonging to the
genus Lt/iia Gray, 1847, and both occupying \eiv small
ranges ;ilong the west coast of the island. Li/iia dclcsscr-
tiaim (Petit de la Saussaye, 1842) is restricted to the
northwestern coast, with all accurately localized records
originating from the island of Nosy Be, while Lijria tu-
learensis Cosel & Blocher, 1977 is restricted to the south-
west, near Tulear [= Toliara, Toliary]. A 1000 km latitu-
dinal gap separates the ranges of these two forms, which
Poppe & Goto (1992) treat as geographical subspecies.
Apart from one doubtful and unconfirmed record of Lijr-
ia lijracfonnis (Swiiinson, 1821) from near Tulear (Poppe
& Goto, 1992:72), no Lijria nor any other volute is known
to occur on the coasts of Madagascar The new species
described here has been obtained in the region of Fort-
Dauphin as a byproduct of lobster and crab fishing using
baited traps. The sheDs of the three kniown specimens
were probably occupied by hermit crabs. The soft parts
are not known. The depth where the specimens were
captured is not known precisely, but based on local fishing
habits it is very likely to be less than 100 m, and probably
between 30 and 50 m.

SYSTEM ATICS

Class Gastropoda

Superfamily Volutoidea

Family Volutidae Rafinesque, 1815

Subfamily L\riinae Pilsbrv & Olsson, 1954

Genus Lijiia Gray, 1847

Lijria patbaili new species
(Figures 1, 2)

Description: Shell large, fusiform, naiTovv, verv sohd
and heavy, consisting oi 2.0 protoconch and 5.75 teleo-
conch whorls (Figure 1). Protoconch (Figure 2) large, bul-
bous, apically depressed, widi small slightly involuted nu-
cleus, moderately con\'e.\ whorls, with smooth, glossy sur-
face, slightly appressed suture. Protoconch/teleoconch
transition shaip. Teleoconch whorls moderately convex,
widi shallow suture. Sculpture consisting of strong, low,
oithocHne axial ribs, distincdy sigmoid on last adult whorl,
discrete, narrow spiral groves in rib interspaces. Sixteen
axi;il ribs on first teleoconch whorls, increasing to 19 on
penultimate whorl, 20 on last whorl, interspaces as broad
as ribs. Thick labial vaiix behind peristome. About 10 spiral
groves on first teleoconch whorl, 15 on diird whorl, more
crowded in subsutunil zone, becoming more obsolete on
penultimate and last whorls. Four stronger, raised cords at
base of last whorl. Aperture ovate, nanow, fonning a nar-
row angle adapicidlv, outer lip smooth, inner lip fonning a
distinct callus over body whorl, expanded, moderately
raised over columella; 9 plaits in columellar region, stron-
ger abapicaUy, 3 indistinct wrinkles in parietal region. Si-
phonal canal broad, short, open. Protoconch yiolet pink.
Teleoconch ground colour pinkish salmon widi complex
pattern of spiral lines and blotches. Spiral lines bnwTi, in-
terrupted in interspaces between ribs, four on spire whorls,
nine on last whorl, evenly spaced, delimiting a subsutural
band twice as broad as distance between lines; brown
blotches occupying two spiral liands, the broad siibsutmiJ
band and a naiTow band, situated abapic;illy of exposed
part of spire whorls and on peripher) of last whorl. Si-
phonal band ;md edge of siphonal canal with axially elon-
gateil browm \iolet stripes. Aperture creamy salmon.

Type material: Holotvpe in MNHN. Paratvpe in die col-
lection of Mr Harald Doute (Bad Sackingen, Gennany).

Type locality: Region of Fort-Daupliin [= Taolanaro],
soudiem Madagascar (ca. 25°01' S, 47°00' E), from fish-
ermen.

Dimension.s: Length 72.0 mm, width 26.0 mm, ap-
erture length 37.0 nun.

Remarks: The parahpe shares the characters of the
holotvpe, but is proportionalK' narrower with a shorter
aperture (length 65.5 mm, width 21.0 mm, aperture
length 28.0 mm), and has three, rather than four, brown
spiral lines on spire whorls, and eight on last whorl. Li/r-

Page 2

THE NAUTILUS, Vol. 113, No. 1

a

^JV.

)»«. »*-i" ?4'" J -'^

Figure I. Lj/rin pathaili
Heialii 72.0 mm.

new species Holot\pe MNHN.

ia patha'Ji di
inbalvinng M
(iruis h: ''-^ i,-
conch, vviii;
coiiiinr-ti-ir t'.r
or pai-\ ::
cies, ii:-] -
patbaii: ;; ': .
reddisli l-kr
with calcarell
leoconch wh<:

ff<='r.s from the Kvo other species of Lt/ria
iciagascar, L. dclcssciiiaiia and L tnlear-
ii;ch larger protoconch, its narrower teleo-

v-r and lower axial ribs, less elaborate

'! ";:<:rictal plaiting, and a more complex col-

■:!aefonms is also a rather slender spe-

. '.'• ■ ."/lour pattern resembles that of L.

'. • •; spiral bands occupied by brown
':" ■ . -A has an even larger protoconch
:. ' • •• . ■ : ,;! termination, more convex te-
iris 0. ' 1 ■'. ;n<erous brown spiral lines.

Figure 2. Lijria pathaili new species. Protoconch of para-
hpe, length 65.5 mm. Doute collection. Scale hne = 5 mm.

Etymology: The specific name honors Dr Patrice
Bail, volute collector and expert, and president of As-
sociation Fran^aise de Conchvliologie. Dr Bail very gen-
erouslv donated the holotype to MNHN.

Distributional and biogeographical remarks: The

size and nloq5hol()g^• of its protoconch suggest that Lt/ria
pathaili, like manv other volutes, has non-planktotrophic
development with intracapsular metamorphosis and
hatches as a crawling juvenile.

The region of Fort-Dauphin is characterized in Mad-
agascar bv its spectacular belts of brown macrophvtes,
ressembhng the algal belts of temperate coasts (A. Cros-
nier, pers. com.); there are no coral reefs in the area.
Published satellite data on winter sea surface tempera-
tures (Piton & Laroche, 1993) confirm a local anomaly,
with temperatures as low as 21.5°C, vs 24- 25°C or more
elsewhere around Madagascar. Although the total range
of Li/ria pathaili is not known, it is tempting to associate
the local hvdroclimatic conditions with the narrow dis-
tribution of certiiin marine biota. Similarlv. Paliiutnts de-
lagoae Barnard, 1926, a commercial species of lobster
(Crustacea: Palinuridae) has a range in Madagascar re-
stricted to near Fort-Dauphin, and other^vise extends
from Mozambique to Natal (Holthuis, 1984).

With the addition of Lifria pathaili. five species of
Lijria are now knowii from the SW Indian Ocean (Fig-
ure 3) and a further two from Saya de Malha Bank
(Bouchet & Bail, 1991). This distribution pattern is co-
herent with the scenario hvpothesized bv Bouchet &
Biiil (1991), who suggested that colonization by lecitho-
trophic larvae, although a rare dispersal event, is re-
sponsible for the existence of morphologically segregate,
allopatric species.

p. Bouchet. 1999

Page 3

10°-

20"-

50"

60°
I

Lyria delessertiana

Lfiia anna

0

c>

Reunion

Mauritius

Fort-Dauphin

Vf W Lyria patbaili

Figure 3. Generalized distributions (hatched areas) of the 5 species of Lyria occurring in the SW Indian Ocean. Paratype of L.
patbaili illustrated. Distribution data from Weaver & duPont (1970) and Poppe & Goto (1992); however, \Vea\er & duPont (1970:
18) give the range of L. delessertiana as extending to the Comoro Islands and the southern Seychelles, and Poppe & Goto (1992:
65) record the range as extending to Diego Suarez [= Antseranana] but this needs confirmation. Two forms of Lyria lyraefonnis
are illustrated, the broad fomi occurs in all the species range from Mozambique to Somalia, the slender form occurs predominantly
in the central part of its range, i.e. Kenya (Bail, pers. com.).

LITERATURE CITED

Bouchet, P and P Bail. 1991. Volutes from Saya de Mallia
Bank: The saga of Lyria siiriiiamcnsis and a new species.
The Nautilus' 105(4);i.59-164.

Holthuis, L. B. 1984. Lobsters. In W. Fischer & G. Bianchi
(eds.), FAO species identification sheets for fishery pur-
poses. Western Indian Ocean. Volume 5. Unnumbered
pages. FAO, Rome.

Piton, B. and ]. Laroche. 199.3. Qiielques caracteristiques hv-
droclimatiques du siid de Madagascar. Bulletin Oceano-
graphie et Peches [La Reunion] 37:46-54.

Poppe, G. and Y. Goto. 1992. Volutes. L'Informatore Piceno,
Ancona. 348 pp.

Weaver, C. S. and ]. E. DuPont. 1970. Living Volutes. A mono-
graph of the Recent Volutidae of the world. Delaware Mu-
seum of Natural History-, Greenville. 375 pp.

THE NAUTILUS 113(l):4-42, 1999

Page 4

A Revision of the Recent Solariellinae (Gastropoda: Trochoidea)
of the New Zealand Region

Bruce A. Marshall

Museum of New Zealand Te Papa

Tongarewa
P.O. Box 467
Wellington
NEW ZEALAND
bmceni@tepapa.govt.nz

ABSTRACT

Twent)'-nine species (17 new) of Solariellinae are recorded
from the Norfolk Ridge, Three Kings Rise, Kermadec Ridge,
and New Zealand. The species are referred to Solariella Wood,
1842 (10 species, 4 of which are new), Bathijinophila Dall,
1881 (3 new), Microgaza Dall, 1881 (1 ne-w),Archiminolia Ire-
dale, 1929 (10, 6 new), and Zctcln Finlay, 1926 (5, 3 new).
Zcminolia Finlav, 1926, is treated as a sviionynn of Solariella.
Bathtpnophila Dall, 1881, is transferred from Margaritinae to
Solariellinae on the basis of radular morphologv Etiialiopsis
Schepman, 1908, is synonvinized with Bathijinophila. the shell
of a paralectot\pe of Solariella {Ethaliopsis} callomphala
Schepman, 1908, is illustrated, and a lectotype is designated
for the species. Lamellitrochus Quinn, 1991, (based on a west-
em Atlantic species) is regarded as a probable s\aionviii of Ze-
tela. and the southern African species Solariella intennissn
Thiele, 1925, and the North Atlantic species Trochiis (Marga-
rita) rhina Watson, 1886 (= Solariella cincta sensu Dautzen-
berg and Fischer, 1896, and Dautzenberg, 1927, not Philippi,
1836) are referred to Zetela. Se.xually dimorphic shell mor-
phology in New Zealand Solariella species is discussed.

Key words: Trochidae, Solariellinae, new taxa. New Zealand,
Kermadec Ridge, Norfolk Ridge, North Atlantic, western At-
lantic, southern Africa.

INTRODUCTION

The trochid .subfamily Solariellinae has a cosmopolitan dis-
tribution ii; siibhttoral to abyssal depths. SolarieUines are
deposit feeders, and most if not all of them ;u"e proficient
sedimetU I, -rrovv'ers. Species examined alive are capable of
sv.'iriiiP.uii'. by rnpidly moving the foot from side to side,
evideiiH) . ., : . r.icape response (Herbert, 1987). Although
seveiiil nprdt.:^ arc consistendy collected ali\e from rugged
substrata ir- ■ . Lirion with bryozoans, sponges, hydroids,
and cvi:. >. iikely that they live and feed in the

surroiMi'.i' .. i .; • j sediments ratlier than in association
withothi" !n. ■';■<€ of the New Zealand .species have

exceptionally p •• i : ii-'nbutions for sediment dwellers,
suggesting higlil)- s; ec.i '-cological requirements. The sex-

es are separate and some species brood within die piiUial
cavity or the umbilicus. Whereas shell moiphology is ire-
quendy litde different from that in Margaritinae and Um-
boruinae and is interspecifically variable, the animal is dis-
tinctive in having tentaculifonn processes on die snout and
a short, broad radula with a low number of rows (ca. 20)
of teeth; few, simple marginal teedi (6-10 pairs); and dis-
tinctive central and lateral tooth morphology. The central
and inner diree pairs of latenJs are short and stout with
angulate cutting ;u-e;is (lateral 3 may have a reduced cutting
area), and lateral 4 is elongate, curved, and typically lacks
secondary cusps. Many species are further characterized
by having die innennost pair of marginal teeth transfonned
during ontogenesis into broad laterornarginal plates.

Abbreviations and text conventions:

AIM Auckland Institute and Museum

AMS Australian Museum, Sydney

cf compare (with)

BMNH The Natural History- Museum, London

NMNZ Museum of New Zealand Te Papa Tongarewa,
Wellington

NZOI National Institute of Water and Atmospheric
Research, Wellington

TW

(number of) teleoconch whorls

USNM National Museum of Natural Historv, Wash-
ington, DC.

ZMA Zoological Museum, Amsterdam

In all measurements of shells, length precedes width.
Unless specified, all material was collected as tlead shells
and is deposited at NMNZ.

B. A. Marshall, 1999

Pages

SYSTEM ATICS

SuperfamiK' Trochoidea Rafinesque, 1815
Family Trochiclae Rafinesque, 1815
Subfamily Solariellinae Powell, 1951
Solaiiellinae Powell. 1951: 102 — nominohpical genus

Solariella Wood, 1842.
Talopiidae Finlas, 1928: 238 — noniinot\pical genus Tnlopia

Gray, 1842 '(in part).
Minoliinae Kuroda, Habe, and Ovama, 1971: 26 —

nominohpical genus Minolia A. Adams. 1860.

Remarks: Inteqiretation of groups of solarielline gen-
era has been comphcated by the fact that the type spe-
cies of Solariella (S. maculata Wood, 1842) is a Phocene
fossil and because some Recent species that resemble it
have adult radulae with lateromarginal plates, whereas
other species lack them. One Recent species that bears
a particularly close resemblance to S. maculata (Herbert.
1997, fig. 204-206) is S. cincta (Philippi, 1836) (Waren.
1993, fig. 2E, as S. inoptanda Locard, 1897 = S. cincta).
the adult radula of which lacks lateromarginal plates (F.
Rubio, pers. comm.). Another similar European species
is Solariella aiiiabilis (Jeffreys, 1865) (figure 40). the
adult radula ot which has broad lateromarginal plates
(Waren, 1993. fig. 7B). Solariella amabilis (= Trochus
ajftnis Friele, 1877) is the type species oi Machaeroplax,
which has generally been inteqireted as a svnonym ol
Solariella (Odliner,' 1912; Thiele, 1929; Marshall.' 1979;
Quinn, 1979; Hickman and McLean, 1990; Waren, 1993;
herein), although Herbert (1987) was more cautious in
his approach.

Waren (1990) has shovvai that solarielline lateromar-
ginal plates appear during radular ontogenesis. He was
unable to ascertain their origin, but it seems likely that
they arise by progressive in-column transformation of ex-
isting teeth such as the innermost pair of marginals rath-
er than by intercalation (which among Trochoidea is
known to occur only in calhostomatids). Whatever their
origin, presence or absence in adults of lateromarginal
plates, or degree of transformation of the teeth from
which they apparently develop, cannot be used alone for
separation of genera or subgenera, since radular onto-
genesis will naturally be accelerated or retarded to vary-
ing degrees from species to species. Whereas latero-
marginal plate morphology in adults is infraspecificallv
stable in all species examined, interspecifically the later-
omarginal plate may be broad, short and bar-like, or the
innermost marginals may be shortened with the cutting
area weakened to varying degrees. These considerations
led Waren (1993) to include the North Atlantic species
Solariella obscura (Couthouy, 1838) and S. varicosa
(Mighels and Adams, 1842) (Waren, 1993, fig. 4A, B,
6A-C) in Solariella together with S. amabilis, which they
resemble in external anatomy but lack lateromarginal
plates. Species with broad lateromarginal plates include
the type species of Microgaza Dall, 1881, Spectamen
Iredale. 1924 (? = Solariella), Zcmim^lia Finlay, 1926 ( =
Solariella), Minolops Iredale. 1929 (? = Spectamen), and
Archiminolia Iredale, 1929. Species with shortened or

otherwise modified innermost marginal teeth include
most species herein referred to Zctcla Finlav, 1926,
Batluimophila Dall. 1881 (Waren, 1990, p. 180; figures
130, 131 herein), and the tvpe species of Siiavotrochus
Dall, 1924, Hazuregtjra Shikama. 1962 (A. Waren. pers.
comm.), and Lamellitroclms Quinn, 1991 (? = Zetcla).
The type species of Zetcla, species of llanga Herbert,
1987, and Solariella varicosa and S. obscura have un-
modified innermost marginal teeth.

Herbert (1987) provided evidence to suggest that lo-
cal phylogenetic radiations (notably llanga off southern
Africa) are recognizable from features of both radular
and epipothal moq)holog\-. Epipochal features include
number and relative size of the epipodial tentacles, pres-
ence or absence of flap-like projections (here termed
epipodial lobes) from the epipodial fringe, and number
of neck lobes. From observations of living specimens,
Herbert (1987) recorded southern African species as
having a tiny right postoptic tentacle, although I was im-
able to detect such a structure in any solarieUines re-
corded herein (preserved material). Herbert (1987. p.
295) reported the e.xistence of llanga species with four
pairs of large epipodial tentacles in adults and three pairs
in juveniles, which indicates that the number may in-
crease during ontogenesis. One group (Zetcla) is distinc-
tive in having four pairs of epipodial tentacles, the two
middle pairs of which are typically much smaller than
the others. Many other species (e. g., of Solariella sensii
stricto and Spectamen) have three pairs of epipodial ten-
tacles and a prominent epipochal lobe on each side be-
tween the anterior pairs of tentacles. Judging from the
fact that epipodial lobes have epipocUal sense organs at
their bases (Herbert, 1987, fig. 118), they would seem
to be modified epipodial tentacles, which in turn sug-
gests that four pairs of tentacles is plesiomoq:)hic.
Whether or not arrested development and/or transfor-
mation of epipodial tentacles occurs during ontogenesis
remains to be established. These considerations, togeth-
er with the e.xistence in Archiminolia oleacea (Hedley
and Petterd, 1906) of three right and four left epipodiiil
tentacles and of three pairs in A. meridiana (Dell, 1953),
both of which have very similar shells and radular mor-
phologies, suggest that the number of epipodial tentacles
has somewhat limited value for supraspecific grouping.

Solarielline shell sculpture may be predominantly spi-
ral or axial, or it may be reticulate. Most species have
rather stable shell sculpture, though Solariella obscura
is exceedingly variable, ranging from strongly reticulately
sculptured to almost smooth (Waren, 1993). Whereas
many species are strongly sculptured throughout, others
become more or less smooth following an initial spirally
sculptured stage. The latter include the type species of
Microgaza. Bathi/mophila, Archiminolia, and llanga.
The tvpe species of Suaiotrocluis is essentially smooth
throughout. Species of Baflu/inophila are distinctive in
that the umbilicus is covered by a thick callus with a
granulate surface (produced by an internal layer of mi-
nute calcareous spheres). There is strong convergence
in shell shape and sculpture in the group. For example.

Page 6

THE NAUTILUS, Vol. 113, No. 1

llanga biradiatula (Martens, 1902) and 7. laevissima
(Martens, 1881) (Herbert, 1987) resemble species of Ar-
chiminolia without umbilical septa in shell morj^hologv',
whereas /. discus Herbert, 1987, resembles the t\pe spe-
cies of Microgaza, yet they differ in radular moqihology
and external anatomy (unknown in Microgaza). Although
solarielhnes (or at least solarielline-like species) have
been recorded from as early as the Late Cretaceous
(Hickman and McLean, 1990) and undoubted solariel-
lines are known from the Eocene (Maxwell, 1992), the
fossil record of the group is poor, and there are no doc-
umented lineages. Since it is thus impossible to establish
character-state polarity, it seems likely that species in
which the teleoconch becomes more or less smooth are
derived from species with stronger, more persistent
sculpture. Loss of sculpture, which presumably enhanc-
es sediment penetration, has probably occurred repeat-
edly in the group, almost certainly independently in So-
lariella obscura, Microgaza/Bathyinophila/Archiminolia,
llanga, and Suavotrochus. and quite possibly indepen-
dently in Microgaza, Batlupiiopliila. and Archiininolia as
well. Influenced bv the fact that most species with la-
teromarginal plates (apomorphic) ha\'e three pairs of
epipodial tentacles (apomorphic), whereas those without
lateromarginal plates have four piiirs of epipodial ten-
tacles (both plesiomoiphic), I consider the prime can-
didate for a basal solarielline group to be Zctcia, all spe-
cies of which have strong spiral and (especially) axial
sculpture. Zetela species are strikingly similar to species
of CaUiotropis Seguenza, 190.3 (Eucyclinae), and, in-
deed, there are some similarities in external anatomy
and radular morphology, the implication being that the
groups have a common origin.

In the absence of any firm exddence for degree of
relatedness, I have avoided the use subgenera or to lump
heavily, and I interpret Solariclla, Microgaza, Batliij-
mophila, Suavotrochus, Zetela, Archiminolia, Hazure-
gi/ra, and llanga as distinct genera. Solaiiclhi apparent!)'
has a worldwide distribution, Microgaza western Atlantic
and western Pacific, Bathi/mophila Atlantic and western
Pacific, Suavotrochus western Atlantic, Zetela world-
wide, Archiminolia western Pacific and possibly Indian
Ocean, Hazurcgi/ra northwestern Pacific, and llanga In-
dian Ocean and possibl)' western Pacific.

Higher classification follows Hickman and McLean
(1990).

Geims Solariclla Wood, 1842

S. •.■•.■ ■..;'•'.■<■ Wood, 1842; 5.31. T\pe species (by iiionot)py): So-
. ■: '.'-■:. :■ acidata Wood, 1S42; Pliocene, England.

M.uy:,.-i- ■•..•■;■,■ I'riele, 1877: 311. Type species (by subsequent
;:: .'-c.--..:' !■ , , of Tn.'on, 1889); Trochus ajfinis Friele, 1877
- /• ;■■;■• a-ncbiUs Jeffreys, 1865; Recent, western Eu-

Nev; .Sr.-.iih vV-Jt'-:.
Zeminolia F;ii!rt>, 1926; 359. Type species (by original desig-

;924; 182, 227. Type species (by original
■:l{iis philippeiisis Watson, 1880; Recent.

nation): Minolin plicatula Murdoch and Suter, 1906; Re-
cent, northern New Zealand.
?Minolops Iredale, 1929: 169. Type species (by original desig-
nation): Minolia pulcherrinm emendata [sic — emendata
treated as a full species on the same page by Iredale,
1929] Iredale. 1924; Recent, New South Wales'

Remarks: The animal of Solariclla amabilis (preserved
material. North Sea, 292-340 m. pers. obs.) has a broad
right neck lobe, two smaller left neck lobes, three pairs
of long, tapered epipodial tentacles, and a broad epi-
podial lobe between the two anterior tentacles on each
side. The animal is essentially similar to those of S. ob-
scura and S. varicosa (Waren, 1993, fig. 4A, B). The
animal and radula (Waren, 1993, fig. 7B) are verv' similar
to those of Zeminolia plicatula (preserved material, pers.
obs.) and southern African Spectamen species (Herbert,
1987, fig. 118; NB inl" and nil" are transposed in all
illustrations of animals in this paper). Unfortunately the
external anatomies of the type species of Spectatnen and
Minolops remain unknown (all available animals dry).
Whereas all of the southern species of Solariclla have
lateromarginal plates, they mav be present or absent in
North Atlantic species (see above).

Spectamen philippensis and Zeminolia plicatula (fig-
ures 1—4) differ principally from S maculata, S. amabilis
(figure 40), and S cincta in haxang much weaker axial
sculpture on the teleoconch. Zeminolia plicatula differs
further by having bv having a wider umbilicus, a lower
spire, sexually dimorphic shells, and by brooding its
voung within the umbilicus. Among New Zealand taxa
here refeiTed to Solariclla sensu stricto, however, there
is rather fluid interspecific gradation between the ex-
tremes in umbilical moiphology, spire length, and sculp-
ture (figures 1-12, 15-39, 41-47). In the apparent ab-
sence of any combination of character states bv which
Solariclla and Zeminolia may be distinguished, it seems
that Zeminolia should be treated as a synonym of Sola-
riclla. All of the New Zealand species of Solariclla have
a calcified periostracum (intritaciilx), as in Lamellitro-
chus (Quinn, 1991), which is probably a synonym oi Ze-
tela (see below).

The type species of Minolops has unusually strong,
crowded axial riblets on the teleoconch and is also dis-
tinctive in having fine axial sculpture on the outer part
of the first half whorl of the protoconch (Herbert, 1987,
fig. 214). The a\i;d sculpture on the protoconch oi Mino-
lops emendata is evidently teleoconch sculpture that has
been predisplaced into the larval stage by a heterochron-
ic process.

Solariclla plicatula (Murdoch and Suter 1906)
(Figures 1-4, 13, 18, 41, 125)

Minolia plicatula Murdoch and Suter 1906: 299, pi 26, fig.

47^9.
Monilca {Minolia) plicatula.— Suter. 1913: 142, pi .33, fig. 12.
Xeminoha plicatula.— Fmhy. 1926: .3.59; Keen. 1960: 262, fig.

167/2; Herbert, 1987: fig. 215, 216.
Isanda {Zeminolia) plicatula.— Wenz, 1938: 318, fig. 719.

B. A. Marshall. 1999

Page 7

Figures 1-12. Shells of species of Solariella. 1-4. S. plicatula (Murdoch and Suter), Rangaunu Bay, 6.3 m, M. 130750. 1, 2.
Females (1, 4.00 X 5.50 mm; 2, 4.00 X 5.65 mm). 3, 4. Males (3, 2.85 X 4.45 mm; 4, 2.95 X 4.55 mm). 5-7. S. vera (Powell).
5. Female, Middlese,x Bank, 186-196 m. M. 130064 (4.25 X 6.40 mm). 6. Female, E of Great Island, 100 m. M. 130065 (4.20 X
6.10 mm). 7. Male, NE of Great Island, Three Kings Islands, 550 m, M. 1.30062 (3.05 X 4.45 mm). 8-12. S. hiteola (Powell). 8,
9. Female, off Northeast Island, Three Kings Islands, 102 m, M. 34254 (2.90 X 4.40 mm). 10. Female, Middlesex Bank, 98-103
m, M. 1.30051 (3..30 X 5.00 mm). 11. Male, Middlesex bank, 98-103 m, M. 1.30051 (3.00 X 4.70 mm). 12. SW of Great Island, 440
m, M. 1.30045 (3.45 X 4.90 mm). Scale = 2.50 jxm.

Pas^e 8

THE NAUTILUS. Vol. 113. No. 1

Spectamen plicntiihnu. — Powell. 1976; 84; Powell. 1979; 64, pi.

17. fig. 19.
Spectamen venim. — Powell. 1979: 64 (remarks on Spectamen

onlv); Havward and Grace, 1981; text fig. 3; Hickman.

1992; 254. Not Powell. 1937.

Type material: Holotype NMNZ M.1124. from t\pe
locality, 22 January 1904. S.S. Aioartm.

Type locality: Off Great Barrier Lsland. .36°08'S,
175°55'E. 201 111.

Other material e.xammed (> 1000 specimens): N of

Nortli Cape. 34°20.0'S. 173°()6.ryE. coOected abve, 163-
168 m, 27 Januaw 1981. RA' Tangawa (12. M. 130088);
off North Cape. 91 m, J. A. Bollons (2. M.2167); E of
North Cape, 34°25.0'S, 173°13,1'E, 327-257 m, 27 Jan-
uaiT 1981. IW Tangawa (21, M. 130086); Great E.xhibi-
tion Bav. 34°33.4'S, 173°04.8'E. collected alive. 63 m. 27
January 1981, RA' Tangawa (manv, M. 113347); Great
E.xliibition Bay, 34°33.6'S, 173°04.9'E, collected ;dive. 66
m. 27 January 1981, RA^ Tangawa (19, M. 130067); SW
of Cape Maria yan Diemen, 34°41.9'S, 172°33.9'E, 103
m, 10 Januan- 1981, IW Tangawa (2, M. 131499); SW of
Cape Maria van Diemen. 34°41.9'S, 172°33.5'E, 103 m,

10 January 1981, RA' Tangawa (2, M. 130087); off Cone
Point, N of Whangaroa, collected alive, 44 m, September
1965, M.V. Ikatcrc (3. M.64330); NW of Whangaroa Har-
bour, 34°54.0'S, 173°42.6'E, 83 m, 26 Januaiy'l981. RA^
Tangawa (manv, M. 130767); off Flat Island, Whangaroa,
66 m, September 1965, M.V. Ikatcre (1, M.64472); off
Cape Brett. 35°08'S. 174°12.5'E. 80 m. 16 Februaw
1974. RA' Achcwn (25, M. 35780); NE of Ninepin Rock,
Bay of Islands, 35°08.8'S, 174°10.3'E, coOected iJive. 75
m, 1 December 1971, M.V Kohnga (6, M.30767); be-
tween Redhead and Ninepin Rock and out. Bay of Is-
lands, 35°09'S, 174°09'E. 46-.51 m. 13 December 1973.
M.V Kokinga (1, M.40758); off Takau Bav. 35°10'S,
174°11'E, collected alive, 80 m, 16 February' 1974, RA^
Acheron (9, M. 43652); off Ahipara, ' 35°10.4'S,
172°35.4'E, coOected alive, 233 m, 11 January 1981, IW
Tangaroa (4, M. 130089); off Takau Bay.' 35°10.5'S,
174°10'E, 37 m, 16 February 1974, RA' Achcwn (7,
M,43501); Bay of Islands, 35°10.6'S, 174°15.4'E, 67-73
m. 12 February 1976, M.V Ki^kinga (4, M.96005); Bav of
Islands, 35°11.2'S, 174°16.7'E, coOected idive, 58-64 m,

11 February 1976, M.V Kijkinga (18, M.96143); Deep
Water Cove, Bay of Islands, 35°11.6'S, 174°18.1'E. col-
lected alive, 23-32 m, 10 February 1976, M.V tUikinga
(many M.95538); Bay of Islands. 35°11.6'S, 174°16.5'E.
54 m', 11 February l976, M.V Kokinga (18. M.95910);
off island at entrance to Deep Water Cove, 35°11.9'S.
J7? -7 I'E. coOected alive, 47-49m, 11 Febmaiy 1976.
■■■I.' . .-. ■/;r.'(i;/3 (12, M. 95720); between Hope Passage and
;■■ ■> ,:..:,ici Cove. 35°12'S, 174°18'E, 37-40 m, 10 De-
Cfn":-. ;;^73. MV Kokinga (1. M.41454); Deepwater
r, V ■: • : ...ctf , 35'12'S. 174°18'E, 33-46 m, 6 December
K'" J. '; ■'. KfJidnga (8. M.41499); between Hope Passage
and i\vJc, )<ock.s, Bav of Islands, 35°12'S, 174°18'E, 37
m, 14 i:)cc.>i!.ber K.*' 3, M.V. Kokinga (1, M.41401); be-
tween and N of Black island and Moturoa, Bd\ of Islands,

35°12'S, 174°06'E, coOected alive, 31 m, 13 December

1973, M.V, Kokinga (5, M.41630); Bay of Islands.
35°12.5'S, 174°16.1'E, 40 m, 11 Februarv 1976, M.V Ko-
kinga (2, M.95642); N of Poor Knights Islands. 35°22'S.
174°43'E, coOected alive. 146 m. 15 February 1974. IW
Achcwn (many. M. 35115); W of Poor Knights Islands.
35°29'S, 174°43.5'E, coOected alive, 110 m,l5 February

1974, RA^ Achewn (18, M. 35229); W of Poor Knights
Islands, 35°32'S, 174°41'E. coOected alive, 121-113 m,
15 Februan- 1974, RA^ Achcwn (24, M.35876); off Poor
Knights Islands. 79 m, March 1971 (3, M.44715); off Poor
Knights Islands. 124 m. September 1965. M.V. Ikatcrc (6.
M.39638): off Hen Island. 36°00.5'S, 174°43'E, 59 m, 13
FebruaiT 1974, RA' Acheron (6, M. 43809); off Hen Is-
land. 48-55 m. J.A. BoOons (4, M.7242); off Trvphena.
Great Barrier Island (1. M.91031); off Cuvier Iskmd, 64
m and 70 m. J.A. BoOons (2. M.2165; 4, M.7241); 5 km
E of Channel Island, Cape ColviOe, 137 m (1, M.2164);
off Aldemien Islands, 165 m, 10 June 1970, B.L. God-
friaiLx (3. M.44386); off Aldennen Islands, 146 m, 26 May
1969, B.L. Godfriaax (3, M.44419); N of Aldennen Is-
lands. 36°47.5'S. 176°00.0'E. 108-113 m. 24 January
1981. RA' Tangaroa (1, M. 130068); SE of Aldermen Is-
lands. 37°00.8'S. 176°12.3'E, coOected alive, 178-248 m,
23 Januai-v 1979, RA' Tangaroa (many M.65151): N of
Mayor Island. 37°15.2'S, 176°14.5'E, 188-193 m, 22 Jan-
uary' 1979, RA' Tangaroa (1, M.6()979); off Mayor Island.
37°16.7'S. 176°17.5'E, 104-109 m. 22 January' 1979. RA'
Tangaroa (5. M.67411); off E side of Mayor Island,
37°18.9'S. 176°16.2'E, 59-74 m, 22 January' 1979, RA^
Tangaroa (5, M.65680); off Mayor Island. 37°22.5'S,
176°22'E, 207-220 m, 27 February 1957, M.V Alert (1,
M. 12887); off SE side Mayor Island, 37-91 m, February
1986, G. Nicholson (1, 'M.92093); off Wliite Island,
37°30.4'S. 177°09.7'E. coOected ;ilive. 8.3-92 m, 19 Jan-
uarv- 1979. RA' Tangaroa (4, M. 130070); Rungapapa
KnoO, WNW of White Island, 37°33.8'S, 176°59.0'E,
188-228 m, 20 January 1979, IW Tangaroa (1,
M. 113498); W of Plate Island. 37°39.1'S, 176°31.5'E. 64-
59m. 21 Januan- 1979. RA' Tangaroa (2. M.67638); be-
tween Motuhora and White islands. 37°40.2'S,
178°53.6'E. coOected alive, 117 m, 2 May 1990, RA' AAa-
(Icmik Alexander Ncsmeijanov (3, M. 112438); N of New
Plymouth. 38°40.2'S. 174°03.9'E. collected alive, 88 m,
13 January 1981, RA' Tangaroa (1, M. 130069); N of New
Plymouth', 38°40.4'S, 174°04.0'E, coOected alive, 88 m,
13 January 1981, RA^ Tangaroa (1, M. 130093); N of New
PKmoudi', 38°55.0'S. 174°09.3'E. 48 m, 13 January 1981,
RA' Tangawa (4, M. 130066).

Distribution (Figures 13, 48): Northern North Is-
land; west coast from Aliipara to New Plvmouth, east
coast from North Cape to White Island. 31-257 m; col-
lected alive at 31-c-a. 248 m from sand and mud.

Remarks: Among New Zealand Solariclh species. S.
plicatiila is ilistinctive in the combination of low spire,
wide umbilicus, strong spiral cords on spire and base,
and the color pattern of dull red sigmoidal bands on the
spire and base. The mtritacaLx of this species resolves as

B. A. Marshall, 1999

Page 9

ib°

Figure 13. Map of Three Kings Islands and northern tip of North Island, New Zealand showing distiibntions of species of
Solark'Ua (250 and 500 ni isobaths indicated): S. plicatiila (Mnrdocli and Suter) (A) and S. hilcola (Powell) (■).

conspicuous, lodal lines when the shell is (naturally)
lightly solution-etched (figure 3).

SolaricUa plicattila is an umbilical brooder, specimens
more than 2.5 mm wide ha\ing pronounced sexual di-
morphism in shell size, shape and umbilicus moq)holog\-
(figures 1-4). With increasing shell size, the umbilical
rim in females becomes strongly angulate instead of re-
maining weaklv angulate or exenlv rounded as in males,
the spiral thread at the umbilical rim becomes stronger
and higher, the mnbihcal wall becomes steeper and
more weakly sculptured or essentially smooth, the shell
becomes shghtly higher relative to its width, and the um-
bilicus niav liecome narrower than in some males,
though the volume (not measured) is greater because of
the steeper wall and the produced rim: shell/umbihcus
width ratio 1.70-2.30, mean = 2.08 (SD = 0.161, ii =
20) in females, and 1.70-2.40, mean = 2.03 (SD =
0.190, n = 20) in males. Females also attain larger size
than males (largest seen 7.00 mm wide, as against 5.50
mm). Note that in the boreal Pacific species Margaritas
vorticifenis (Dall, 1873), the onlv kniown trochid umbil-

ical brooder other than the present SolaricUa species,
the umbihcus is both wider and more voluminous in fe-
males than in males (Lindberg and Dobberteen, 19S1;
Lindberg, 1985). Dobberteen and Ellmore (1986) found
that shell chmoq^hism in Mariiaritcs vorticifenis is ex-
pressed even at the embiyonic stage, and although shell
dimorphism is not obvious in specimens of S. plicattila
less than 2.5 mm wide, it remains to be checked.

In a sample ot 240 specimens more than 2.5 mm wide
taken off Rangaunu Bay at 63 m depth (NMNZ
M. 130750), the female: male ratio is 1.12:1. Specimens
with yolk-y egg capsules (8 or 10) within the umbilicus
were obtiiined during the months of December January
(M.65151, M. 130067), and May (Havwird and Grace,
1981 - misidentified as S. vera), and shelled larvae oc-
curred during December and February (M. 35115,
M41401).

TJie radula (figure 125) is t\pical of the genus, with
liroad lateroniarginal plates. The jaw plates are extreme-
ly thin, with very weak elements along the leading edge
onl\-.

Paee 10

THE NAUTILUS, Vol, 113, No. 1

Figure 14. Map of Three Kings Islands and northern tip of North Island, New Zealand showing distributions of species of
Solarii'lla (250 and 500 ni isobaths indicated): S. vera (Powell) (■), S. exigiia new species (•) and S. basilica new species (o).

Solahclla vera (Powell, 1937)
(Figures 5-7, 14, 42, 126)

Zetninolia vera Powell. 19.37: 179, pi. 51, fig. .3

Spectamen venim. — Powell, 1976: 84; Powell, 1979: 64. pi. 17,

fig. 17.
NOT Spectamrn ventin. — Powell, 1979: 64 (remarks on Spec-
Uiinen only); Hay-ward and Grace, 1981: te.xt fig. 3; Hick-
man, 1992: 254 = Solariella plicatula.

Tj'pe iriaterial: HoloUpe (3.55 X 5.30 mm, 3.30 TW,
umbilicus 2.00 mm wide) BMNH 1962962, from t\pe
locality, Ki Angust 1932, R.R.S. Discovery II.

Type IochIjH: Off Spirits Bav, 34°21.0'S, 172°48.0'E,
59 m.

Other n>fitfv-«a£ .-ivamined (152 specimens NMNZ):

37 km NT:; of ( n^a! Island, Three Kings Islands,
33°58.0'S, ! V:"-.iO.G'i:. 550 m, 25 June 1978, RA' Tan-
garoa (28, U.li'Jyl-, Middlesex Bank, NW of Three
Kings Islands, 33"5 '..'S, 17r45.3'E, collected alive,
186-196 m, 31 janiarv 1981, RA' Tanoaroa (20,

M. 1.30064); Middlesex Bank, .34°02.1'S, 171°45.8'E, col-
lected alive, 221-206 m, 31 Januaiv 1981, R/V Tangaroa
(5, M. 1,30063): 22 km ENE of Great Island, Three Kings
Islands, 34°05.0'S, 172°24.6'E, 200 m, 24 June 1978,
RA' Tangaroa (92, M. 130061); .33 km E of Great Island,
.34°14.2'S, 172°.32.4'E, 100 m, 24 June 1978, RA' Tan-
garoa (7, M. 1.30065).

Distribution (Figure 14): Middlesex Bank, Three
Kings Islands, and off Spirits Bay, 59-550 m; collected
alive at 186-221 m from comminuted brvozoan/shell
substratum with .sponges, corals, hvdroids, etc.

Remarks: Solariella vera resembles S. plicatula in col-
or pattern and shell shape, but differs in having much
finer, more numerous spiral threads on the teleoconch.
With the exception of a few specimens from the extreme
south ol its range (see below). S. vera lacks the low,
rounded axial pleats on the spire and base characteristic
of S. plicatula. The protoconch in S. vera is typically
white. rareK' pale \ell(jw. The first one t)r hvo teleoconch

B, A, Marshall, 1999

Page 11

whorls may be yellow or white; subsequent whorls are
tvpically white with narrow, widely spaced, pale red axial
bands that are confined to the spire and do not extend
bcN'ond the peripherv; the liands are tvpicallv alisent
from a broad subsutural zone but do extend to the su-
ture in a few specimens (e. g., M. 130065). A few spec-
imens lack color bands. No specimens have been found
brooding voung within the umbihcus, though from the
size of the umbihcus, it mav also be an umbilical lirood-
er. The shell appears to be sexually dimorphic, putative
females ha\ing an angulate instead of rounded umbihcal
rim bordered Ijy a considerably stronger spiral thread
than in putative males (figures 5-7). Unlike S. plicatiila
and S. lutcola, which have sex ratios of about 1:1, the
ratio of putative females to males in S. vera would seem
to be more than 18.4:1. Whether the five putative males
are simply phenotspic \'ariants of females that otheiAvise
happen to resemble males in shell morphology is un-
clean Powell's (1979), Hayward and Grace's (1981), and
Hickman's (1992) records of brooding in S. vera are
based on misidentified specimens of S. plicatiila.

The radula (figure 126) is typical of the genus, with
rather narrow lateromarginal plates. The jaw plates are
thin, with rather well-developed elements covering most
of the central area from the leading edge almost to the
back edge.

Solaiiella hiteola (Powell, 1937)
(Figures 8-12, 13, 19, 43, 127)

Zeminolia hiteola Powell, 1937: 179, pi. .51, fig. 1, 2.
Spectamen luteolum. —?owe\\, 1976: 84; Powell, 1979: 64, pi.
17, fig. 1.5, 16.

Type material: Holotype BMNH 1962960 and para-
t)pe (1962961), from type locality, 17 August 1932,
R.R.S. Discovery II.

Type locality: Off Three Kings Islands, 34°13.3'S,
172°12.0'E, 260 m.

Other material examined (477 speeimen,s NMNZ):

Middlesex Bank, NW of Three Kings Islands, 33°57.0'S,
171°45,4'E, collected ahve, 98-103 m, 31 January 1981,
IW Tangaroa (20, M. 1.30051); King Bank, N of Three
Kings Islands, 33°57.0'S, 172°19.0'E, 128 m, 1 Febnian'
1981, RA' Tangaroa (9, M. 130048); King Bank.
33°57.4'S, 172°19.4'E, collected alive, 128-123 m, 1
February 1981, RA^ Tangaroa (11, M.72040); 37 km NE
of Great Island, Three Kings Islands, 33°58.0'S,
172°30.6'E, 550 m, 25 June 1978, RA' Tangaroa (4,
M. 130053); 18 km N of Great Island, 33°59.2'S,
172°13.6'E, 155 m, 23 June 1978, RA' Tangaroa (23,
M.93551); Middlesex Bank, 33°59.9'S, 171°45.3'E, 186-
196 m, 31 January 1981, RA^ Tangaroa (11, M. 1.30056);
Three Kings Trough, NW of Three Kings Islands,
34°00'S, 17r55'E, 805 m, 15 July 1962, R.N.Z.FA. Tni
(20, M. 1.30054); Middlesex Bank, .34°00.9'S, 17r44.7'E,
collected alive, 201-216 m, 31 Januan,' 1981, RA' Tan-
garoa (52, M. 130047); N of Three Kings Islands,
34°01'S, 172°07'E, 622 m, 18 Febmai>- 1974, WV Ach-

eron (2, M. 34620); Middlesex Bank, .34°01.2'S,
171°44.4'E, collected alive, 206-211 m, 31 January 1981,
RA^ Tangaroa (12, M. 130044); Middlesex Bank,
34°01.4'S, 17r45.2'E, collected alive, 201-216 m, 31
Januar\' 1981, RA' Tangaroa (5, M. 130052); 13 km N of
Great 'island, 34°()1.8'S, 172°12.0'E, 508 m, 23 June
1978, RA^ Tangaroa (7, M. 1.30046); Middlesex Bank,
34°02.0'S, 171°44.0'E, 246-291 m, 31 Januai-v 1981, R/

V Tangaroa (44, M. 130050); 37 km NW of Great Island,
34°()2.0'S, 171°48.4'E, 188 m, 27 June 1978, RA' Tan-
garoa (7, M. 130049); Middlesex Bank, 34°02.1'S,
17r45.8'E, collected alive, 221-206 m, 31 Januarv 1981,
RA' Tangaroa (50, M. 1.30055); 22 km ENE of Great
Island, 34°05.0'S, 172°24.6'E, 200 m, 24 June 1978, R/

V Tangaroa (43, M. 130059); off Northeast Island, Three
Kings Islands, .34°08.5'S, 172°11'E, collected alive, 102
m, 18 FebruaiT 1974, RA' Acheron (7, M.34254); off
Three Kings Islands, .34°10'S, 172°12'E, 252 m, 19 Feb-
i-uaiy 1974, WV Acheron (9, M.34019); off Three Kings
Lslands, .34°11'S, 172°10'E, 91 m, 19 Febmar\' 1974, W
y Acheron (3, M..337.53); SE of Great Island, ■34°14.1'S,
172°09.0'E, collected alive, 192-202 m, 1 February
1981, RA' Tangaroa (10, M. 130057); NW of Cape Rein-
ga, 34°14.2'S, 172°32.4'E, 100 m, 24 June 1978, RA'
Tangaroa (35, M. 130058); SE of Great Island, ;34°14.8'S,
172°13.6'E, 173-178 m, 2 Felmiai-v 1981, RA/ Tangaroa
(9, M. 130091); 39 km SW of Great Island, 34°17.6'S,
171°45.3'E, 427 m, 21 June 1978, RA' Tangaroa (22,
M. 130043); 20 km NW of Cape Maria van Diemen,
34°20.0'S, 172°30.0'E, collected alive, 100 m, 20 June
1978, RA' Tangaroa (15, M.93744); SE of Three Kings
Islands, 34°20.2'S, 172°21.8'E, collected ahve, 121 m, 2
February 1981, RA' Tangaroa (21, M. 130090); 37 km
SW of Great Island, 34°20.4'S, 17r48.2'E, 440 m, 21
June 1978, RA' Tangaroa (18, M. 130045); N of Cape
Reinga, 34°21'S, 172°37'E, 88m, 19 Februai>' 1974, R/

V Acheron (4, M.35915); 28 km S of Great Island,
34°24.0'S, 172°16.8'E, collected alive, 120 m, 20 June
1978, RA' Tangaroa (4, M. 1.30092).

Distribution (Figure 13): Middlesex Bank, King
Bank, Three Kings Islands, and off Cape Reinga, 88-
805 m; collected ahve at 98-202 m from comminuted
bryozoan and shell substrata with sponges, hydroids, cor-
als, etc.

Remarks: Solaiiella hiteola is rather variable in sculp-
ture, protoconch size, color and color pattern. The spire
mav be sculptureil throughout with weak, crowded spiral
threads, or all spirals other than a subsutural spirid may
become obsolete on the second teleoconch whorl. The
protoconch mav be white or chrome vellow and ranges
in width from 500 to 620 |xm (mean = 560 jjim). The
first teleoconch whorl m;i\ he chrome vellow or pinkish
white, and subsequent whorls are either uniform white
or pinkish white or pinkish buff with the addition of
narrow, pale red sigmoidal axial bands that entirely tra-
verse the whorls to the umbilical rim, extending onto
the umbilicid wall in some specimens. Specimens from
shallower than 200 meters are tvpicalK' the most strongly

Page 12

THE NAUTILUS, Vol. 113, No. 1

sculptured and pigmented, whereas specimens from
depths greater than 400 meters are tx-pically uniform
white and weakK sculptured.

The radula (figure 127) is closest to that of S. plica-
tula, differing in having more broadly angulate cutting
areas on the lateral teeth. The jaw plates are extremely
thin, xvith weak elements covering the anterior third.

S. hitcola broods its voung within the umbiUcus and
has sexually dimorphic shells, shells of females having a
more voluminous umbihcus with a flatter, steeper wall
than males, and a more prominent spiral cord at the
umbilical rim (figures 8-12). The umbifical rim is strong-
ly angulated in both sexes, unlike S. pUcatiila in which
this is a characteristic of females alone. One specimen
examined had a shelled embrvo deep within the umbi-
licus, and another had two (M. 13005.5), though to judge
from the volume of the umbilicus, the number of young
brooded at any one time is probably considerably greater
than this.

Compared with S. vera, which it most closelv resem-
bles, S. luicola thffers in that spiral sculpture on the spire
is more or less obsolete or consists of finer, closer, more
numerous spiral threads, in that the protoconch is gen-
erallv larger (mean = width 560 \x.m. cf 510 jim), and
in that the color bands (when present) are more strongly
sigmoidal and tra\'erse the spire and the base instead of
only the spire. Solariella hitcola and S. vera have been
taken together at se\'eral stations throughout their rang-
es, living svmpatricallv at 221-206 meters depth on Mid-
dlesex Bank (M. 130055, M. 130063). Specimens from
the south side of the Johnson Trough (figure 13;
M. 130092, M. 130058, M.93744, M. 130090, M.35915),
between the Three Kings Islands and Cape Reinga, at-
tain smaller size than northern shells, and a few of them
have rounded axial folds, characteristics that are en-
hanced to the southeast. Southern specimens with axial
folds resemble S. plicatula. though thev are otherwise
indistinguishable from northern populations of S. hitcola
and they are not strictly moq^hologically intermediate.

Solariella cxigtia new species
(Figures 14-17, 21, 44)

Description: Shell up to 4.60 mm wide, wider than
high, of moderate thickness, widelv umbilicate. Proto-
conch white; first 1.2-1.3 teleoconch whorls yellow; sub-
sequent whorls white with dull red, irregular, tvpically
sub(^uadrate, axial maculations on abapical two thirds ot
'ivr- wliorls Protoconch 430-480 fjLm wide, sculptured
u.' .'ranules and a fine spiral thread, tip of apical
'•i. . -lU rounded. Teleoconch up to 3.0 convex
V.;.:.'. ::•■- Ahorl rather evenly convex, subsequent
wiit'i' •. ^'' '. ;i,:ontal or outwardly adapically sloping
sutu'.,' ■. :, ) ■ t ;idult whorl sUghtlv flattened above
and l> ■rv. ;r,>led periphery. Primarv sculpture of
proininent ■-•■■.;;] r 'reads that multiplv by intercalation,
8-13 on spi:t: asH' 16-25 on base in adults, innermost
basal spiral at uniiiijicul rim; spire spirals stronger and

angulate in section, with interspaces wider than each spi-
ral; basal spiral finer and rounded in section, with inter-
spaces narrower than each spiral except beside umbilical
rim. Secondare' sculpture of weak axial pleats confined
to adapical third on spire on last adult whorl, fine col-
labral growth fines throughout. Unibificus perspective to
protoconch, diameter 41—46% of adult shell diameter,
rim angulate; wall sculptured with spiral threads and fine
collabral growth lines, last whorl more or less flat over
outer two thirds. Aperture subcircular, fips thin and sim-
ple, thickest at angulation of umbilical rim.

Type material: Holotype NMNZ M. 131480 and 6
paratvpes NMNZ M.43711, from tvpe locafitv', IS Feb-
ruarv'l974, WW Acheron. Paratvpes (16 NMNZ): middle
Southeast Bay, Great Island, 34°10'S, 172°08'E, collect-
ed afive, 42-46 m, 18 Februarv 1974, RA' Acheron (14
NMNZ M.33833, 1 AMS), Southeast Bay, Great Island,
30 m, 5 February 1993, F. Brook (2, M. 117426).

Type locality: Middle Southeast Bay, Great Island,
Three Kings Islands, 34°10'S, 172°08'E, 42-46 m.

Other material examined: Off Rangaunu Bay,
.34°49.6'S, 17.3°15.0'E, 23 m, 27 Januan- 1981, RA' Tan-
garoa (7, M. 1.30071).

Distribution (Figure 14): Off Three Kings Islands,
and off Rangaunu Bay, 23-55 m, living at 42—46 m on
gravel; dead shells on comminuted brvozoan/shell or
shell substrata.

Remarks: Solariella exigua most closely resembles S.
plicatula and S. t era, differing from both in having finer,
closer, more numerous spiral threads on the spire, a
smaller protoconch (width 430-480 |i,m: mean = 462
(xm, cf 470-530 |xm: mean = 498 (xm and 500-530 \im:
mean = 510 \i.m), and in attaining smaller size (maxi-
mum width 4.60 mm, cf 7.00 mm and 5.20 mm). It
differs further from both in showing no obvious sexual
dimorphism in the shell.

Etymology: Little (Latin).

Solariella basilica new species
(Figures 14, 20, 22, 23)

Description: Shell up to 8.90 mm wide, wider than
high, of moderate thick-ness, widely umbificate. Proto-
conch white; first 1.0-1.3 teleoconch whorls vellow, fad-
ing over next half whorl to white, subsequent whorls
uniform white. Protoconch 600 |xm wide, smooth, tip of
apical fold bluntly rounded. Teleoconch up to 3.75 con-
vex whorls; first whorl rather evenlv convex, subsequent
whorls with horizontal sutural ramp with rounded rim,
last adult whorl slightlv flattened above and below
rounded peripherv. Priman' sculpture of fine, crowded
spiral threads that multiply by intercalation, angulate in
section on spire, rounded in section on base, 17 on pen-
ultimate whorl and about 30 on base (subadult holo-
tvpe), 20 on penultimate whorl and 22 on base (adult
paratype), innermost basal spiral at umbifical rim. Fine

B. A. Marshall. 1999

Page 13

Figures 15-29. Slii^lLs of species of Solariclla. 15-17, 21. S. cxi'^iiti new species. 15, 17, 21. HoloUpe, Southeast Bay. Great
Island, Three Kings Islands. 42-46 in, M. 131480 (2.40 x 4.00 nun). 16. Paratvpe, Southeast Bay, 55 ni, M.33S33 (2.90' x 4.55
mm). IS. S. pUeatiila (Murdoch and Suter), off Poor Knights Islands, 146 m, M. 35115. 19. S. hiteola (Powell), off Northeast Island,
Three Kings Islands, 102 m, M.34254. 20, 22, 23. S. basilica new species, holotype. Southeast Bay, 55 m, M. 131557. 24, 25. .S.
semireticulata (Suter). 24. E of Fort\- Fours, Chatham Islands, 238 m, M. 10837. 25. SE of Nugget Point, Otago, 140 m, M.59359
(3.55 X 4.80 mm). 26-29. S. tnjphenensis (Powell), off Te Aral Point, 35 m, M. 130840 (4.30 X 5.40 mm). Scale 28 = 1 mm.
others = 250 (j.ni.

Page 14

THE NAUTILUS, Vol. 113, No. 1

coUabral growth lines throughout. Umbilicus perspective
to protoconch, diameter 33% (adult)-38% (subadults) of
shell diameter, rim angulate, projecting inwards at ma-
turity to overhang outer umbilical wall: wall of each
whorl steep, sculptured with low spiral threads and fine
coUabral growth lines. Aperture subcircular, Ups thin and
simple, thickest at angulation of umbilical rim.

Tj'pe material: Holotvpe (2.50 X 3.80 mm, subadult)
NMNZ M. 131557 and 2 subadult paratypes (M.33828),
from tvpe localit)', 18 FebruaiT 1974, RA^ Aclicron. Par-
atopes: Middlesex Bank, NW of Three Kings Islands,
33°59.9'S, 17r45.3'E, 186-196 m, 31 January 1981, R/
V Tangawa (1 adult, M. 131481); NW of Cape Reinga,
34°14.2'S, 172°32.4'E, 100 ni, 24 June 1978, RA' Tan-
garoa (1 subadult, M. 131539).

Type locality: Outer Southeast Ba\-, Great Island,
Three Kings Islands, 34°10'S, 172°08'E, 55 m.

Distribution (Figure 14): Off Great Island, Three
Kings Islands, and Middlesex Bank, 5.5-196 m (dead),
on gravel and comminuted br)ozoan/shell substrata.

Remarks: Solariella basilica resembles S. exigua in
having crowded spiral threads on the spire and base but
differs in having finer, more numerous threads at equiv-
alent stages of shell growth, in attaining much larger size
(ma.ximum width 8.90, cf 4.60 mm), in having a wider
protoconch (600 |xm, cf 430—480 (xm) and a narrower
umbiLcus (33-38% instead of 41-46%c of shell width),
and in the last few teleoconch whorls being imiform
white instead of axially banded. Solariella basilica differs
from S. vera in having finer, more numerous spiral
threads on the spire and base (basal spirals obsolete in
S. vera). Empty shells of S. basilica occurred togedier
with S. exigtta at the type locahts' and with S. vera at
Middlesex Bank (M. 131481, M. 130064).

Etymology: Roval (Greek).

Solariella semireticulata (Suter, 1908)
(Figures 24, 25, 48)

Monilea (Minolia) semireticulata Suter, 1908: 22. pi. 2, fig. 1;

Suter, 191.3: 142, pi. .33, fig. 13.
Spectamen semireticulata. — Powell, 1926: 44, 4.5.
Zeminolia semireticulata. — Finlay, 1926: 360; Powell, 19.5.5: .55.
Spectamen semireticidatum. — Powell, 1976: 84; Powell, 1979:

64. pi. 17, fig. 18.

Type material: Lectotype (Boreham, 19.59) NZGS
TM467.

Type U>cai'>y: Off the Snares Islands, 91 m.

Other ik;;'^' rial examined (88 specimens NMNZ):Peg-
asus Canvor NE of Banks Peninsula, 43°31.0'S.
173°30 5'E, oUected afive, 256-293 m, 21 Februaiy
1979, RA" Al,;:'!V.i (i, M.94717); Papanui Canyon, off
Otago Peni-isuia, 4.5''51'S, 171°00'E, .348-220' m, 15
Februar)- 1S79. 1 ■ A' Acheron (2, M.94629); Saunders
Canyon,' (jff East ■::''u-:o, 45°56'S, 170°54'E, 360 m, 30
June 1975, W\^ Munida (15, M.4S847); offTaiere, Dun-

edin, 46°12.0'S, 170°41.5'E, 150 m, 1 September 1976,
RA^ Miinida (2, M.65802); Saunders Canyon, off East
Otago, 45°56'S, 170°54'E, 420 m, 27 October 1967, R/
V Miinida (1, M.28426); SE of Nugget Point, East Ota-
go, 46°40'S, 170°00'E, 140 m, 27 March 1978, RR.V.
James Cook, coll. L. Paul (20, M.59359); 0.8 km SW of
Hare's Ears, Doubtful Sound entrance, 45°17.2'S,
166°49.3'E, collected alive, 146 m, 7 Januai>- 1977, RA'
Acheron (24, 58783); E of Fom- Fours, Chatham Is-
lands, 44°04'S, 17.5°23.5'W, collected afive, 238 m, 1
February 1954, M.V. Alert (13, M. 10837).

Distribution (Figure 48): East coast of South Island,
Doubtful Sound, Chatham Rise, also Snares and Auck-
land islands (Powell, 1955), 91-i21 m; collected ahve at
146-238 m from sand and shell substrata.

Remarks: Solariella .semireticulata differs from the su-
perficially similar species S. plicatiila in having stronger
teleoconch sculpture, in being uniform white, and in that
the umbifical rim is consistently rounded. Unfike S. pli-
catula, S. .semireticulata exliibits no obxdous shell dimor-
phism and is probably not an umbifical brooder.

Solariella trijphenensis (Powell, 1930)
(Figures 26-29, 45, 49)

Zeminolia tnjphenensis Powell, 1930: 534, pi. 62, fig. 16-18.
Spectamen trtjphenense. — Powell, 1976: 84; Powell, 1979: 65,
pi. 17, fig. 12-14.

Type material: llolotype AIM AK72175, from t\pe
locality.

Type locality: Tiyjihena Bay, Great Barrier Island, 11
ni.

Other material examined (many specimens NMNZ):

Great Exhibition Bav, 34°33.4'S, 17.3°04.8'E, 63 m, 27
Januaiv 1981, RA^ fangaroa (51, M. 113337); off Twin
Rocks," Bay of Islands, 35°10'S, 174°18'E, 46-73 m, 10
December 1973, M.V. Kokinga (2, M.42245); off Mo-
tuwheteke Island, Bav of Islands, 35°12'S, 174°17'E, 22-
31 m, 14 December' 1973, M.V. Kiikinga (1, M.41S64);
SE of Te Arai Point, Mangawhtu, Northland, 36°10.5'S,
174°42.0'E, collected alive, 35 m (dredged for beach re-
plenishment at Mission Bav, Auckland) 3 March 1996,
B.A. Marshall and N.J. Peterson (10, M. 130840), B.R
Hazelwood, 1996 (51, M. 131392), P Poortman (many,
M. 131712); off Mavor Island, 37°16.7'S, 176°17.5'E,
104-109 m, 22 Januarv 1979, RA' Tangaroa (10,
M.67431) off E side of Mavor Island, 37°18.9'S,
176°16.2'E, 59-74 m, 22 Januarv 1979, RA^ Tangaroa
(102, M.65681).

Distribution (Figure 49): Northeastern North Is-
land, from Great Exhibition Bay to Mayor Island, 11-
109 m, collected iilixe at 35 m on clean sand and shell
substratum.

Remarks: Among New Zealand Solariella species, S.
tnjphenensis is characterized by its large size and strong

B. A. Marshall, 1999

Page 15

Figures 30-40. Shells of species of Solariclhi. 30-33. S^ henthicola (Powell), Middlesex Bank, 221-206 iii, M. 130079 (5.30 X
5.80 mm). 34-37. S. peristicta new species, holotype, NW of Cape Reinga, 78 m, M. 131563 (4.55 X 5.05 mm). 38, 39. S. flavida
new species, holot>'pe, NW of Cape Reinga, 100 m, M. 131.564 (3.30 X 4.40 mm). 40. S. amabilis (Jeffreys), Snorre oil field, N
North Sea, 292-.340 m, Swedish Musenni of Natnral History (4.20 X 4.30 mm). Scales 32, .36 = 1 mm, others = 250 |xm.

.sculpture, and in heint; pale huff with a deep red (sel-
dom brown) color pattern of suhsutural niacidations and
irregular, wavy axial bands that extend from the macu-
lations to about midway across the base. S. tn/phcncnsis
ranges into shallower water than other New Zealand
species, perhaps as shallow as 11 meters, and it has an
extremely patchy distribution.

SolaricUa henthicola (Powell, 1937)
(Figures 30-33. 46, 48, 12S)

Zriniimliii hcnlliicola Powell, 1937: ].S0, pi. 51, fig. 4;
Spccfaincn hciiihicol/i. —VoweW. 1976: 84; Powell, 1979: 64.

Type material: Holot\pe BMNH 1962963, from type
locaht), 17 August 1932, R.R.S. Discoicnj II.

Page 16

THE NAUTILUS, Vol. 113, No. 1

Figures 41—17. Shells ot species of Solariella. 41. S. plicatula (Murdoch and Suter), off Rangaumi Bd\. 6.3 m, M. 1.30750 (4.00
X 5.6.5 mm). 42. S. vera (Powell), ENE of Great Island. Three Kings Islands, 200 m, M. 1.30061 (5.05 X '7.50 mm). 43. S. luteola
(Powell), Middlesex Bank, 9S-103 m, M. 1.30051 (4.40 X 6.40 mm). 44. S. exigua new species, paratope. Southeast Bay, Three
Kings Islands, 55 m. M.338.33 (2.S5 X 4.50 mm). 45. S. tnjpheiwnsis (Powell), off Te Arai Point, .35 m, M. 1.30840 (4.80 X 5.70
mm). 46. S. benthicola (Powell). Middlesex Bank, 221-206 m, M. 1.30079 (5.-35 X 5.60 mm). 47. S. peri.sticta new species, NW of
Cape Reinga. 78 m, M. 131492 (4.15 X 4.60 mm).

Type locality: Off Three Kings Islands, 34°13.3'S,
172°12.0'E, 260 m.

Other material examined (114 speeimen.s NMNZ):

18 km N of Great Island, Three Kings Islands,
33°59.2'S, 172°13.6'E, 1.55 m, 23 June 197S, IW Tau-
garoa (4, M. 1.30073); Middlesex Bank. NW of Three
Kings Islands, .33°.59.9'S, 171°45.3'E, 186-196 ni, 31
January 1981, IW Tangaroa (15, M. 130076); Three
Kings Trough, NW of Three Kings Islands, 34°00'S,
17r.55'E, 805 m, 15 Julv 1962, R.N.Z.RA. Tui (30,
M.48959); Middle.sex Bank, .34°0O.9'S, 171°44.7'E, col-
lected ahve, 201-216 m, 31 January' 1981, IW Tangaroa
(2, M. 130077); N of Three Kings Islands, .34°01'S
172°07'E, 622 m, 18 Felmiarv 1974, IW Acheron (2
M. 1.30083); 13 km N of Great Island, .34°01.8'S
172°12.0'E, .508 m, 23 June 1978, RA^ Tangaroa (6
M. 131501); Middlesex Bank, .34°02.0'S, 17r44.0'E
246-291 m, 31 January 1981, RA^ Tangaroa (10
M. 1.30081); 37 km NW of Great Island, .34°02.0'S
17r48.4'E, 188 m, 27 June 1978, IW Tangaroa (2
M. 130075); Middlesex Bank, .34°02.1'S, 171°45.8'E, col-
lected alive, 221-206 m, 31 January 1981, RA' Tangaroa
(5, M. 1.30079); SE of Great' Island, .34°14.1'S,
172°09,0'E, collected alive, 192-202 m, 1 Februarv
1981, K\ Tangaroa (4, M. 1.30078); NW of Cape Reinga,
.34°14.2'S, 172°.32.4'E, 100 m, 24 June 1978, RA^ Tan-
garoa {.2., M. 130074); 39 km SW of Great Island,
-! i7,6'S, 17r45.3'E, 427 m, 21 June 1978, RA^ Tan-
gur-y; 1, M. 130082); N of North Cape, .34°20.0'S,
r7:V()(i 6E, collected alive, 16.3-168 m, 27 JanuaiT 1981,
R/V Tangiiroa (S, M. 131502); SE of Three Kings Islands,
34°20.2'S, 172"21.8'E, collected alive, 121 m,"2 Febru-
ary- 1981, HA' Tangaroa (3, M. 130080); 28 km S of Great
Island, 34°24.0'S,' 172n6.8'E, 120 m. 20 June 1978. W
V Tangaroa (1, M. 130072); E of North Cape. 34°25.0'S,

173°13.1'E, 327-257 m, 27 Januaiy 1981, RA' Tangaroa
(22, M.48959).

Distribution (Figure 48): Off Three Kings Islands
and off North Cape, 100-805 m, collected alive at 121-
216 m on comminuted brvozoan/shell substratum with
sponges, corals, hydroids, etc.

Remarks: Among species from the New Zealand re-
gion, Solaiivlla benthicola most closely resembles S. tnj-
phencnsis, differing principally in being smaller relative
to the number of whorls (maximum wdth 5.90 mm, 4.3
TW cf. 6.70 mm, 4.1 TW); in ha\ing crisp axial riblets
rather than low, rounded axial folds (jn the spire; and in
being more sparsely pigmented. Specimens from the ex-
treme south of its range and many from extreme depths
have stronger, more widelv spaced axial riblets than oth-
er material but seem likely to be conspecific.

The radula (figure 128) is tvpical of the genus, with
rather narrow lateromarginal plates. The elements on
the extremely thin jaw plates are weak and confined to
the anterior half, weakening posteriorly.

Solariella pcristkia new species
(Figures .34-37, 47, 49, 129)

Description: Shell up to 5.80 mm wide, slightly wider
than high, of moderate thickness, umbilicate, almost lus-
treless. Periostracum apparently calcified (intritacalx).
Protoconch white. Teleocoiich pale biift or white, spiral
cords widi small, irregular yellowish brown or pale red
spots; paler, irregular, wavy axial bands on base and outer
part of umbilicus; some specimens pinkish white with
irregular, wavy, red axial bands that extend from suture
into umbilicus; greenish nacre showing through trans-
lucent outer shell layers. Protoconch 130 jjim wide.

B. A. Marshall. 1999

Page 17

Figure 48. Map of New Zealand region showing distribu-
tions of species of Solariella: S. plicatuln (Murdoch and Suter)
(•), S. semireticulata (Suter) (A), and S. benthicola (Powell)
and S. floviclfi new species {ie).

sculptured with 4 fine, crisp spiral threads and sparse
granules, tip of apical fold pinched. Teleoconch up to
4.25 whorls. First whorl evenly convex, subsequent
whorls with horizontiil sutural ramp with angulate rim.
rather evenly conve.x from ramp rim to suture within
umbilicus. Primarv sculpture of prominent, narrow spiral
cords that multiply by intercalation and enlarge to re-
semble adjacent spirals, interspaces several times wider
than each spiral; 4 on first 1.5 spire whorls, adapical 3
similar, abapical spiral more or less covered bv succeed-
ing whorls; adults with 6 on penultimate whorl and 11-
14 on base and umbilical wall. Second and subse(juent
whorls traversed by low, rounded axial pleats, strongest
on adapical half of each spire whorl, evanescent abapi-
cally. Fine, crisp, crowded collabral growth lines
throughout; fine, crisp, interstitial spiral threads com-
mencing after second whorl, becoming crowded
throughout. UmbiUcus perspective to protoconch, rim
evenK' roimded, wall of each whorl con\'ex, sculptured
with spiral cords and fine collabral and spiral threads.
Aperture subcircular, lips thin and simple.

Radula (figure 129) tyj^ical of the genus, lateromar-

ginal plates rather narrow. Jaw plates ven thin, weak
elements confined to the anterior half.

Type material: Holotype NMNZ M. 13156.3 and 3
paratopes M. 13 1492, from type locality, 2 February
1981, WV Tanooroa. collected alive. Parafspes (14):
Middlesex Bank, NVV of Three Kings Islands, ■33°57.0'S,
17r45.4'E, collected alive, 98-103 m, 31 Januarv 1981,
IW Tangaroa (5, M. 131493): King Bank, N of Three
Kings Islands, 33°57.4'S, 172°19.4'E, collected alive,
128-123 m, 1 February 1981, RA^ Tangawa (8 NMNZ
M. 131494, 1 AMS).

Type locality: NW

172°27.8'E, 78 m.

of Cape Reinga, 34°25.0'S,

Other material examined (100 specimens NMNZ):

King Bank, N of Three Kings Islands, 33°57.4'S,
172°19.4'E, 128-123 m, 1 February/ 1981, RA' Tangawa
(13, M. 130084); Three Kings Trough, N\\' of Three
Kings Islands, .34°00'S, 171°55'E, 805 m, 15 Julv 1962,
R.N.Z.EA. Tui (5, M. 130085): N of Three Kings Islands,
34°01'S, 172°07'E. 622 m, 18 Februarv 1974,' RA^ Ac/i-
cron (6, M.34619); Middlesex Bank, NW of Three Kings
Islands, 34°01.2'S, 17r44.4'E, 206-211 m, 31 January
1981, RA' Tangaroa (3, M. 131496); Middlesex Bank,
34°02.0'S. 171°44.0'E, 246-291 m, 31 Januarv 1981, R/
V Tangaroa (5, M. 131495); 11 km NW of Great Lsland,
Three Kings Islands, 34°06.5'S. 172°04.7'E. 310 m, 30
June 1978, RA' Tangaroa (2, M.61461); off Northeast
Island, Three Kings Lslands, 34°08.5'S, 172°11'E, col-
lected alive, 102 m, 18 February 1974, WW Acheron (13,
M.34253); outer Southeast Bav, Great Island, 34°10'S,
172°08'E, 55 m, 18 FebmaiT 1974, IW Acheron (4,
M.33825); off Three Kings Islands, 34°10'S, 172°12'E,
252 m, 19 Februaiy 1974. IW Acheron (5. M.34024);
Arch Pinnacle, Princes Islands, Three Kings Islands, 40
m, 10 March 1983, F. Brook (1, M. 109336): 39 km SW
of Great Island, 34°17.6'S, 171°45.3'E, 427 m, 21 June
1978, RA' Tangaroa (5, M. 131500); N of Cape Reinga,
34°21'S, 172°37'E, 88 m, 19 Febniaiv 1974, WV Ach-
eron (4, M. 35912): off RangaunuBay, 34°42.8'S,
173°14.5'E, 63 m, 27 January 1981, RA' Tangaroa (32,
M. 130751); NW of Whangaroa Harbour, 34°54.0'S,
173°42.6'E, 83 m, 26 Januan' 1981, RA' Tangaroa (1,
M. 130768); ENE of Great Barrier Island, \icinit\- of
35°43'S, 176°18'E, 329 m, 21 July 1962, R.N.Z.F.A.' Titi
(1, M. 44047).

Distribution (Figure 49): Off Three Kings Islands,
and northeastern North Island as far south as Great Bar-
rier Island, 40-805 m, collected alixe at 78-128 m on
comminuted bryozoan/shell substrata with sponges, cor-
als, hydroids, etc.

Remarks: Compared with Solariella tn/plicnensis,
which it most resembles, S. peristicta differs in attaining
smaller size (width up to 5.80 mm cf. 6.70 mm) and in
having crisper, more crowded collabral axial riblets on
the last few whorls that protluce a fine reticulation with
the secondarv spiral threads. Solariella peristicta resem-

Page 18

THE NAUTILUS, Vol. 113, No, 1

Figure 49. Map of New Zealand region showing distribu-
tions of species of Solariella: S. tn/pbcncnsis (Powell) (A) and
S. pcristicta new species (■).

bles the svmpatric species S. hcntliicola in size, shape,
and radular morpholog)' but has much coarser axial
sculpture.

Etymology: Dappled (Greek).

Solariella jiatida new species
(Figures 38, 39, 48)

Description: Shell up to 5.05 mm wide, sBghtly wider
than high, ol moderate thickness, umbihcate. Periostrac-
um apparently calcified (intritacabc). Protoconch white.
First 2.5 teleoconch whorls \ellow. sutural ramp white;
suf)sequent whorls white with pale yellow subsutural
Tiaciilafions and pale yellow, irregular axial bands that
>.'•;;• •i^•^^, ■■:■ below maculations and extend to either
aLo"' ini.iwuv across base or deep into umbilicus. Pro-
toconch iO'J-43fJ (xm wide, sculptured with 5 fine, crisp
spiral frM^-.i-Jj ;!nd crowded granules, tip of apical iold
moderal-l. ;:;,)i hed Teleoconch up to 3.75 whorls with
narrow, moie or less liorizontal sutural ramp; rather
evenly convex troin /amp to suture within umbilicus. Pri-
mary sculpture of prominent, narrow spiral cords that
multiply by intercalation and enlarge to resemble adja-

cent spirals, strong on spire, strongest on inner third of
base and umbifical wall, interspaces several times wider
than each spiral, finer and crowded on outer part of
base; 4 on first spire whorl, adapical 3 similar, abapical
spiral more or less covered by succeechng whorls; adults
with 8 or 9 on penultimate whorl, 20-25 on base and
umbihcal wall. Second and subsequent whorls traversed
by weak, inconspicuous, rounded axial pleats, strongest
on spire, evanescent on base. Fine, crisp, crowded col-
labral growth fines throughout; fine, crisp spiral threads
commencing after first whorl, becoming crowded and
covering interspaces on spire, base, and umbilical wall.
Umbihcus perspective to protoconch, rim evenly round-
ed, whorl of each whorl convex, sculptured with spiral
cords and fine collabral and spiral threads. Aperture sub-
circular, lips thin and simple. Radula unknown (only
available live-taken specimens decayed).

Type material: Holotvpe NMNZ M. 131564 and 29
paratvpes (28 NMNZ MT31512, 1 AMS), from t\pe lo-
cality, 24 June 1978, RA' Tangaroa, collected alive. Par-
at\pes (15 NMNZ): Middlesex Bank, NW of Three
Kings Islands, 3.3°57.0'S, 17r45.4'E, 98-103 m, 31 Jan-
uarf 1981, RA' Tangaroa (3, M. 131508); Middlesex
Bank, 33°59.9'S. 17r45.3'E, 186-196 m, 31 Januarv
1981, RA^ Tangaroa (8, M. 131507); Middlesex Bank,
34°01.4'S, 171°45.2'E, 201-216 m, 31 JanuaiT 1981, R/

V Tangaroa (1, M. 131510); Middlesex Bank, ■34°02.1'S.
171°45.8'E, 221-206 m, 31 January 1981, RA' Tangaroa
(2, M. 131511); 37 km NW of Great Island, Three Kings
Islands, 34°02.0'S, 17r48.4'E, 188 m, 27 June 1978, R/

V Tangaroa (1, M. 131509).

Type locality: NW of Cape Reinga, 34°14.2'S,
172°32.4'E, lo'o m.

Other material examined (34 specimens NMNZ):

King Bank, N of Three Kings Islands, 33°57.4'S,
172°19.4'E, 128-123 m, 1 February 1981, RA' Tangaroa
(5, M.72047); 37 km NE of Great Island, Three Kings
Islands, 33°5S.0'S, 172°.30.6'E, 550 m, 25 June 1978, R/

V Tangaroa (3, M.93904); Middlesex Bank, NW of
Three Kings Islands, 34°01.2'S, 171°44.4'E, 206-211 m,
31 Januarv 1981, RA' Tangaroa (2, M. 131506); Middle-
sex Bank,'34°02.0'S, 171°44.0'E. 246-291 m, 31 Januar\-
1981, RA' Tangaroa (14, M. 131505): 22 km ENE of
Great Island, .34°05.0'S, 172°24.6'E, 200 m, 24 June
1978, IW Tangaroa (8, M.93456); off Three Kings Is-
lands, .34°10'S, 172°12'E, 252 m, 19 Febniarv 1974. IW
Acheron (1, M.34020); NW of Cape Reinga, 34°22.8'S,
172°24.6'E, 121 m, 2 Febmary 1981, WV Tangaroa (1,
M. 131503).

Distribution (Figure 48): Off Three Kings Islands
and off Cape Reinga, 98-550 m, collected alive at 100
m (ui comminuted bryozoan/shell substratum with
sponges, corals, hvdroids, etc.

Remarks: Compared with Salarwlla tryplicncnsis and
S. pcristicta, which it resembles in shape, S. flavida dif-
fers principallv in ha\ing finer priman- spiral cords on

B. A. Marshall, 1999

Page 19

Figures 50-61. Shells of species oi BathipnophHa. 50-53. B. gravida new species 50, 51. HoloUpe, N Tliree Kings Rise, 1137-
1150 m. NZOI H6S0 (9.50 X 12.6 mm). .52, 53. ParaUpe. N Three King-s Ri,se, 1216-13S5 m, NZOI PI 123 (10.0 X 13.4 mm).
54, 55. B sp. cf. oravida new species, N Three Kmgs Rise, 1216-1385 m, NZOI U602 (12.0 X 13.0 mm). 56, 57, 59. B. callomphala
(Schepman), paralectot\pe, Sulu Sea, 4.50 m, ZMA 3.08.050 (4.60 X 6.50 mm). 58, 60, 61. B vnlentia new species, hol()t>pe, N
Three Kings Rise, 157()-1,563 m, NZOI H681 (7.40 x 8.50 mm). Scale .52 = 250 |xm, others = 1 mm.

the spire (8 or 9, cf. 5-8 and 6, respectively, on penul-
timate whorl in adults) and in having a yellow color pat-
tern. S. flavida and S. pcristkia have been taken togeth-
er as empty shells at five stations, suggesting that they

live in close pro.ximitv to one another, it not s)inpatri-
cally.

Etymology: Yellowish (Latin).

Page 20

THE NAUTILUS, Vol. 113, No. 1

Genus Bathijmophila Dall, 1881

Bathi/mophila Dall, 1881: 102. Tyjie species (by monotypy):
Margarita (Batlujmophila) euspira Dall, 1881; Recent, At-
lantic.

Ethaliopsis Schepman, 1908: 51. T\pe species (bv monotypy):
Solariclln lEtlwIiopsis) callomphala Schepman, 1908; Re-
cent, Indonesia.

Remarks: The radula of the tvpe species of Bathi/mo-
pliilii is unknown, and Batlu/mi)plula has traditionally
been referred to Margaritinae because the shell bears a
general resemblance to species oi Margarifcs Gray, 1847
(Quinn, 1979). The tvpe species of Bathijmophila and
B. bairdi (Dall, 1889) (Quinn, 1979, figs. 1-4) are closely
similar to Ethaliopsis callomphala (Schepman, 1908)
(figures 56, 57, 59) in gross shell fades. All three are
distincti\'e in having spiral sculpture on the earliest te-
leoconch whorls, subsequent teleoconch whorls smooth
apart from (usually) a subsutural row of nodules, and the
umbilicus filled bv a heavy callus with a granular surface
and a small nodule near the abapical extremity. Bathij-
mophila gravida new species appears to have exception-
ally variable shell morphology and to encompass within
its variation forms that are scarcely distinguishable from
B. ciispira or B. bairdi in gross moqihology. The radula
of B. gravida new species (figures 130, 131) is essentially
similar to that of E. callomphala (Schepman, 1908, pi.
9, fig. 9) but is distinctive in having a flagellum-like pro-
jection on the lateromarginal plate. Ethaliopsis is here
treated as a svnonym of Bathijmophila, because there
seems to be no way of justif)ing continued separation.
The Bathijmophila radula is strongly similar to those of
the t\pe species ni Microgaza (Marshall, 1979, fig. 9J-
L) And Archiminolia (Herbert, 1987, fig. 209) but differs
in that the lateromarginal plate is narrower and has a
terminal flagellum-like projection.

The animal of a paralectotyj^e of B. callomphala has
unpigmented eyes (blind). On the right side is a rounded
neck lobe, from which the eye stalk is not differentiated;
and three epipodial tentacles, the anterior one broad
with a rounded tip and projecting posteriorly from a
broad membrane; middle tentacle posterior to a convo-
luted membrane, smallest; posterior tentacle long and
slender. Left side with ven' small eye stalk, two small,
tapered neck lobes, and three large epipodial tentacles,
the anteiior one separated from the posterior two by a
broad epipodial lobe. The animals of the three other
Bathi/mophila species examined (see below) are essen-
tially similar, having a broad right neck lobe, two smaller
left neck lobes, three well-developed pairs of epipodial
tentacles, and a broad epipodial lobe between the two
anterior pairs of tentacles.

There is rather smooth transition in shell morphology
between Bathi/viophila species and the type species of
Archiminolia among representatives from the New Zea-
iaittl region. For example, Archiminolia alabida (Mar-
.' . '^ !9"9', A. tenuiseptiim new species, and A. diade-
■. snecies have the umbilicus covered by an ex-
. ■ ■; ■■.<■ the inner lip as in Bathijmophila euspira, yet

the covering is a thin, smooth, translucent septum rather
than thick pad with a granular surface, and the shell is
otherwise similar to the type species oi Archiminolia. On
the other hand, some apparent forms of Bathipnophila
gravida have an open umbificus at maturity and are thus
extremely similar to Archiminolia species. Whereas
there seems little doubt that Ethaliopsis is a synonym of
Bathijmophila and that Bathijmophila and Archiminolia
are closelv relatetl to Microgaza, 1 hesitate to suggest
that Archiminolia is a synonym of Bathijmophila or Mi-
crogaza or that Bathijmophila and Archiminolia might
be more appropriate as subgenera of Microgaza. Be-
cause of these uncertainties, I accept Bathipnophila, Mi-
crogaza, and Archiminolia as discrete genera, defined as
follows: Bathijmophila — high to low spire, narrow um-
bilicus infilled by heavy callus with granular structure
and with one or \^vo small bosses near base, some spe-
cies with fine interstitial axial riblets on earh' teleoconch,
lateromarginal plate (innermost marginal) with terminal
flagellum; Microgaza — low spire, umbificus open and
very wide, externallv bounded by rounded axial folds
separated by incised grooves, no interstitial axial riblets
on early teleoconch, lateromarginal plate broad and sim-
ple; Archiminolia — high to moderate spire, narrow um-
bificus open or covered by a thin septum, no interstitial
axial riblets on early teleoconch though fold-like axial
costae present in some species, lateromarginal plate
broad and lacking terminal flagellum.

Microgaza, Bathijmophila, and Archiminolia species
are strikingly simflar to Solariclla niida (Dafl, 1896)
(Hickman and McLean, 1990, fig. 72A, 75B), the
smoother forms of S. obscitra (Couthouy, 1838) (Waren,
1993, fig. 5, 7), and some species of Ilanga Herbert,
1987 (Herbert, 1987) in shefl morphology, the radulae
of which, however, lack lateromarginal plates (see re-
marks on Solariclla).

Besides the species discussed herein, the only other
known members of Bathipnophila are Solariclla micans
Dautzenberg and Fischer, 1896 (Waren, 1990, p. 180),
from the northeastern Atlantic, and an unde.scribed
western Adantic species (J.F. Quinn, pers. comm.).

Bathijmophila gravida new species
(Figures 50-53, 75, 130, 131)

Description: Shell up to 16 mm wide, wider than
high, rather thick and heavy, pofished, white, umbilicus
filled by thick callus. Proto'conch 430 [xm wide, sculp-
tured with a few fine spiral threads, tip of apical fold
broadly rounded. Teleoconch up to 5.3 whorls; first
whorl convex, subsequent whorls weakly convex on spire
and base, periphery evenly rounded, second and third
whorls with narrow horizontal sutural ramp with angu-
late rim. Spire on first 1.0-2.5 \yhorls sculptured with
fine, similar spiral cords, and much finer, crowded in-
terstitial axiiil riblets, 7 cords on first whorl, typically
multiplying to 8-10 by intercalation, all spirals other
than that at ramp rim weakening and vanishing together
with riblets. On third whorl ramp rim spiral enlarging to

B. A. Marshall. 1999

Page 21

occupy position of former siitural ramp, with rounded
nodules, cord splitting to form 2 similar cords with
rounded nodules, nodules obsolete on last adult whorl.
Base beside umbilical callus typically with 1 or 2 low or
obsolete spiral cords with rounded nodules, nodules dis-
crete or continuous across interspace. Umbilicus rather
narrow with tight!)' rounded rim, outer part completely
covered by thick callus with finely granulate surface,
small rounded boss near abapical extremity. Aperture
subcircular, outer lip thin at rim, moderately thickened
within, simple, inner lip continuous with umbilical callus.
Animal (idcohol, form C, NZOI U602) lacking eye pig-
ment. Right side with flap-like neck lobe fused to eye-
stalk, and 3 long, slender epipodial tentacles, anterior
tentacle separated from posterior 2 by broad epipodial
lobe. Left side with 2 small, tapered neck lobes and 4
epipodial tentacles, anterior tentacle longest and narrow-
ly tapered, second tentacle very small, posterior tenta-
cles of intermediate size, second tentacle extending from
edge of broad epipodial lobe between first and third ten-
tacles. Radula (holotype, figures 130, 131) with 9 p;iirs
of marginal teeth per transverse row, central and lateral
teeth with strongly serrate cutting areas, innermost mar-
ginal with flagellum-like terminal cusp on outer edge.

Type material: Holotvpe H68() and 5 paratypes
NZOI P1127, from type locality, 6 Febmary 1988, IW
Rapuhia (U584), collected afive.' Paratypes (10): N Three
Kings Rise, 3ri9.9'S, 173°05.1'E, 1570-1563 m, 9 Feb-
marv 1988, IW Rapuhia (1, NMNZ M. 131565; 5, NZOI
U6dl/P1128): N Three Kings Rise, 31°30.7'S,
172°49.8'E, 1216-1385 m, 9 February 1988, IW Ra-
puhia (2, NMNZ M. 131566; 2, NZOI U602/P1123)

Type locality: N Three Kings Rise, 31°26.3'S,
172°35.fi'E, 1137-1150 m.

Other material examined (25 specimens): N Three

Kings Ri.se, 3()°21.5'S, 173°08.7'E, 1474-1365 m. 7 Feb-
niaiT 1988, RA^ Rapuhia (1 form B, NZOI U595); N
Three Kings Rise, 30°32.3'S, 172°55.3'E, 1097-1082 m,
7 February 1988, RA^ Rapuhia (2 form A, NZOI U593);
N Three Kings Rise, 30°40.9'S, 173°51.5'E, 1335-1520
m, 8 February 1988, IW Rapuhia (1 form A, NZOI
U598); N Three Kings Rise, 3ri9.9'S, 173°05.1'E,
1570-1563 m, 9 February 1988, RA' Rapuhia (4 form
B, 8 form C, 4 form D, NZOI U601); N Three Kings
Rise, 31°30.7'S, 172°49.8'E, collected alive, 1216-1385
m, 9 Februarv 1988, RA' Rapuhia (1 form C, NZOI
U602); S Norfolk Ridge, 34°44.1'S, 169°25.0'E, 1123-
1064 m, 3 February 1988, IW^ Rapuhia (4 form A,
NZOI U571).

Distribution (Figure 75): Southern Norfolk Ridge
and northern Three Kings Rise, 1082-1570 m, collected
alive at 1137-1150 m (form A) and 1216-1385 m (form
C) from rocky substrata with foraminiferal sand.

Remarks: Batlu/iuophila gravida differs from the lec-
totype (the specimen figured by Schepman here select-
ed, ZMA 3.08.049, Siboga station 100, Sulu Sea, 450 m)

and paralectotypes of B. callomphala (Schepman, 1908)
principally in attiuning much larger size (maximum
width 16 mm, cf 6.50 mm), and in having a smaller
protoconch (width 350 |xm, cf 430 |xm). The shell of a
paralectotype of B. caUomphala is illustrated here (fig-
ures 56, 57, 59), and some features of the external anat-
omy are described above.

Four shell moqjhotvpes are present in the sample
from NZOI station U601: form A (7 specimens, figures
50-53, includes type material) — width up to 16 mm
(est.), lengtli/width ratio 0.72-0.88 (4 specimens), weakly
convex spire whorls, two nodular subsutural spirals, one
or two rows of nodules on inner base, smooth periphery,
umbilicus infilled; form B (4 specimens, figures 54, 55)
width up to 12 mm (est.), length/width ratio 0.88 (1
specimen), 1 nodular subsutural spiral, inner base
smooth or with weak axially elongate notlules, periphery
smooth, umbilicus infilled; form C (8 specimens) — width
up to 12 mm, length/width ratio 0.82-0.92 (2 speci-
mens), no subsutural or basal spirals or nodules, smooth
periphen', umbilicus infilled in six specimens, open in
two; form D (4 specimens) — width 14 mm, lengtlVwidth
ratio 0.86 (1 specimen), no subsutural or basal spirals or
nodules, smooth peripheiy, umbilicus open, early teleo-
conch spirals also more persistent than in other forms.
All other samples examined are form A shells with the
exception of NZOI station U602, which comprises four
form A and one form C with an infilled umbificus. The
single shell from station U595 resembles ff)rni B but has
no nodules on the subsutural spiral. The single shell
from station U598 has no spiral cord or nodules on the
base but otherwise resembles form A.

Given the occurrence of additional variants at NZOI
stations U602 and U598, and reports of similar variation
in B. cuspira (Dall, 1881) by Jeffreys (1883) and Quinn
(1979), I am not convinced that forms A-C represent
distinct species. The status of form D is even less clear.
Assuming that all specimens really are conspecific, ex-
treme polymorphism may be due to one or more of se\-
eral factors such as sexual dinioiphism, inflax of genetic
material over time from remote sources, translocation
from various depths, or differential Mayrian founder ef-
fect on waxing and waning spatially and temporarily iso-
lated populations. Most of the shells are old and friable
to varying degrees and may have accimiulated in self-
buffering calcareous secUments over hundreds or thou-
sands of years. Form A, and forms B and C are strikingly
similar respectively to B. hairdi (Dall, 1889), which is
particularly \'ariable in sculpture, and B. cuspira. both of
which have been collected in the Yucatan Channel at
1170 meters depth (Quinn, 1979), the implication being
that the latter two may fikewise be forms of a single
species.

Etymology: Laden, heavy (Latin).

Bathijiuopluhi valcutia new species
(Figures 58, 60, 61, 75)

Description: Shell up to 8.60 mm wide, wider than
high, rather thick and heavy, polished, white, umbilicus

Page 22

THE NAUTILUS, Vol. 113, No. 1

Figures 62-74. Sliells ot species of Bathijriuipliila and Airhiminolia. 62-65. B. cisphala new species, holotvpe, W'anganella Bank,
Norfolk Ridge, 449-442 m, NZOI H682 (4.20 X 5.15 mm). 66-70. A. tcnui.wptum new species, N Three Kings Rise"; 790-780 m.
66, 68, 69. HoloUpe, NZOI H6S3 (11.0 X 16.7 mm). 67, 70. Parat>pe, NZOI P1124 (11.2 X 17.0 mm). 71-73. .A, diadema new
species, holohpe. Wanganella Bank, Norfolk Ridge, 757-660 m, NZOI H684 (lO.O X 12.3 mm). 74. A. alahida (Marshall), holotype.
Silent I seamonnt, Kermadec Ridge, 695-689 m", NZOI H255 (8.30 X 10.5 mm). Scales 63, 65 = 1 mm. 64 = 250 |jLm.

B. A. Marshall, 1999

Page 23

» i;9- u i?B" » m"

Figure 75. Map of northern New Zealand region showing distribution of species of Bathymophila. Archiminolia. and Microgaza
(500, 1000 and 2000 m isobaths indicated): B. gravida new species (♦), B. valentia new species (o), A. alabida (Marshall) (■), B.
asphala new species (•), A. temiiseptum new species (*), A. diadenui new species (□), M. norfolkensis new species (O).

fiUed by thick callus. Protoconch 430 |jLm wide, sculpture
unknown (surface etched), tip of apical fold broadly
rounded. Teleoconch up to 4.8 whorls; first whorl
strongly convex with narrow, horizontal sutural ramp;
subsequent whorls becoming almost flat on spire; low
suprasutural angulation commencing after second whorl,
angulation vanishing on penultimate or last whorl in
atlults so that adult peripherv is rounded, base weakK-
convex. First whorl sculptured with 2 or 3 fine, similar
spiral threads, 1 on ramp, 1 on ramp rim, another beside
ramp rim in 1 specimen, crisp on first whorl, obscure on
second whorl; on third whorl resoKdng us 2 low, lirf)ad,
rounded, subsutural spiral cords, eventually with round-
ed nodules; periphery on last adult whorl with 3 or 4
obscure or well-defined, narrow, smooth, rounded spiral
cords, 1 surmounting suprasutural angulation, in some
specimens another on its adapical side, 2 on outer base;
base with or without row of rounded nodules at inner
third, broad, smooth, rounded swelling beside umbilical
rim. Umbilicus narrow though completely covered by

thick callus with coarsely granulate surface, strong
rounded, spirally elongate boss near abapical extremity.
Aperture subcircular, outer lip thin at rim, moderately
thickened within, simple, inner fip continuous with um-
bilical callus. Animal unknown.

Type material: Holotype NZOI H681 and 2 para-
types (1, NMNZ M. 131611; 1 NZOI P1129), from type
localitv, 9 February 1988, RA' Rapuhia (U601). Paratype
(1): N Three Kings Rise, 31°30.7'S, 172°49.8'E, 1216-
1385 m, 9 February 1988, RA' Rapithia (NZOI U6()2/
PI 130).

Type locality: N Three
17.3°()5.1'E, 1570-1563 m.

Distribution (Figure 75):
Rise, 1216-1570 m (dead).

Kings Rise, 31°19.9'S,

Northi

Thr

Kings

Remarks: Bathijtnophila valentia closely resembles B.
gravida in gross fades, differing in attaining smaller size
and in being smaller relative to the number of whorls

Page 24

THE NAUTILUS, Vol. 113, No. 1

(width up to 8.60 mm, 4.8 TW, cf. 16,0 mm, 5.3 TW)
and in having a strong, rounded spiral swelling beside
the umbilical rim. BathijinophiJa valcntia occurred to-
gether with B. gravida (all empty shells) at the two sta-
tions from which it is known, and there is no indication
of intergradation.

Etymology: Strength, vigor, value (Latin).

Baflu/inoplula asphala new species
(Figures 62-65, 75)

Description: Shell up to 8.2 (est.) mm wide, wider
than high, rather thick and heavy, pohshed, white, um-
bihcus filled bv thick callus. Protoconch .350 |jLm wide,
sculptured with 5 crisp, similar spiral threads, tip of api-
cal fold broadly rounded. Teleoconch up to 4.75 whorls,
first 1.5 whorls strongly convex, subsequent whorls
broadly convex above and below strongly rounded pe-
riphery', suture channelled before last adult whorl. Spire
on first 1.5 whorls sculptured with fine, similar spiral
threads, typicallv multiplying bv intercalation, weakening
and vanishing on next hidf whorl. Subsequent whorls es-
sentiallv smooth apart from subsutural row of low,
rounded nodules on third and first half of fourth whorl;
and moderate to weak, rounded axial pleats on inner
third of base, umbilicus liounded bv low, rounded swell-
ing. Umbilicus rather narrow with tightlv rounded rim,
partly or completely covered by thick callus with finely
granulate surface, 2 rounded, spirallv elongate bosses
near abapical extremity, another near adapical extremity.
Aperture subcircular, outer lip thin at rim, moderately
thickened within, simple, inner lip continuous with um-
bilical callus. Animal unknown.

Type material: Holot\pe NZOI H682 and 9 para-
types (1, NMNZ M. 131612; 8, NZOI P1131), from type
locality, 25 January 1977, IW Tangaroa (P13). Paratype
(1): Wanganella' Bank, Norfolk Ridge, 32°40.8'S,
167°26.8'E, 757-660 m, 25 January 1977, IW Tangaroa
(NZOI P8/P1132).

Type locality: Wanganella Bank, Norfolk Ridge,
.32°10.5'S, 167°21.2'E, 449-i42 m.

Other material examined: SSE of Phifip Island,
Norfolk Island, 29°33'S, 168°07'E, 326 m, 11 July 1962,
R.N.Z.F.A. Tui (4 juveniles, NMNZ M.224838)'; Wan-
ganella Bank, Norfolk Ridge, E slope, 32°.35.3'S,
167°41.8'E. 437-422 m, 29 January 1981, WW Tangaroa
(3 juveniles, NMNZ M.234119); N of Three Kings Is-
lands, 34°01'S, 172°07'E, 622 m, 18 February 1974, R/
V Acheron (fragment of adult, NMNZ M. 34608).

Distribution (Figure 75): Norfolk Island, Wanganel-
la Bank and off Three Kings Islands, 326-757 m (dead).

Remarks: Bathijnwphila asphala differs from B. gra-
vida in attaining smaller size (width up to ca. 8.2 mm,
cf 16 mm), in having a smaller protoconch (width 350
\i,m. cf 430 jjLm), and in having two bosses instead of
one near the base of the umbilical callus. Though very

incomplete, the shell from off the Three Kings Islands
resembles the tvpe material in protoconch size and
sculpture of the umbilical callus.

Etymology: Steadfast (Greek).

Genus Archiminolia Iredale, 1929

Archiminolia Iredale, 1929: 170. Type species (by original des-
ignation): Monilea oleacea Hedley and Petterd, 1906; Re-
cent, eastern Australia.

Remarks: As discussed above, Archiminolia species
resemble species of Bathijmophila and Microgaza in
gross shell and radular mor|:)hologv, differing from
Bathijmophila species in diat the umbilicus is open or
covered by a thin septum at maturity, in having a broad
lateromarginal radular teeth without a terminal flagel-
lum, and in having the rim of the inner fip smooth ratlier
than granulate. They differ from Microgaza species in
having a narrower umbilicus and a higher spire. The an-
imal of the type species has a large left neck lobe and
three large, tapered epipodial tentacles on the left side,
a broad epipodial lobe between the bases of the two
anterior tentacles, and a small flap in front of the ante-
rior tentacle. On the right side are two small, tapered
neck lobes, and four long, tapered epipodial tentacles.
The radula of the tvpe species was illustrated by Herbert
(1987, fig. 209).

Apart from A. oleacea (Hedley and Petterd, 1906) and
the species recorded below, other species apparentlv re-
ferable to Archiminolia include Solaiiella zacallcs Mel-
vill and Standen, 1903 (Persian Gulf); Solariella oliva-
ceostrigata Schepman, 1908 and Solariella zacalloides
Schepman, 1908 (Indonesia); and the southern Japanese
species Microgaza fulgcns. Dall, 1907, Ethaliopsis iri-
dcsccns Habe, 1961, Ethaliopsis katoi Kuroda and Habe
in Habe, 1961, Microgaza ziczac Kuroda and Habe in
Kuroda, Habe and Oyama, 1971, and Solariella (Micro-
gaza) opalina Shikama and Hayashi in Shikama, 1977.
Their animals, however, are unkiiovvm, and some may
prove to belong in llanga Herbert, 1987.

Archiminolia alahida (Marshall, 1979)
(Figure 74)

Solariella (.Microgaza) alahida Marshall, 1979: .5.32. fig. 4K-M,

9M-0.
?Solariella (Microgaza) sp, cf alahida. — Marshall, 1979: 53.3,

fig. 5A-C.

Type material: Holohpe NZOI H255, from type lo-
cahty, 5 September 1974, WV Tangaroa (J659), collected
alive. Paratvpes (2): Silent 1 seamount, 35°02.0'S,
179°15.8'E, 803-788 m, 5 September 1974, RA' Tan-
garoa (J66(): 1, NMNZ M. 131540; 1, NZOI).

Type locality: Silent 1 Seamount, southern Kermadec
Ridge, 35°00.6'S, 179°15.1'E, 695-689 m.

Other material examined (3 specimens): N Three
Kings Rise, 3r30.7'S, 172°49.8'E, 1216-1.385 m, 9 Feb-

B. A. Marshall, 1999

Page 25

Figures 76-89. Shells ol species of Arrhiminolid. 76-80. A. diadcina new species, parat\pes, W'aiiguiiellu Bunk, Norfolk Ridge,
757-660 m, NZOl P112.5 (76-78, 9.20 X 1,3.0 mm). 81. A. dawsoni (Marshall), holotype, NE of Raoul Island, Kermadec Islands!
14.5 m, NZOI H257 (7.40 X 9.20 mm). 82, 85. A. oleacea (Hedley and Petterd), N of Eraser Island, Queensland, 229-192m, AMS
C.l;3.3269 (13.0 X 16.5 mm). 83, 84. A. episcopniis new species 83. Holotype, SW of Great Island, Three Kings Islands, 427 m,
M. 131.541 (7.50 X S.20 mm). 84. Paratyjje, SW of Great Island, 440 m. M. 131.546. 86. A. hurleiji (Marshall)', holotype, NE of
Raoul Island, 145 ni, NZOI H2.56 (9.00 X 11.0 mm). 87-89. A. meridimm (Dell), Taiaroa Canyon, off E Otago 549 m M 9055
(97-6.60 X 7.90 mm). Scales 80, 89 = 250 jxin; all other figures = 1 nnii

Page 26

THE NAUTILUS, Vol. 113, No. 1

ruaiy 1988, RA^ Rapuhia (1, NZOI U602); Three Kings
Trough, NW of Three Kings Islands, 34°00'S, 171°55'E,
805 m. 15 July 1962, R.N.Z.F.A. Tui (1, NMNZ
M.59362); NE of Raoul Island, Kermadec Islands,
28°34.9'S, 177°50.0'W, 508-510 m, 26 July 1974, RA'
Tanoaroa (1, NZOI K828/2).

Distribution (Figure 75): North Three Kings Rise
and Kermadec Ridge, 508-1385 m, collected aHve at
689-695 m.

Remarks: Archiminolia alabida differs from all spe-
cies oi Archimirwiia, other than two of the new species
described below, in that the umbilicus is covered by a
thin, translucent septum at maturity. It is transferred
from Soloiiclla (Microgaza) to Archiminolia rather than
to BatJujifiophila because the septum is thin and smooth
instead of thick and more coarsely granulate and because
the lateromarginal plates lack a terminal flagellum.

A single broken shell from off Raoul Island, Kermadec
Islands (Marshall, 1979, fig. 5A-C), certainly adult, is
more strongly sculptured than the t\pe specimens and
lacks an umbilical septum but is otherwise similar They
may well be conspecific, because a similar degree of var-
iation is exliibited by some other Archiminolia species.

Archiminolia teniiiscptum new species
(Figures 66-70, 75, 132)

Description: Shell up to 17 mm wide, wider than
high, rather thin, highlv polished and shpperv, umbilicus
covered by thin septum. Protoconch and first 2.5 teleo-
conch whorls translucent white; subsequent spire whorls
pink with irregular darker a.xial bands and a narrow,
white suprasutural band; base white, nacreous through
translucent outer shell layer, inner lip and umbilical sep-
tum white. Protoconch 430-450 |xm wide, sculptured
with 2 fine, crisp spiral threads and minute granules, tip
of apical fold broadly rounded. Teleoconch up to 5 con-
vex whorls, suture channelled on first 3.5 whorls, chan-
nel rim angulate, periphery strongly convex, base weakly
convex. Spire sculptured with spiral threads and round-
ed collabral a.xial folds. Spiral threads on first 3 spire
whorls 6, weak or very weak, either vanishing early on
fourth whorl or enlarging on third whorl, multipKing by
intercalation and persisting throughout. Axial folds stron-
gest abapicall}^ and coronating suture, gradually resoKing
on second or third whorl, weakening on fourth whorl
and vanishing early or late on fifth whorl, entirely tra-
versing spire on second and tliird whorl or third and
most of fourth whorl, thereafter increasingly evanescent
abapicaliy until restricted to subsutural zone. Base at in-
ner third with 6-<S weak spiral threads and low, rounded
axial folds, elsewhere smooth apart from obscure spiral
lines. Umbilicus rather narrow widi tightly rounded rim,
almost or entirely covered by thin translucent septum
that spreads from inner lip; inner Up rim with finely
granulate surface. Aperture subcircular outer lip thin at
rim, weakly thickened within, simple. Animal with pig-
mented eyes. Right side with broad neck lobe that is

almost completely fused with eye stalk and extends to a
projecting flap at the base of anterior epipodial tentacle,
and 3 long, narrowly tapered epipocbal tentacles, ante-
rior 2 separated by a broad epipodial lobe. Left .side with
2 small neck lobes and 3 posteriorly enlarging epipodial
tentacles, bases of posterior neck lobe and anterior 2
epipodial tentacles connected by a broad membrane.
Jaw plates thin; jaw elements small, strongest at anterior
edge, weakening posteriorly, extending almost to poste-
rior margin though extremely weak in posterior third.
Radula typical of the genus (figure 132).

Type material: Holotype NZOI H683 and 1 paratvpe
NZOI PI 124, from type locality, 5 Februaiy 1988, IW
Rapuhia (U.582), collected afive.

Type location: N Three Kings Rise, 31°51.7'S,
172°26.0'E, collected alive, 790-780 m.

Distribution (Figure 75): Northern Three Kings
Rise, living at 780-790 m.

Remarks: Archiminolia tenuiseptum resembles Ar-
chiminolia alabida in general facies, differing in attaining
larger size and in being larger relative to the number of
whorls (width up to 17 mm: 5 TW, cf 10.7 mm: 5 TW)
and in having a sfightly larger protoconch (width 430-
450 |JLm, cf 400-420 |xm). Archiminolia tenuiseptum dif-
fers further in having a lower spire, stronger ;L\ial sculp-
ture, and stronger sculpture on the inner base. The par-
atvpe has a strong growth scar early on the third teleo-
conch whorl, and it is unclear whether or not the
sculptural chfferences on subsequent whorls between it
and the holotyj^e are due to damage to the mantle.

Etymology: Thin-walled (Latin), alluding to the um-
bihcal septum.

Archiminolia diadcma new species
(Figures 71-73, 75)

Description: Shell up to 13.5 mm wide, wider than
high, rather thin, highly polished, narrow umbificus open
or covered by thin septum. Protoconch and first 3 teleo-
conch whorls translucent white; subsequent spire whorls
pale pink with or without irregular darker axial bands
and narrow, white suprasutural band; base white, nacre-
ous through translucent outer shell layer, inner hp and
imibilical septum white. Protoconch 370 (xm wide,
sculptured with 4 or 5 fine, crisp spiral threads, tip of
apical fold broadly rounded. Teleoconch up to 5.2
whorls, all but last adult whorl with narrow, more or less
horizontal sutural ramp with angulate rim, periphery and
early spire whorls strongly convex, last few whorls in
adults weakly convex above and below periphery. Spire
sculptured with narrow but prominent spiral cords and
weak, rounded collabral axial folds. Spiral cords multi-
plying by intercalation, 7 or 8 at end of third whoil and
9-13 at end of fourth whorl on spire, gradu;illy weak-
ening on mid fourth or early fifth whorl before becoming
obsolete; adapical spiral at ramp rim, with prominent.

B. A. Marshall, 1999

Page 27

Figure 90. Map of New Zealand region showing distribution

of species oi Archiminolia (200 and 1000 ni isobaths indicated);
A. wanganellica new species {*), A. cpiscopalis new species and
A. regalis new species (■), A. meridiana (Dell) (A), and A.
diplax new species (O).

roundly conical nodules after second whorl. Axial sculp-
ture weak before third whorl, becoming obsolete on last
adult whorl, strongest at ramp rim, weakening and van-
ishing abapically across spire whorls. Base with 4 or 5
rounded spiral cords at inner third, and strong, rounded
axial folds, elsewhere smooth apart from oliscure spiral
lines. Umbihcus rather narrow with tightly rounded rim,
open in juveniles, in adults open or entirely covered by
thin translucent septum that spreads from inner lip, in-
ner hp rim with finely gramilate surface. Aperture sub-
circular, outer Up thin at rim, weakly thickened within,
simple. Animal unknown (only available specimen de-
cayed and broken into pieces during extraction).

Type material: Holotype NZOI H6cS4 and 7 para-
types (2, NMNZ M.272597; 5, NZOI PI 125), from t\pe
locality, 25 January 1977, RA' Tunoaroa (NZOI PS), col-
lected ahve.

Type locality: Wanganella Bank, Norfolk Ridge,
32°40.8'S, 167°26.8'E, 757-660 m.

Distribution (Figure 75): Wanganella Bank, south-
em Norfolk Ridge, hving at 660-757 m.

Remarks: Aichiininolia diadciiia most closely resem-
bles A. (dabida and A. tcnuisepfitin in general facies. It
chffers from A. tcnuiscptum and resembles A. alabida in
size relative to the number of whorls and chffers from
both in having a shghtlv smaller protoconch (width 370
|jLm, cf 4.30-450 |xm and 400-420 ji.m), in that the sub-
sutural spiral is stronger ami chstinctl)' nodular, in that
the spiral and axial sculpture at the inner third of the
base is much stronger, and in having paler shell colora-
tion.

Etymology: Crown (Greek), an allusion to the strong

subsutural row of nodules.

Archiminolia meridiana (Dell, 1953)
(Figures 87-90, 133)

Zeminolia meridiana Dell. 1953: 42, fig. 1, 2; Dell, 1956: 48;

Dell, 1962: 75.
Spectamen meridianum. — Powell, 1976: 84; Powell, 1979: 64.

Type material: Holotvpe NMNZ M.5666, from type
locality, 4 November 1950, R.R.S. Discovcnj II.

Type locality: Chatham Rise, 43°48'S, 178°58'W, 361 m.

Other material examined (Several hundred speci-
mens): NE of Mayor Island, Bav of PIent\, 37°I0'S,
176°23.5'E. collected dive, 732 m, 28 Februaiy 1957,
M.V. Aleri (1, M. 11257): off Gable End Foreland,
38°3S.7'S, 178=41. 1'E, collected alive, 755-725 m, 21
JanuaiT 1981, RA' Tangaron (9, M. 130095); off Cape
Kidnappers, 39°52.8'S, 177°36.5'E, collected alive, 785-
882 m, 21 Januan' 1981, RA^ Tangaroa (7, M. 130097);
off Cape Kidnappers, 39°5S.55'E, 178°14.18'E, collect-
ed alive, 900 m, Januan' 1994, F.V. San Maniikaii. coll.
M. Friar (4, M.118128); off Cape Kidnappers, 39°59.7'S,
178°08.0'E, collected alive, 665 m, 7 May 1973, RA^
Tangaroa (1 NZOI stn. G941); SE of Cape Campbell.
41°55.8'S, 174°40.7'E, 434-446m, 14 January 1979, RA'
Tangaroa. (5, M. 60378); steep outer slope of Conway
Ridge, S of Kaikoura, 42°37.2'S, 173°42.0'E, collected
ahve, 503 m, 13 January 1979, RA' Tangaroa (2,
M.59735); NE of Memoo Bank, W Chatham Rise,
42°38.1'S, 176°16.3'E. 980-1000 m, 11 Januan- 1979, R/
V Tangaroa (1, M.595S1); NE of Mernoo Bank,
42°38.2'S, 176°10.5'E, collected ahve, 999-984 m, 11
January 1979, RA' Tangaroa (3, M.59721); off Kaikoura,
■42°45.2'S, 173°44.8'E, 805 m, 4 November 1979, IW
Tangaroa (2 NZOI stn. S213); off Chatham Islands,
42°46.59'S. 177°15.91'W, 987-996 m, 23 Julv 1994,
FR.V. Tangaroa (1, M. 130101); off VVaiau River mouth,
S of Kaikoura, 42°55'S, 173°43'E, collected alive, 549-
586 m, 18 March 1976, RA^ Acheron (many, M.50870);
Mernoo Bank, 42°59.4'S, 175°30.5'E, 112 m, 23 Januaiy
1954. M.V. Aleri (1. M.9128); up steep wall of Pegasus
Canyon, NE of Banks Peninsula, 43°14'S, 173°.39'E, col-
lected alive, 1006-512 m, 27 September 1976, WV Ach-
eron (9, M.52780); Chatham Rise, 43°14'S, 176°11'E.

Page 28

THE NAUTILUS, Vol. 113, No. 1

collected alive, 366 m, 23 January 1954, M.V. Alert (10,
M. 130102); head of Pegasus Canvon, NE of Banks Pen-
insula. 43°24'S, 173°26'E, 366-.311 m, 18 March 1976,
BJV Acheron (1, M.51032); Chatham Ri.se, 43°32'S,
178°38'E, collected ahve, ca. 549 m, 24 January 1954,
M.V. Alert (5, M. 130098); NE of Kaingaroa, Chatham
Islands, 4.3°35'S, 176°03.5'W, collected"^ alive, 220-229
m, 7 Fehmai-v 1954, M.V Alert (1. M. 130105); Chatham
Rise, 43°38'S', 177°19'E, 531 m, 11 February 1954, M.V
Alci-f (1, M. 130100); Chatham Rise, 44°04'S, 178°04'W,
collected alive, 476 m, 10 February 1954, M.V. Alert (5,
M. 130099); SE of Banks Peninsula, 44°28.0'S,
174°54.8'E, collected ;ilive, 692 m, 5 October 1979, R/

V Tangaroa (2 NZOI stn. S143); SE of Pitt Island, Chat-
ham Islands, 44°32'S, 176°05'W, 302 m, 3 February
19.54, M.V Alert (17, M. 130104); SE of Banks Peninsula,
44°33.9'S, 174°51.2'E, collected ahve, 750 m, 4 October
1979, RA' Tangaroa (5 NZOI stn. S140); SE of Banks
Peninsula, 44°35.4'S, 174°49.6'E, collected aUve, 785 m,
4 October 1979, RA^ Tangaroa (13 NZOI ,stn. S138); SE
of Pitt Island, Chatham' Islands, 44°35.5'S, 176°04'W,
collected alive, 604 m, 3 February 1954, M.V Alert (27,
M. 130103); SE of Banks Peninsula, 44°50.17'S,
174°31.33'E, collected ahve, 760 m, 5 October 1979, R/

V Tangaroa (8 NZOI stn. S147); head of Waitaki Can-
von, off Oamaru, 45°10'S, 171°30'E, 29.3-256 m, 19
March 1976, WW Acheron (3, M. 51295); Karitane Can-
yon, off East Otago, 45°37.5'S, 17r06.0'E, collected
alive. 476-640m, 28^ March 19.54, M.V Alert (6, M.9136);
head of Karitane Canyon, 45°38.5'S, 171°05.0'E, 585-
5.30 m, 19 March 1976, VJ\ Acheron (1, M.51143); off
East Otago, 45°45'S, 171°02'E, 520-600 m, 1980, RA^
Munida (2,, M. 100609); Taiaroa Canvon, NE of Otago
Peninsula, 45°45.4'S. 17r05'E, collected ahve, 549 m,
16 August 1955, M.V Aleii (many, M.9055); Taiaroa
Canyon, 45°45.6'S, 171°05'E, 549 m, 23 January 1957,
M.V. Alert (19, M. 11256); Papanui Canyon, off Taiaroa
Head, 45°46'S, 171°03'E. 660 m, 1 September 1976, RA'
Munida (19, M.58479); off Otago Peninsula, 45°.50.S5'S,
171°01.71'E, .555-604 m, 4 June 1992, RA' Munida
(8, M. 131531); off Otago Peninsula, 45°51.05'S,
171°00.90'E, collected ahve, .500-589 m, 4 June 1992,
IW Munida (2, M. 130745); Canyon E, off East Otago,
collected alive, 421 m, 27 October 1967, RA' Munida (6,
M.26292); off Otago Peninsula, 4.5°51.71'S, 171°01.7S'E,
collected ahve. 619-5.50 m, 4 June 1992, IW Munida
(19, M. 130096); between Pukaki Rise and Auckland Is-
lands, 49°.5()'S, 170°14'E, collected alive, 593 m. 4 Sep-
tember 197S RA' Tangaroa (1 NZOI stn. S16); SE of
Campbell Island, 5.3°32.8'S, 172°16.6'E, collected aUve,
580 m, 2 Januan- 1971, RA' Tangaroa (1 NZOI stn.
G927).

Distribulion (Figure 90): North and South Island
east coast, from Mayor Island to Otago Peninsula, Chatham
Rise and Chatham IslantLs, and off Auckland and Campbell
islands, 220-1000 m; collected ahve at 220-999 m.

Remarks: The animal (alcohol) has on its right side a
well-developed neck lobe extending from the eyestalk to

the base of the first epipodial tentacle, more strongK-
projecting anteriorly and posteriorly; three long, narrow-
ly tapered epipothal tentacles, and a broad, rounded epi-
podial lobe between the two anterior tentacles. On the
left side are two neck lobes, the anterior slender, the
posterior subtriangular, and three long, narrowly tapered
epipodial tentacles, between which the epipochal fringe
is particularly well developed. The jaw plates are well-
developed. Jaw plate elements are strongest at the an-
terior edge and progressively weaken posteriorly. The
radula (figure 133) is typical of the genus.

Following an initial spirally lirate stage, the whorls of
Archiminolia meridiana may be entirely smooth or have
a subsutural zone of rounded axial phcae that eventually
weaken and vanish, with or without roimded plicae at
the umbilical rim or the addition of a narrow horizontal
subsutural ramp with an angulate rim; the spire a.xials
and subsutural ramp become obsolete late on the pen-
ultimate whorl or early on the last whorl in adults. The
shell may be uniform nacreous white, or the spire on
the last one or two whorls may be pale pinkish buff, with
or without pale axial bands, the base white below a sharp
peripheral boundary. All combinations occur together
and completely intergrade in several large samples.
Specimens from off North Cape (NMNZ M. 131.559,
131.562, juveniles and tragmentarv' adults) chlfer from
southern shells in being more deeply pigmented, in con-
sistently lacking subsutural axial plicae, and in having
smaller protoconchs (width 470-.5OO |jLm, cf 570-630
(xm). It is unclear whether these northern specimens
represent a disjunct local population of A. meridiana or
a distinct species, and better, geographically intermedi-
ate material (if such e.xists) is required.

Arcliiininolia meridiana is not to be confused with a
superficially similar though larger species (width up to
22 mm, cf 14 mm) with a rounded instead oi angulate
umbihcal rim, which occurs on the Bounty Platform at
705-722 meters depth and is an undescribed species of
Margarites Gray, 1847 (Margaritinae) (NZOI stations
1684, 1685).

Archiminolia diplax new species
(Figures 90, 92-95)

Description: Shell up to 11.0 mm wide, wider than
high, of moderate thickness, stout, polished, narrowly
umbilicate, white. Protoconch 600 |xm wide, minutely
granulate and with 2 fine spiral threads, tip oi apical fold
broadly rounded. Teleoconch up to 5 (est.) whorls; first
whorl strongly convex; subsequent whorls with narrow,
sloping sutural ramp with angulate rim and pronounced
suprasutural angulation; spire whorls strongly flattened
between angulations; !>oth angulations with rounded, ax-
ially elongate nodules, angulations and nodules obsolete
on last adult whorl; periphery rounded, base weakly con-
vex. First few whorls sculptured with crisp, similar spiral
lirae, 7 or 8 on spire, weakening and vanishing at end
of second whorl. Inner third of base with rounded axial
pleats, 2 low, rounded spiral cords at umbihcal rim. First

B. A. Marshall, 1999

Page 29

2 whorls with niicrosculpture of minute, crisp granules;
fine coUabral and spiral growth liTies throughout. Um-
bilicus rather narrow, rim tightlv rounded, wall weaklv
convex, with fine collabral growth lines. Apertme sub-
circular, outer lip thin at rim, moderately thickened
within, simple, inner lip rather strongly thickened
against umbilical rim. Radula unknown (only live-taken
specimen is juvenile).

Type material: Holotype NMNZ M. 60166, from type
locaht^', 23 Januarv' 1979, RA' Tangflroa. Paratvpes (5
juveniles NMNZ).'SE of Aldermen Islands, 37°04.3'S,
'176°26.6'E, collected alive, 807-872 m, 23 Januaiy 1981,
WW Tanoaroa (M. 75065).

Type locality: NE of Mayor Island, 37°09.1'S,
176°24.4'E, 753-826 m.

Other material examined: 1 ju\enile NMNZ, E of
Great Barrier Island, 36°12.0'S, 176°19.4'E, 486-655 m,
25 Januaiy 1981, IW Tanoaroa (M. 131558).

Distribution (Figure 90): Off Great Barrier, Alder-
men, and Mayor islands, 486-872 m, collected alive at
807-872 m from mud.

Remark.s: This species is extremely distinctive in ha\'ing
a suprasutural angulation on the spire whorls, rounded,
axially elongate nodules on both the submedian and sub-
sutural angulation, and rounded axial pleats on the inner
base. The sculpture of the third teleoconch whorl
(where sculpture is transitional from juvenile to adult) is
unknown, the t)pe material comprising bxo jusenile par-
atypes with 1.75 and 1.8 teleoconch whorls, and the
adult holotype, the first three whorls of which are de-
corticated.

Etymology: Double (Greek), an allusion to the t\vo an-
gulations on the spire.

Archiininolia cpiscopalis new species
(Figures 83, 84, 90)

Description: Shell up to 10.6 mm wide, wider than
high, rather thin, polished, narrowly umbilicate. Proto-
conch and first 2.5 teleoconch whorls white, subsequent
whorls pale buff with pale, irregular reddish brown axial
bands that extend from suture to extreme peripheiy or
slightly above it, sharply tmncated abapically, base white.
Protoconch 500 |xm wide, tip of apical fold broadly
rounded; sculptured with fine, crisp, crowded granules
and 4 fine spiral threads. Teleoconch up to 4.6 whorls;
first whorl con\ex, next ca. 2.5 whorls with narrow su-
tural ramp with angulate rim overhanging suture, ramp
obsolete thereafter, below subsutural rim spire whorls
and periphery rather evenly rounded, base more weaklv
convex. Spire on first 2-3 whorls sculptured with fine,
similar spiral cords that multiply by intercalation, 8 on
first whorl, typically multiplying to 12-15 before becom-
ing obsolete, rim of sutural ramp with or without low,
rounded axially elongate nodules after second whorl.
Fine collabral and spiral "rowth lines throughout. Um-

bilicus rather narrow, rim tightly roundeil and projecting
inwards to overhang convex wall. Aperture subcircular,
outer lip thin at rim, weakly thickened within, simple,
inner lip strongly thickened and finely granulate against
umbilical rim. Animal unknown.

Type material: Holot\pe NMNZ M. 131541 and 2
paratopes M. 131542, from type locality 21 June 1978,
IW Tanoaroa. ParatyjDes (8): Middlesex Bank, NW of
Three Kings Islands, 33°59.9'S, 17r45.3'E, 186-196 m,
31 Januaiy 1981, IW Tangama (3, M. 131544); 37 km
SW of Great Island, 34°20.4'S, 171°48.2'E, 440 m, 21
June 1978, RA' Tanzania (4, NMNZ M. 131546; 1 AMS).

Type locality: 39 km SW of Great Island, Three Kings
Islands, 34°17.6'S, 171°45.3'E, 427 m.

Other material examined (38 broken or immature
specimens NMNZ): 37 km NE of Great Island, Three
Kings Islands, 33°58.0'S, 172°30.6'E, 550 m, 25 June
1978, RA' Tanoflwa (2, M. 131548); Middlesex Bank, NW
of Three Kings Islands, 33°59.9'S, 171°45.3'E, 186-196
m, 31 January 1981, RA' Tangaroa (7, M. 131545); Three
Kings Trough, NW of Three Kings Islands, 34°00'S,
171°55'E, 805 m, 15 July 1962, ' R.N.Z.FA. Tui (4,
M. 59363); 13 km N of Great Island, 34°01.8'S,
172°12.()'E, 508 m, 23 June 1978, RA^ Tangoroa (2,
M. 131550); Middlesex Bank, 34°02.0'S, 17r44.0'E, 246-
291 m, 31 Januaiy 1981, RA' Tangaroa (4, M. 131551);
Middlesex Bank, .34°02.1'S, 17r45.8'E, 221-206 m, 31
Januaiy 1981, RA' Tangaroa (1, M. 131552); 22 km ENE
of Great Island, 34°05.0'S, 172°24.6'E, 200 m, 24 June
1978, IW Tangaroa (5, M. 131553); 39 km SW of Great
Island, 34°17.6'S, 171°45.3'E, 427 m, 21 June 1978, RA'
Tangaroa (10, M. 131543); 37 km SW of Great Island,
34°20.4'S, 17r48.2'E, 440 m, 21 June 1978, IW Tan-
garoa (3, M. 131547).

Distribution (Figure 90): Off Three Kings Islands,
186-805 m (dead), in comminuted Bryozoa and shell.

Remarks: Compared with Archiininolia mcridiana,
the color bands in A. cpiscopalis are stronger, the sutural
ramp is outwardK adapicallv sloping rather than hori-
zontal, and the axial sculpture in the subsutural zone
consists of nodules rather than short costae. Archinii-
nolia cpiscopalis differs further in having a smaller pro-
toconch (width 500 [xm, cf 570-630 |xm).

Etymology: Overseer (Latin).

Archiininolia regalis new species
(Figures 90, 96-99)

Description: Shell up to 10.3 mm wide, wider than
high, sturdy, narrowly uml)ilicate, highly polished. Pro-
toconch white. Ground color of first 2.5-3.0 teleoconch
whorls pale yellow, last whorl white or p;ile greenish or
dull reddish white. Second and subsequent whorls with
irregular pale dull red or olive green wavy axial bands
that extend from sutme to about midway across base and
that are interrupted over middle third of spire w'horis

Page 30

THE NAUTILUS, Vol. 113, No. 1

Figure 91. Map of New Zealand region showing distribution
of species of Zetehi (200 and 1000 m isobaths indicated): Z.
textilis (Murdoch and Suter) (A), Z. tangaroa new species (▼),
Z. variabilis Dell (■), and Z. anneciens new species (*).

dried animal) characteristic of the genus, lateromarginal
tooth broad.

Type material: Holot)pe NMNZ M. 131.513, from
tvpe locahtv', 31 Januarv 1981, RA' Tangaroa. Parat\pes
(9): Middlesex Bank, .33°.59.9'S, 171°4.5.3'E, 186-196 m,
31 Januarv 1981, RA' Tangaroa (2, M. 131518); 13 km N
of Great Island, Three Kings Islands, .34°01.8'S,
172°12.0'E, 508 m, 23 June 1978, IW Tangaroa (1,
M. 131515); Middlesex Bank, .34°02.0'S, 171°44.0'E,
246-291 m, 31 January 1981, IW Tangaroa (3, NMNZ
M. 131517; 1 AMS); off Northeast Island, Three Kings
Islands, .34°08.5'S, 172°11'E, 102 m, 18 February 1974,
VJW Acheron (2, 131514); SE of Great Island, .34°14.8'S,
172°13.6'E, collected alive, 17.3-178 m, 2 Februar)'
1981, RA' Tangaroa (1, M. 131516).

Type locality: Middlesex Bank, NW of Three Kings
Islands, 34°02.1'S, 171°45.8'E, 221-206 m.

Other material examined: 103 mostly immature or
fragmentary specimens in 20 lots from off Three Kings
Islands, NMNZ.

Distribution (Figure 90): Off Three Kings Islands,
91-805 m, collected alive at 17.3-178 m on comminuted
bryozoan substratum with sponges, corals, hydroids, etc.

Remarks: Archiininolia rcgalis most closely resembles
the Kermadec Ridge species A. dawsoni in general facies
but chffers principally in having a larger protoconch
(width 500-.530 (im, cf. 330 |jLm) and in being larger
relative to the number of whorls (width 9.70 mm, 4.3
TW, cf. 9.10 mm, 5.2 TW). Arcliiminolia rcgalis is the
most brightly and variablv colored solarielhne in the
New Zealand region. A similar species occurs on Wan-
ganella Bank, Norfolk Ridge, and is described below.

Etymology: Regal (Latin).

and at middle of outer half of base to form more or less
discrete subsutural, peripheral, and basal zones; subsu-
tural zone and adapical e.xtremity of peripheral zone
most deeply pigmented; umbOical rim and inner lip por-
cellaneous white. Protoconch 500-530 |j,m wide, sculp-
tured with 5 fine, crisp spiral threads and sparse gran-
ules, tip of apical fold broadly rounded. Teleoconch up
to 4.5 whorls; narrow, more or less horizontal sutural
ramp widi tightly rounded rim; convex from ramp to
umbilical rim tliough distinctlv flattened abo\'e and be-
low periphery'. Spire on first 2 whorls sculptured with
fine spiral cords that multiply by intercalation and en-
large to resemble adjacent spirals, about 20 near end of
second whorl, weakening and becoming obsolete early
on third whorl. Low. rounded a.xial pleats on subsutural
ramp. Fine collabral growth fines throughout. Umbificus
perspective to protoconch, rim tightly rounded, wall al-
most vertical, wall of each whorl shallowlv concave be-
side rim, convex within. Aperture subcircular, outer fip
thin at rim, moderately thickened within, simple, inner
lip strongly thickened. Radula (optical preparation from

Archiminolia wanganeUka new species
(Figures 90, 100-103)

Description: Shell up to 8.60 mm wide, wider than
high, sturdv, narrowlv umbilicate, highly pofished and
sfipperv'. Protoconch white; first 2 teleoconch whorls
very pale red; subsequent whorls white with pale yellow-
ish brown, irregular, wavy axial bands that extend from
suture or middle of spire whorls to midpoint of base,
predominant green of nacreous layer showing through
translucent outer shell la\er; umbilical rim and inner Up
porcellaneous white. Protoconch 470^80 |xm wide,
sculptured with about 5 fine spiral threads and minute
granules, tip of apical fold liroadK rounded. Teleoconch
up to 4.6 whorls; second and third whorls with narrow,
more or less horizontal sutural ramp with tightly round-
ed rim; convex from ramp to umbilical rim, weakl\- flat-
tened above and below periphery. Spire on first 2 whorls
sculptured with fine spiral threads that multipK by in-
tercalation and enlarge to resemble adjacent spirals, 1.'3-
19 near end of second whorl, becoming obsolete early
on third whorl. Fine collabral growth lines throughout.

B. A. Marshall, 1999

Page 31

Figures 92-103. Shells of
75.3-.S26 111, M. 60166 (S.SO x
mm). 96-99. A. rvoalis

pecies ot Anhiininolia 92-95. A. diyilax new specie.s 92. Holohpe (adult), NE of Mavor Lsland,

11.0 mm). 93-95. Ju\eiiile paratype, SE of Aldennen Islands, Sb7-S72 m, M. 75065 (2.50 X 2.70

new .species 96, 97. Middlese.x Bank, 221-206 m, M.l;3151:3 (7.70 x 9.70 mm). 98, 99. Paraty]ie, off

Northeast Island, Three K.mm Islands, 102 m. M. 1.31514. 100-103. A

m. 100. Parat>pe,
others = 250 (jLm.

ncUica new species, Wanganella Bank summit, 133

M. 131.537 (7..50 X 8,30 nnn). 101-103. Ilolotspe, M. 131,5.36 (6.20 X 6.90 mm). Scale bars 98, 102 = .500 \x.m.

Page 32

THE NAUTILUS, Vol. 113, No. 1

Umbilicus perspectixe to protoconch, rim tightly round-
ed, wall almost \'ert^ical, wall of each whorl shallowly con-
cave beside rim, convex within. Apeiture subcircular,
outer hp diin at rim, moderately thickened within, sim-
ple, inner hp strongly ihickened.

T\pe material: Holot\-pe NMNZ M. 131.536 and 6
paratypes (3, NMNZ M.'l31.5.37; 1 AMS), from type lo-
cality, 30 January 1981, IW Tangaroa, collected ahve
(dried).

Type locality: Summit of Wanganella Bank, Norfolk
Ridge, .32°.39.2'S. 167°31.7'E, 133 m.

Other material examined: Wanganella Bank, sum-
mit, 32°32.2'S, 167°30.7'E, 113 m, 29 Januaiy 1981, R/
V Tangaroa (1 immature, NMNZ M. 2.57230); slope of
Wanganella Bank, 32°10.5'E, 167°21.2'E, 449-442 m,
25 Januar\- 1977, RA' Tangawa (7, NZOl P13); Wan-
ganella Bank, summit, .32°39.2'S, 167°31.7'E, 133 m, .30
January 1981, IW Tangaroa (7 broken or immature,
M. 131538); Wanganella Bank, summit, .32°40.0'S,
167°.33.6'E, 133 m", 30 Januaiv 1981, IW Tangaroa (8
immature, NMNZ M. 131602).'

Distribution (Figure 90): Wanganella Bank, Norfolk
Ridge, 113-449 m, collected alive at 133 m from sub-
stratum of white sand and rhodolith gravel with small
red algae.

Remarks: Archimiuolia wangancUica differs from the
similar species A. dawsoui and A, regalis in protoconch
size (width 470-480 |jLm, cf 330 fim and .500-530 [xm),
and size relative to the number of whorls (width 8.60
mm, 4.6 TW, cf. 9.10 mm, 5.2 TW and 10.4 mm, 4.3
TW); it differs further from A. regalis in that the earhest
teleoconch whorls are pale red instead of v ellow.

Etymology: Named after the type locahty.

Archiininolia dawsoni (Marshall, 1979)
(Figure 81)

Solariella (Microgaza) dawsoni Marshall 1979: .5.3.3, fig. .5D-F.

Type materia!: Holotype (H257) and paratype NZOI,
from t>pe locality, 25 July 1974, IW Tangaroa (KS25).

Type localitv-: Kermadec Ridge, NE of Raoul Island,
28°47.8'S. 17r4r,S'W. 145 m.

Di,stribution: Kermadec Ridge NE of Raoul Island,
145 m.

Remarks: The iiolotvpe and paratype remain the only
specimens of Arckiminolia dawsoni seen to date. Su-
perficially similar species from off the Three Kings Is-
lands (A. regalis new species) and Wanganella Bank (A.
wangancUica new .species) are described above. The pro-
toconch of A. dawsoni is exceptionally small for the ge-
nus (width .3-30 |j.m).

Archiminolia hurleyi (Marshall, 1979)
(Figure 86)

Solariella (Microgaza) hiirleiji Marshall, 1979; 5.34, fig. 5G-I.

Type material: HoloUpe NZOI H256, from tvpe lo-
cahty, 25 July 1974, RA^' Tangaroa (KS25).

Type locality: Kermadec Ridge, NE of Raoul Island,

28°47.8'S, 177°47.8'W, 145 m.

Distribution: Kermadec Ridge NE of Raoul Island,

145 m.

Remarks: This species is transferred from Solariella
(Microgaza) to Archiminolia because of its general re-
semblance to the type species. As with A. datvsoni, the
protoconch is unusualh' small among Archiminolia spe-
cies from the southwestern Pacific (width 330 iJ-m).

Genus Microgaza Dall, 1881

Microgaza Dall, 1881: 50. Tvpe species (by original designa-
tion): Microgaza rotella I3all, 1881; Recent, West Indies.

Remarks: As discussed above, species of Microgaza
resemble Bathi/mophila and Arcliiminolia species but
differ from Bathijmophila species in that the umbihcus
does not become covered by a thick, granulate septum
at maturity and have broad lateromarginal radular teeth
without a terminal flagellum; species o{ Microgaza cUffer
from both groups in having a wider umbilicus. The spe-
cies described below is closer to the West Indian tvpe
species in shell facies than to anv species here referred
to Bathijmophila or Archiminolia. The only other named
species known from outside the Caribbean is M. nava-
kaensis Ladd, 1982, described from die Pleistocene of
Santo, Vanuatu, which also occurs hving off southern
New Caledonia (AMS C.328490, MNHN). I have ex-
amined additional, unnamed species from off Western
Austraha (AMS C. 170029) and off Crocker Island in the
Arafura Sea (AMS C..328491).

Microgaza noifolkcnsis new species
(Figures 75, 104-106)

Description: Shell up to 16.3 mm wide, wider than
high, thin, widelv umbilicate, highK- polished. Proto-
conch and first 2.5 teleoconch whorls white; next 1.5
whorls pinkish buff, last half whorl white with narrow,
irregular wavy axial band on spire, base white. Proto-
conch 400 |jLm wide, tip of apical fold broacUv rounded.
Teleoconch up to 5 whorls, first 2 whorls stronglv con-
vex, subsequent whorls more weaklv convex on spire and
base, periphery' strongly convex. Spire on first 2 whorls
sculptured with 6 crisp spiral threads, interspaces wider
than each thread, rapicUv becoming obsolete thereafter;
numerous fine, crowded, rounded spiral threads com-
mencing on fourth whorl, crisp on spire and periphery,
more weakly defined on base; low, rounded axial costae
commencing about midway through second whorl, en-
Hrelv traversing spire whorl at first, progressively enlarg-

B. A. Marshall, 1999

Page 33

ing and becoming obsolete abapically to form siibsutural
zone of rounded axial pleats, becoming olisolete on first
half of third whorl or late on fourth whorl. Inner quarter
of base and umbilical rim with close, rounded axial pleats
separated bv grooves. Umbilical rim strongly angulate,
surmountetl bv rounded spiral cord, umbilical wall even-
ly and moderately convex, sculptured with crisp spiral
lirae and coUabral growth lines. Umbilicus perspective
to protoconch. Aperture ovate, lips thin at rim, weakly
thickened within, simple. Animal unknown.

Type material: Holotyi^e NZOI H685 and paratype
NMNZ M. 131567, from t\pe locaht\, 30 Januaiy 1977,
RA' Tangaroa (P45).

Type locality: Off Norfolk Island, 2S°42.0'S,
167°57.8'E, 527-508 m.

Distribution (Figure 75): Off Norfolk Island, 527-
508 m (dead).

Remarks: Microgaza noifolkcnsis is strikingly similar
to the tropical western Atlantic species Microgaza rotclla
(Dall, 1881) in general facies but differs in attaining
more than twice the size (width up to 16.3 mm, cf 6.80
mm), in having a larger protoconch (width 400 |xm, ct.
330-350 |j.m), in having a more broadly rounded pe-
ripheiy, and in having stronger spiral lirae on the last
adult whorl. Microgaza uavakacnsis Ladd, 1982, from
the Pleistocene of Santo, Vanuatu, differs in lieing small-
er relative to the numiier of whorls antl in ha\ing a color
pattern of crowded, wavy axial bands (Ladd, 1982, pi
34, fig. 1-5).

Genus Zctcia Finlay 1926

Zetela Finlay, 1926: .359. Type species (by original designation):
Minolin textitis Mindoch and Siiter, 1906; Recent, New
Zealand.

?Lnmi'IUtrochii.s Qninn. 1991: 81. T\pe species (by original des-
ignation): Margarita Inincllosa \'errill and Smith, 1S80;
Recent, eastern North America.

Remarks: The tyjDe species of Zctcia is distinctive in
having strong, reticulating spiral and axial sculpture and
crisp granules on the teleoconch and in not having the
innermost marginal modified to form a lateromarginal
plate (figures 134-138). The animal of Zctcia tcxtili.s
(presen'ed material, NMNZ M. 58797) has a broad,
rounded neck lobe on the right side connected to the
eyestalk, a tiny tentaculiform neck lobe below the left
eyestalk, and four epipodial tentacles on each side, the
anterior and posterior pairs long and slender, the middle
pairs minute. The periostracum is apparently calcified
(intritacalx) in all species, though as in Solariclla (short
of examination of cross sections of fractured shells), its
presence is only revealed bv the effects of solution etch-
ing in a few species.

The southern African species Solariclla iutcrnmsa
Thiele, 1925 (Herbert, 1987, fig. 115— "ml" and "Inl"
are transposed) and the closely related species Z. kopiia
new species differ from Z. tcxtilis in haying an extra

(posterior) neck lobe on the left side but are otherwise
similar Like Z, tcxtili.s, both species lack lateromarginal
plates in the radula, though unlike Z. tcxtili.s, the tooth
that gives rise to it has a reduced cutting area. According
to Quinn's (1991) composite description of the external
anatomy, species of Lumcllitrochiis (^uinn, 1991, have
Kvo left neck lobes and three epipodial tentacles on each
side, the middle one long or short. The conchtion in the
type species {Margarita lamello.sa Verrill and Smith,
1880) was not specified, though from studv of preserved
material kindly lent by J.F. Quinn, there are actually four
pairs of epipodial tcTitacles, the two middle pairs much
smaller as in Z. tcxtili.s. All of these Atlantic species re-
semble Z. tcxtili.s in having strong axial sculpture, at least
on the earlv teleoconch, a protoconch with a bulbous tip
antl fine spiral threads, and a teleoconch microsculpture
of minute granules. The type species and most other
species referred to Lamcllitrochii.s bv Quinn (1991) dif-
fer markedly from Z. tcxtili.s in that the later spire whorls
are more weakly convex and scidptured with three an-
gulating primary spiral cords (abapical spiral more or less
covered by succeeding whorls). Differences in shell mor-
phology between L. lamcllo.su.s and Z. tcxtili.s, however,
are bridged via L. fcnc.strafii.s Quinn, 1991, L. carinatit.s
Quinn, 1991, L. hicoronatiis Quinn, 1991, and Zetela
variabilis Dell, 1956. Since Lamcllitrochus and Zetela
species also have similar external anatomies and radulae,
it seems fikely that Lamcllitrochii.s is a synonym of Zc-
tcla. A New Zealand Zetela species resembhng Atlantic
Lamcllitrochus species is described below.

The shells of Zetela tcxtili.s and Lamcllitrochus lamel-
losus are similar to the affiui.s form (figure 40) and the
holotvpe (Waren, 1993, fig. 2A), respectively, of Solari-
clla amahilis (Jeffreys, 1865), in which, however, latenJ
four is relatively larger and the innermost marginal has
been transformed into a broad lateromarginal plate
(Waren, 1993, fig. 7B), and there are three p;urs of sim-
ilar epipodial tentacles and a rounded flap-like stnicture
between the two anterior pairs of epipodial tentacles. As
discussed above, S. amahilis apparently belongs in So-
lariclla sensu stricto.

Zctcia tcxtilis (Murdoch and Suter, 1906)
(Figures 91, 109, 110, 134)

MinoUa tcxtilis Mnrdocli and Snter, 1906: 298, pi. 26, fig. 4.5,

46.
Monilea {Minolia} tcxtilis.— Suter. 191.3: 143, pi. .33, fig. 14,

14a,
Zetela texiilis.—Fm\a.y. 1926: 359.
hniula (Zetela) textilis.—Wenz. 193S: 318, fig. 720.
Solariclla (Zetela) tcxtilis.— Keen, 1960: 262, fig. 167/1; Powell,

1974: 207; Powell, 1979: 64, pi. 17. fig. 20.

Type material: Holotvpe NMNZ M.1123, from type
localitv', S.S. Auania.

Type locality: Off Great Barrier Island, 36°08'S,
17.5°55'E, 201 m, 22 Januaiy 1904.

Other material examined (ca. 1000 specimens

Page 34

THE NAUTILUS, Vol. 113, No. 1

256-293 m, M.94763 (4.6.5 x 4.70

Figures 104-1 15. Shells of species of Micro^nza and ZcUla^ 104-106. Microaaza noifolkemis new species, holot>pe, ott Norfolk
Island, .527-508 m, NZOI H685 (16.3 x 7.00 mm). 107, 112, 113. Zetela vnriahilis Dell. 107, 113. Pegasus Canyon, NE of Banks
Peninsula. 1006-512 m, M.52779 (107, juvenile; 113, 4..35 X 4.90 mm). 112. Pegasus Canyon,
mm). 108. Zetein am,

Zetela textilis {Wwdovh „ ,. ^,-vr^T uccc / < -rrr ^ =; c;i=;\

556-311 m M 35478 111, 114. Zetela tangawa new species, holot>pe, SE of Banks Penmsula, /60 m. NZUl Hb86 (4./,'3 x .^.DD)
Zetela kopua new species, paratype, N BounU' Trough, 1373 m, NZOI PI 126. Scale bar 110 = 120 jxm, others = 250 i^m.

vie'ctens new species, holotvije, off Great Barrier Island, 486-655 m, M. 137596 (5.80 x 5.05 mm). 109, 110.
loch and Suter). 109. SE of The Aldermen, 178-248 m, M.65154 (4.35 x 4.30 mm). 110. Off Mayor Island,

115.

B. A. Marshall, 1999

Page 35

NMNZ): E of North Cape, 34°25.0'S, 173°13.1'E,
collected alive, 327-257 m, 27 January- 1981, IW Tan-
garoa (71, M. 131497); SE of Aldermen Islands,
37°()().,S'S, 176°12.3'E, collected ;ilive, 178-248 in, 23 Jan-
udiy 1979, RA^ Tangaroa (many, M.65154); SE of Alder-
men Islands, 37°01.0'S, 176°14.8'E, collected alive, 357-
312 m, 24 January 1981, IW Tangaroa (1, M, 131498);
NNW of Mayor Island, 37°()8.7'S, 176°14.2'E, 356-380
m, 23 Januan- 1979, IW Tangaroa (2, M. 6 1047); NW of
Mayor' Island, 37°09.6'S, 176°12.2'E, 293-.348 m, 22 Jan-
uary 1979, IW Tangaroa (1, M.61028); NW of Mayor
Island, 37°11.5'S, 176°10.0'E, collected alive, 198-273 m,
22 Januan- 1979, IW Tangaroa (35, M.6656()); off Mavor
Island, 37°12'S, 176°17'E, collected alive, 256-311 m', 9
FebniaiT 1974, WV Acheron (9, M. 35478); SE of Mayor
Island, 37°21.9'S, 176°20.9'E, collected alive, 203-248
m, 22 January- 1979, IW Tangaroa (3, M.61319); off
Mayor Island,' Bay of Plenty, 37°22.5'S, 176°22'E, 207-
220 m, 27 February 1957, M.V. Alert (10, M. 12868); N
of Motuhora Island, 37°35.1'S, 176°59.5'E, collected
alive, 248-213 m, 20 Januar)- 1979, RA' Tangaroa (6,
M.65022); N of Motuhora Island, 37°37.8'S, 176°59.9'E,
collected alive, 129-139 m, 20 January- 1979, RA^ Tan-
garoa (21, M,67785); ENE of Tokomaru Bay, 37°59.7'S,
178°40.0'E, collected alive, 144 m, 16 Januaiy 1979, R/
V Tangaroa (43, M. 59846); ENE of Tolaga Bay,
38°15.2'S, 178°38.6'E, collected alive, 139 m, 16 January
1979, IW Tangaroa (29, M.59797); SE of Cape Kidnap-
pers, 39°55.5'S, 177°30'E, 183 m, 20 May 1952, M.V.
Ki^tuhu (1, M.8748).

Distribution (Figure 91): North Island east coast,
from North Cape to Cape Kidnappers, 129-380 m; col-
lected alive at 139-327 m from mud.

Remarks: Zctcia tcxtili.s attains 5.60 mm in width, is
uniform translucent white, and has a highly distinctive
sculpture of prominent axial lamellae that are narrowly
and sharjily angulate in section and strong, rounded spi-
ral cords, with sharply nodular intersections on the spire
and at the umbilical rim.

Zetela variabilis Dell, 1956
(Figures 91, 107, 112, 113)

Zetela variabilis Dell, 19.56: 48, fig. 70, 74.
Solariella (Zetela) variabilis.— FoweW, 1976: 84; Powell, 1979:
64.

Description: Shell up to 5.75 mm wide, wider than
high or about as high as -wide at maturity, of moderate
thickness, umbilicate, white. Protoconch 530-570 |jLm
wide, tip of apical fold broadly rounded; sculptured
■with fine, crisp, irregular granules, and 3 fine, crisp spi-
ral threads. Teleoconch up to 3.60 whorls, first whorl
rather evenly convex; subsequent whorls convex below
angulation at rim of gently sloping subsutural ramp,
base distinctly flattened, umbilical rim angulate. Sculp-
tured with rounded spiral cords and strong, rounded
axial costae. Spiral cords numbering 5 or 6 at end of
first whorl, abapical spiral either fully ex-posed through-

out or ;ilnu)st covered by succeeding whorls; on sub-
se(juent whorls all spirals strong and similar, or 1 or 2
on side of spire ma\- vanish on second whorl then re-
appear later at varying stages in the same positions,
these remaining weaker than adjacent spirals, 4-7 on
adult penultimate whorl, and 3-9 on base; rounded
nodules at intersections with a.\ial costae, strong on
spire, strongest at umbilical rim, of intermediate
strength on other spirals, nodules at umbilical rim less
numerous than axials on spire. Axial costae strong on
spire and at umbilical rim, almost obsolete on outer
base, evanescent over outer umbilical wall. First whorl
with microsculpture of numerous fine, crisp axial
threads, weakening and vanishing early on next whorl.
Umbilicus funnel-shaped, narrow within, rim moder-
ately angulate; outer part of wall of last whorl almost
flat, convex within, sculptured with inwardly weakening
spiral cords and a.xial riblets. Aperture subcircular, lips
thin at rim, moderately thickened within, simple. Ani-
mal (alcohol, NMNZ'M.65164) with broad flap-like
neck lobe on right side fully fused with eyestalk, 1 small
tentaculiform left neck lobe, and 4 epipodial tentacles
on each side; anterior and posterior tentacles on left
side long and slender, middle 2 minute; anterior ten-
tacle and posterior 2 tentacles on left side long and
slender, anterior second tentacle much smaller. The
radula (optical preparation) is similar to that of Z. tex-
tilis.

Type material: Holotype NMNZ M.9768, from type
locality, 3 February- 1954, M.V. Alert.

Type locality: SE of Pitt Island, Chatham Islands,
44°32'S, 176°05'W, 284 m.

Other material examined (40 specimens NMNZ):

Off Waiau River mouth, S of Kaikoura, 42°55'S,
173°43'E, 549-586 m, 18 March 1976, VJW Acheron (6,
M. 50888); up steep wall of Pegasus Canyon, NE of
Banks Peninsula, 43°14'S, 173°39'E, collected alive,
1006-512 m, 27 September 1976, RA^ Acheron (15,
M. 52779); Pegasus Canyon, NE of Banks Peninsula,
43°25.0'S, 173°26.0'E, collected alive, 485-476 m, 21
February 1979, RA' Acheron (2, M.65164); Pegasus
Canyon,' 43°31.0'S, 173°30.5'E, 2.56-293 m, 21 Feb-
ruary 1979, VJV Acheron (15, M.94763); Chatham Rise,
43°r4'S, 176°11'E, 366 m, 23 January 1954, M.V Alert
(1, M. 10839); SE of Pitt Island, Chatham Islands,
44°32'S, 176°05'W, 284 m, 3 February 1954, M.V Alert
(1, M. 10838).

Distribution (Figure 91): Off Banks Peninsula and
Chatham Islands, and Chatham Rise, 256-1006 m; col-
lected alive at 476 m and 512-1006 m from mud with
shell and/or coral.

Remarks: Zetela variabilis resembles Z. textilis in size
and in being translucent white, but differs principally in
that the axial costae on the spire whorls are broadly
rounded in section instead of sharply and narrowly an-
gulate. The original specimens oi Zetela variabilis are all

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THE NAUTILUS, Vol. 113, No. 1

Figures 116-123. Zetela kopua new species. 116. Holotype, N Bounty Trough, 1386 m, holotvpe NZOI H687 (9.00 X 9.50
mm). i!7. N Three Kings Rise, 1570-1563 m, NZOI U601 (9.10 X 9.35 mm). 118. S Lord Howe Rise, 2417-2421 m, NZOI
U226 (10.0 (est.) X 9.40 mm). 119. Paratype, N Bounty- Trough, 1373 m, NZOI P1126 (width 4.70 mm). 120-122. S Lord Howe
Rise, 1573 m, NZOI U198 (120, width 4.60 mm; 121, '6.80 X 6.50 mm; 122, 8.30 (est.) X 7.85 mm). 123. S Lord Howe Rise,
1186 ni, NZOI U197 (7.85 X 7.35 mm).

juveniles (holotvpe 2.25 X 2.70 mm, 2.5 teleoconch
whorls), and the .specie.s i,s redescribed from adult ma-
terial. It is aptly named, for adults prove to have excep-
tionally variable sculpturi'. Despite the great differences
between the e.xtremes (figures 112, 113), none of the
differences are consta)it and it seems clear that all rep-
resent a single highly polymorphic species.

Zetela annectens new species
(Figures 91, 108)

Description: Shell (holotvpe) up to 5. SO mm high,
slight!) liigher than wide at inatuiits', of moderate thick-
ness, umbihcate, chalky white (lightly solution etched).
Protoconch about 500 jim wide, tip of apical fold broad-

B. A. Marshall, 1999

Page 37

Figure 124. Map of New Zealand region show distribution
of Zetcla kopim new species (200 and 1000 m isobaths indi-
cated).

ly rounded, ,sculpture unknown (surface and rim etched
away). Teleoconch up to 4.5 whorls, first whorl rather
evenly convex; subsequent whorls with gently sloping su-
tural ramp with strongly angulate rim, weakly conve.x
above and below angulate peripheiy, umbilical rim an-
gulate. Sculpttired with roimded spiral cords and narrow,
more or less coUabral a.\ial riblets, rounded conical nod-
ules at intersections. Earliest whorls eroded but with at
least 4 spiral cords, spiral 2 (numbering abapically) be-
coming obsolete on third whorl; cords on subsequent
whorls numbering 3 on spire and 4 on base. Spire spirals
at shoulder angulation, submedially and at peripheiy,
strongly nodular; submedian and peripheral spirals
strongest and similar, summit of peripheral spiral ex-
posed on spire; basal spirals similar, more weakly nod-
ular than spire spirals. Axial riblets traversing spire and
base, evanescent within umbihcus, with increasing shell
size increasing numbers intercalating between shoulder
angulation and imibilical rim, so that munbcr of nodules
on spirals between shoulder spiral and umliilical rim be-
comes about 3 times greater than on shoulder spiral and
umbilical rim. Umbihcus deep, funnel-shaped, angulate
rim with strong, roundlv conical nodules; wail convex,
sculptured with 3 spiral threads on outer part and axial

riblets. Apeiture subcircular, lips thin at rim, weakly
thickened within, inner lip with moderate mecUan an-
gulation, separated from outer lip by strong basal an-
gulation. Animal unknown (shell of holotype too fragile
to risk extraction).

Type material: Holotype NMNZ M. 137596 and 1 ju-
venile paratope M. 137597, from type locahty, 25 January
1981, RA^ Tangaroa. collected alive.

Type locality: E of Great Barrier Island, 36°12.0'S,
176°19.4'E, collected alive, 486-655 m.

Distribution (Figure 91): Off Great Barrier Island,
486-655 (alive), in mud with foraminiferans and coral.

Remarks: Zciela annectens is strikingly similar to
western Atlantic species referred to LamcUitrochiis by
Quinn (1991), from all of which it differs, however, in
having only 4 spiral cords on the base and no secondarv*
spiral sculpture on the last few adult whorls.

Etymology: Linking, joining (Latin), an allusion to
similarity to Western Atlantic species.

Zetcla kopita new species
(Figures 115, 116-124, 135, 136)

Description: Shell up to 14.2 mm wide, slightlv wider
than high, thin, rather narrowlv umbilicate, translucent
white. Protoconch 600-620 \x,m wide, tip of apical iold
broadh- rounded; sculptured with fine, crisp, irregular
granules and 3 fine, crisp spiral threads. Teleoconch up
to about 5 whorls, first whorl evenly convex; subse<juent
whorls with narrow horizontal sutural ramp defined by
angulate rim, strongly and rather evenly convex from
ramp rim to suture within umbihcus. Reticulately sculp-
tured with narrow, crisp spiral cords that multiplv by
intercalation and enlarge to resemble adjacent spirals
and crisp, collabral axial riblets that are about as strong
as spirals, interspaces wider than each spiral and axial,
intersections finely beaded, spirals and axials roundlv an-
gulate in section. Axials continuous on first 2 whorls;
thereafter strongest on and close beside subsutural
ramp, increasing numbers of adilitional axials intercalat-
ing at irregular interviils in broad zone between abapical
side of ramp rim and inni-r third of base, so that even-
tuallv there are about twice as manv axials as in subsu-
tural zone and on inner third of base. Spiral cords num-
bering 6 or 7 on first whorl, abapical spiral more or less
covered by succeechng whorls, 19-25 on adult penulti-
mate whorl. Umbilicus funnel-shaped, narrow within,
rim evenly rounded; wall of last whorl evenly convex,
sculptured as on base. Aperture subcircular, lips thin at
rim, weaklv thickened within, simple. Animal (alcohol,
NZOI stn S154) with broad, rounded right neck lobe, 2
small tentaculiform left neck lobes, and 4 epipodial ten-
tacles on each side, anterior and posterior pairs long and
slender, median pair much smaller, eyes pigmented.
Radula (figures 135, 136) with the formula ca. 6 + 4 +
1 + 4 + ca. 6. Central and lateral teeth stout will)

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THE NAUTILUS, Vol. 113, No. 1

B. A. Marshall. 1999

Page 39

strongly flanged and groined interlocking hases, laterals
1—3 ontwardK' elongating hnt with eutting areas out-
wardly decreasing in size, lateral 4 considerahK' longer.
Cutting areas narrowly tapered, serrate on central tooth,
outer edges (only) serrate on lateral teeth, terminal cusp
on each strongest, that on lateral 4 longest. Marginal
teeth slender, terminal cusp \en long and slender, a tinv
cusp on outer etlge at about distal (juarter

Type material: Holotype NZOI H687 and l.'> para-
types (3, NMNZ M. 131532: 10, NZOI PI 133), from
type locaht), 27 October 1979, IW Tatip,aroa (S153),
collected alive. Paratvpes (12): Bounty Trough, E ofOa-
niaru, 45°24.2'S, 17.3°59.8'E, collected alive', 1373 m, 27
October 1979, RA' Tangaroa (2, NMNZ M. 131533; 10,
NZOI S154/P1126).

Type locality: Bountv Trough, E of Oamani, 45°21.rS,
173°35.8'E, 13Sfi m.

Other material examined (20 specimens): N Three
Kings Rise, 3n9.9'S, 173°05.1'E, 1570-1563 m, 9 Feb-
ruary 1988, IW Rapuhid (1, NZOI U601): Norfolk-
Ridge, 34°05'S, 168°10'E, S 1488-1556 m, 20 March
1968, M.V. Taranui (1, NZOI E870): Lord Howe Rise,
SE of Lord Howe Island, 34°59.3'S, 162°11.21'E, col-
lected alive, 1573 m, 26 September 1982, IW Tangaroa
(2, NMNZ M. 131535; 3, NZOI U19S); Lord Howe Rise,
SE of Lord Howe Island, 34°09.8'S, 163°36.7'E, 1186
m, 25 September 1982, RA' Tangaroa (1, NMNZ
M. 137598; 4, NZOI U197); Lord Howe Rise, SE of
Lord Howe Island, 38°37.3'S, 165°36.0'E, 2417-2421
m, 16 October 1982, RA' Tangaroa (1, NMNZ
M. 131534; 3, NZOI U226); Hik-urangi Trench, off Cas-
tlepoint, 41°11.9'S, 177°19.6'E, collected alive, 2200-
2328 m, 18 June 1980, RA^ Tangaroa (3, NZOI P971).

Distribution (Figure 124): Southern Lord Howe
Rise, southern Norfolk Ridge, northern Three Kings
Rise, southern Hikurangi Trench, and northern Bount\'
Trough, 1186-2421 m, living at 1373-2328 m in mud
and foraminiferal ooze.

Remarks: Specimens from the Lord Howe Rise differ
from specimens from southeastern New Zealand in hav-
ing a submedian angulation on the spire, an angulate
umbilical rim, and stronger, more widely spacetl a\ial
costae on the early teleoconch whorls, on the subsutural
zone on later whorls, and at the umbilical rim (figures
116-123). Specimens from 2417-2421 meters depth on
the Lord Howe Rise differ from specimens from both
southern New Zealand and shallower depths on the
Lord Howe Rise (1186-1573 ni, figures 120-123) in hav-

ing h\() low angulations that di\ide the spire whorls into
thirds The single shell Irom the northern Three Kings
Rise (figure 117) has an angulate umbilical rim, a weak
supramedian angulation, an extremely weak submedian
angulation, only a single spiral cord on the umbilical
wall, and a larger protoconch (width 700 |xm, cf 600-
620 |xm) and considerabK' weaker avial costae on the
early teleoconch whorls than in specimens from either
Lord Howe Rise or southern New Zealand. The single
incomplete shell from the southern Norfolk Ridge has
evenly convex whorls, an angulate umbilical rim, and nu-
merous spirals on the umbilical wall. Given the distances
between Lord Howe Rise, northern Three Kings Rise
and southeastern New Zealand and the fact that the
populations are not significantK' isolated by bathvmetry,
1 am not conxanced that ilistinct species are involved and
assume that the various forms are local populations of a
single poKmorphic species and that nioqihologically and
geographicalK- intermediate populations may eventually
be discovered.

Forms of Z. kopiia from the Lord Howe Rise are
strikingly similar to Trochus (Margarita) rhina Watson,
1886 (.syntypes BMNH 1887.2.9.302-10), from bathyal
and abvssal depths in the northeastern Atlantic, but T
rhina differs in having a smaller protoconch (width 500
(xm, cf. 600-620 |xm) and a rounded angulation on die
innbilical wall, against which the inner lip is much thick-
er and roimdly angulate. Specimens of T rhina were
well illustrated by Dautzenherg and Fischer (1896, pi.
20, fig. 15-17) and Dautzenberg (1927, pi. 5, fig. .35-
37), who misidentified them as SoIaiicUa cincta (Philip-
pi, 1836), an entirely different species (Waren, 1993,
figs. B, C, E). Lord Howe Rise specimens are also sim-
ilar to S. intcrmissa Thiele, 1925, from off southern Af-
rica at 340-1280 meters depth (Herbert, 1987), but
Lord Howe Rise and New Zealand specimens differ in
having a larger protoconch (width 600-620 |xm, cf. 450-
500 |xm) and a thinner inner lip. Trochus rhina, S. in-
tennissa, and Z. kopua obviously belong in the same spe-
cies group. Other possible congeners are Solariclla de-
licafa Dall, 1919 (holotyjx- u'SNM 205780- Kosuge,
1972, pi. 2, fig. 6) and similar northwestern Pacific spe-
cies described by Bagirov (1995), all of which differ from
Z. rhina, Z. intcrmissa, and Z. kopna. however, in that
ittial riblets are not added bv intercalation between the
subsutural zone and the umbilical rim and in other
sculptural details. Bagirov (1995) did not describe the
animals or illustrate the marginal teeth of his species but
did state that all oi the marginal teeth are similar, so it
would seem that they lack lateromarginal plates.

Figures 125-132. Radulae of species of Solariella and Batln/inopliila. 125. SolarielUi plicatiila (Murdoch and Suter), off Poor
Knights Islands, 121-11.3 ni, M..3,5876. 126. .S. vera (Powell), Middlesex Bank, 221-206 m, M. 130063. 127. S. hiteola (Powell),
Middlesex Bank, 221-206 m, M. 1.300.55. 128. S. benthicola (Powell), Middlesex Bank, 221-206 m M. 1.30079. 129. S. pensticta
new species. King Bank, 128-123 m, M. 131494. 130, 131. Bnthi/mophihi grovida new species, ex holot\pe, N Three Kings Rise,
11.37-1150 m. 132. B. tenuiseptum new species, ex holotvpe, N Three Kings Rise, 790-780 m. Scales 125-129 = 50 |jim, 1.30-1.32
= 100 fjLm.

Page 40

THE NAUTILUS, Vol. 113, No. 1

Figurti 1Ljl;-138. Radulae ot species of Airhiiniiiolia and Zetcla. 133. Archiininolin iiwrkliana (Dell), otTCape Kidnappers, 900
m, M,]i812S. 134. Zetela textilis (Murdoch and Suter), off Mayor Island, 256-311 ni, M, 35478. 135, 136. Z, kopua new species,
e.x holohpe, Bounty Trough. E of Oamani, 13S6 m. 137, 138. Z tan^aroa new species, NE of Menioo Bank, Chatham Rise, 999-
984 ni, ^.59727. Scale 1.35 = 100 fjim, others = 50 (xm.

EUinolog^: Deep (Maori).

Zetela tangawa new species
(Figures 91, 111. 114, 1.37, 138)

Description: SheO up to 7.80 mm wide, slightly wider
than high, thin, ratlier narrowly umbilicate, translucent
white. Protoconch 630 |i.m wide, obscurely and minutely
granulate, a fine spiral thread beside suture, tip of apical

fold broacUv rounded, bulbous. Teleoconch up to 4.0
whorls, strongly and rather evenly convex from suture
on spire to suture within umbihcus. Sculptured with
prominent crisp spiral cords that multipl\- bv intercala-
tion and enlarge to resemble adjacent spirals, and weak
coUabral a,\ial riblets, intersections with well-developed
nodules after first whorl, nodules on spire roimdh' con-
ical on second whorl, roundly conical or becoming fully
rounded on next whorl: nodules on adult liase rounded

B. A. Marshall, 1999

Page 41

and weaker than on spire, roundly conical and as strong
as on spire on uniltilical wall. Spiral cords ninnbering 7
on first whorl, abapical spiral more or less covered by
succeeding whorls, 10 on adult penultimate whorl, in-
terspaces wider than each spiral. Axials fold-like on spire
on first 1.5 whorls, thereafter almost obsolete on spire
and base, though stronsj on umbilical wall. Umbilicus
funnel-shaped, narrow within, rim evenly rounded, wall
of last whorl evenly convex, strongly sculptured. Aper-
ture subcircular, lips thin at riin, weakly thickened with-
in, simple. Animal (alcohol, NZOI stn. S147) with broad,
rounded right neck lobe and small tapered left neck
lobe, 4 epipodial tentacles on each side, anterior and
posterior pairs long and slender, middle pairs minute.
Radula (figures 137. 138) with the formula ca. 6 + 4 +

I + 4 + ca. 6. Central and lateral teeth stout with
strongly flanged and grooved, interlocking bases, laterals
1-3 outwardly elongating but with cutting areas out-
wardlv decreasino; in size, lateral 4 considerablv longer
and with longer cutting area. Cutting areas narrowlv ta-
pered, serrate on central tooth, outer edges (only) ser-
rate on lateral teeth, terminal cusp on each strongest.
Marginal teeth slender, terminal cusp very long and slen-
der, below which up to 6 minute cusps, fewer on outer
teeth.

Type material: Holotvpe NZOI H.686 and 8 para-
types (2, NMNZ M. 131536; 6, NZOI P1136), from t\pe
locality, 25 October 1979, IW Tangaroa (S147), collected
ahve. Paratypes (10): SE of Cape' Campbell, 42°00.8'S,
174°41.0'E; 939-1019 m, 14 Januaiy 1979, IW Tanga-
roa (1, M.60436); NE of Memoo Bank. W Chatham
Rise, 42°3S.2'S, 176°10.5'E, collected alive, 999-984 m,

II January 1979, RA^ Tangaroa (2, M.59727); SE Banks
Peninsula, 44°46.67'S. 174°91.33'E, collected alive, 692m,
5 October 1979, RA' Tangaroa (2, NZOI S143/P1134):
SE Banks Peninsula, 44°59.00'S, 174°82.67'E, collected
alive, 785m, 4 October 1979, RA^ Tangaroa (5, NZOI
S138/P1135).

Type locality: SE of Banks Peninsula, 44°30.1'S,
174°18.8'E, 760 m.

Other material examined: Oil Auckland Islands,
51°10'S, 166°37'E, 490-510 m, 31 October 1994, F.V.
Petersen, coll. M. Marinovich, fountl alive amongst sam-
ple of cirrate octopods from bottom trawl (1, M. 131554).

Distribution (Figure 91): Off western Chatham Rise,
692-1019 m, and off Auckland Islands, 490-510 m; col-
lected alive at 490-999 m from foraminiferal mud.

Remarks: Compared with Zctcia variabilis, which it
most closely resembles, Z. tangaroa differs principally
by having a larger protoconch, weaker axial sculptme,
and in lacking a basal angulation.

Etymology: Named after IW Tangaroa.

ACKNOWLEDGEMENTS

For the loan oi type and other material, I am grateful
to R. Germon and M.G. Harasewych (National Museum

of Natural History, Washington D.C.), S. Gofas (Muse-
um National d'Histoire Naturelle, Paris), B.F. Hazel-
wood (Auckland), B.W. Ha)-ward (formerly of Auckland
Institute and Museum), D.G. Herbert (Natal Museum,
Pietermaritzburg), I. Loch (Australian, Museum, Syd-
ney), R.G. Moolenbeek (Zoological Museum, Amster-
dam), S. O'Shea (National Institute of Water and At-
mospheric Research, Wellington), P. Poortman (Auck-
land), A. Waren (Swedish Museum of Natural History,
Stockholm), and KM. Way (The Natural History Mu-
seum, London). For discus.sions on Solariellinae, I am
grateful to D.G. Herbert, J.F. Quinn (Florida Marine
Research Institute, St. Petersburg), and A. Waren.
Thanks to N. Heke and M. Hall (Museum of New Zea-
lantl, Wellington) for photographv and photographic
printing, respectively, and to K. Reader (Victoria Uni-
versity, Wellington) and W St. George (Institute of Geo-
logical and Nuclear Sciences, Lower Hutt) for access to
scanning electron microscopes.

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Powell. A. W. B, 1979, New Zealand Mollusea, Marine, land
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Quinn, J, F. Jr 1979. Biological results of the University of
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zoogeography of the gastropod family Trochidae collected
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Quinn, J. F, Jr, 1991, Lamcllitrochus. a new genus of Solariel-
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Suter, H, 1908, Additions to the marine molluscan fauna of
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Suter, H. 1913. Manual of the New Zealand Mollusea, With
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Thiele, J. 1929. Handbuch der Systematischen Weichtierhmde
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Tiyon, G, W 1889, Manual of conchology. Volume II. Acad-
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Waren, A. 1990. Ontogenetic clianges in the trochoidean (Ar-
chaeogastropoda) radula, with some phylogenetic imph-
cations. Zoologica Scripta 19:179-187.

Waren, A. 1993. New and little known Mollusea from Iceland
and Scandinavia. Part 2. Sarsia 78:159-201.

Wenz, W 1938, Gastropoda. Teil 1: Allgemeiner Teil und Pro-
sobranchia, In: SchindewoH, OH, (ed) Handbuch derPa-
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Wood, S, y. 1842, A catalogue of shells from the Crag, Annals
and Magazine of Natural History 9:527-544.

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THE€7NAUTILUS

CONTENTS

Volume 113, Number 2

Juh/ 14. i, 9.9,9

ISSN \)02H- 1344

Christopher B. Bovko The authorship and status of type specimens of Urocoptis

Gary Rosenberg alleni and three species of Cerion from Cuba (Gastropoda:

Urocoptidae and Ceriidae) 43

Luiz Ricardo L. Simone Anatomv and systematics oi Anticorbula fluviatilis (H.

Adams, 1860) (Bivalvia: Lvonsiidae) from the Amazon
Basin, Brazil and Peru 48

Pablo E. Penchaszadeh Egg capsules in tlie genus Adcloiiwlon (Caenogastropoda:

Patricia Miloslavich Volutidae) from the Atlantic coast of South America 56

Mario Lasta

Paulo Marcio S. Costa

John B. Wise Reassignment of f/cim/fl luorrisoni Bartsch, 1947 from the

family AcLdidae to the Ebahdae (Gastropoda:
Heterobranchia) 64

Book Review 71

Notices 72

THE NAUTILUS 113(2):43-i7, 1999

Pafie 43

The Authorship and Status of Type Specimens of
Urocoptis alleni and Three Species of Cerion from
Cuba (Gastropoda: Urocoptidae and Ceriidae)

Christopher B. Boyko

Department of Zoology

Universit) of Rhode Island

Kingston,' RI 02881-0816 USA

and

Department of Invertebrates

American Museum ot Natural Histor\'

Central Park West at 79th Street

New York, NY 10024 USA

cboykoCS'amnh.org

Gary Rosenberg

Department of Malacology
Academy of Natural Sciences
1900 Benjamin Franklin Park-way
Philadelphia, PA 19103 USA

ABSTRACT

Urocoptis alleni, Cerion pancicostdtiim. Cerinn nllcni. and Cer-
ion victor (Urocoptidae aiwl Ceriidae) are pulmonate gastro-
pods from Cuba whose names were introduced solelv in plate
captions. Although usually attributed to Torre, 1929, all four
are Torre manuscript names introduced h\ Pilsbr\, 1929. Prob-
lems with lectot)pe designations for these names are addressed
as is the status of supposed type material in several U.S. mu-
seums.

Ket/ worcl.s: Pulmonata, Urocoptidae, Ceriidae, Tetrentodon.
Pilsbrv, Torre, Cuba.

INTRODUCTION

During the preparation of the second part oi a t\pe cat-
alog of mollusks in the American Museum of Natural
Hi,stoiy (e.g., Boyko & Sage, 1996), questions arose
about the ty{3e status of specimens of Urocoptis alleni
(now placed in Tetrentodon. Urocoptidae), as well as
Cerion paiicicostatuni, Cerion alleni, and Cerion victor
(Ceriidae) and the authorship of those names. These
names were introckiced in a plate caption in Tlie Nau-
tilus (1929, 42(3), pi. 4; figure 1, herein), and, although
attributed to [Carlos de la] Torre, we suspected they
should be attributed to Henr)' A. Pilsbry, who was then
editor of the journal. Although many authors continue
to use the name Cerionidae Pilsbry, 1901, because of its
long-standing usage, we use Ceriidae Fleming, 1818, as
it is the technically correct form of the family name.

Torre created manv manuscript names, as pointed out
by Clench and Turner (1962:.3-4), who stated that "it is
exceecUngK' luilortunate that Carlos de la Torre had giv-
en actually hundreds of names to species or sub.species
which were never described. Many Cuban collectors
have sold or exchanged a vast number of specimens un-

der these manuscript names." Pilsbiy (1929d:449) also
noted one occasion when Torre stated that he had "gi\'en
unpubhshed names " to species Pilsbn- had under study.
Welch (1934), in naming Cerion ram.scleni, wrote "the
e.xpected descriptioTi not being received from Dr. Torre,
I am here describing it under his name." Also, a man-
uscript catalog of landshells by M. K. Jacobson (1949-
1957: entr\- 8633) at AMNH Usts Urocoptis alleni as
"Torre MS." Jacobson was knowledgeable about the tax-
onnm\' of Cuban land snails, and his notation suggests
that Pilsbiy used an unpublished Torre manuscript name
for this species. The wherabouts of Torre's manuscript(s)
containing these names, or if anv such manuscript ever
existed, is unknown. Since the contents of the fype series
in such cases can depend on who authored the names,
we investigated further.

Acronyms:

AMNH Department of Invertebrates, American Mu-
seum of Natural History, New York;

ANSP Department of Malacology,, Acadenn of Nat-
ural Sciences ot Philadelphia, Penn.sylvania;

DMNH Delaware Museum of Natural Historv, Wil-
mington;

FMNH Field Museum of Natural Histor); Chicago, Il-
linois;

ICZN International Code of Zoological Nomencla-
ture;

MCZ Museum of Comparatixe Zoologv, Hanartl
University, Cambridge, Massachusetts;

USNM Department of Invertebrate Zoology, United
States National Museum (now National Mu-
seum of Natural History), Smithsonian Insti-
tution, Washington, D.C.

Page 44

THE NAUTILUS, Vol. 113, No. 2

AUTHORSHIP OF NAMES

The question of authorship for Urocoptis alleni and the
three "Torre" Ccrion species introduced in The Nautilus
(1929, 42(3). pi. 4, figs. 7-13; figures 7-13, herein) ari.ses
because no article by Torre appeared in volume 42 nl
that journal. His name is not listed in the index to that
volume or La Rocque's (1963) author index for the jour-
nal. There is no indication that Torre was the author oi
the plate captions otlier than his name appearing alter
those of the species. The names were not marked as new
nor described; only locality data were given.

Partial descriptions of Urocoptis alleni were given by
Pilsbry (1929c:141) and jauine and Torre (1972:1530),
but no full description of the species has been published
to date, presumably because the species is "muy carac-
teristica, no pudiendo contnndirse con ningima otra"
(Jaume and Torre, 1972:1530). A written description of
Cerion victor was published bv (Jlench and Aguavn
(1952:417, 419) and C. alleni has been described under
three different synonymous names (see Clench, 1957:
133). Ccrion paiicicostatiini has not been described as
such, although a subspecies, ('. p. luirrin^toni Agua\'o
and Sanchez Roig, 1953, has been described (see Rich-
ardson, 1992:78). However, these names are available
from the original publication in The Nautilus (1929,
42(3), pi. 4, figs. 7-13) because ICZN Article 12(b)(7)
allows names introduced before 1931 to be accompanied
only by an ilhistratioTi.

Cuban Caracolus illustrated on the same plate as the
four taxa in (juestion (pi. 4, figs. 1-6, and figures 1-6,
herein) were treated bv Pilsbrv ( 1929a) in tlie same issue
of the journal, but lie did not mention the Urocoptis or
Cerion species. In the next issue, ht)wever, Pilsbiy
(1929c: 141), cited the figure of Urocoptis alleni, gave
its radular formula and remarked on its similarity to U.
plicata [= Tetrciitodon pliealiis (Pfeiffer, 1.S56)] and U.
sealarina [= T. scalarina (Shuttleworth, 1852) |. Pilsbry's
authorsliip <il both articles (1929a and 1929c) that cite
plate 4 suggests that he prepared the plate and is re-
sponsible for the availabilitv of the names introduced
therein. As editor of The Nautilus, Pilsbry often com-
bined figures from several articles on one plate to save
space. For example, Pilsbiy (1929a:80), stated "To com-
plete Plate 5 a few Cuban shells collected bv Mr. d'Alte
A. Welch and the writer in northwestern (^aniagney are
figured." Perhaps he neglected to mention at the same
point that he had completed plate 4 with figures of U.
alleni. to be treated in the next issue, and several Cuban
Cerion.

Since evidence from The Nautilus was not fully con-

clusive for determining authorship of these foiu' species,
external evidence was sought. We turned to four soinces:
the pubhshed work of subsequent authors, the labels for
supposed components of the ty|)e series at AMNH,
ANSP DMNH, FMNH, MCZ, and USNM, catalog en-
tries for specimens at ANSP, and the ANSP archives.

Published Infoiination: Clench and Turner (1962)
did not list Urocoptis alleni, Cerion paucicostatuni, C.
alleni. or C. victor among Pilsbry's tiixa. All authors who
ha\'e used these names have credited them to Torre (see
hill .synonymy in Richardson, 1991:81; 1992:8, 77, 109).
Aguavo and Jaume (1950) included these names in their
list of Torre's titxa, but Bartsch (1950) did not include
plate 4, figures 7-13 in his hst of Torre's works. Because
these authors did not criticallv evaluate the vahdity of
Torre's authorship of these names, no inference of his
authorship can be drawn from their works. Torre (1929)
did publish a paper on Cuban Urocoptidae, but he did
not mention U. alleni in it.

Catalog and Label Infonnatioii: In the vial with the
illustrated specimen of U. alleni (ANSP 146901) is a clay
and cardboard mounting strip of the style u.sed by Pils-
biy, suggesting that Pilsbry prepared the specimen for
illustration. As editor of the journal, however, he often
would have prepared plates for other authors, so it is
possible that the species was illustrated at Torre's re-
quest. The ANSP catalog entiy for this lot, however,
reads "Urocoptis alleni Torre ms, " with no mention of
t\pe status. The specimens were collected by P. J. Ber-
inudez and C. C. Allen in 1927 and donated by Ber-
mudez on October 22, 1928. Because Torre did not col-
lect or donate the lot from which the illustrated speci-
men was chosen, it seems unlikely that he was involved
ill preparing the plate. The original labels for U. alleni
indicate that the specimens are "cotvpus" of "Urocoptis
alleni Torre, mss, " while the original labels for C. alleni
and C. paucico.statuni do not indicate type or manuscript
status. There is no original label for C. victor Among
the three Cerion species on the plate, Torre donated
onlv specimens of C. victor; the others were collected
and donated by C. Curry Allen.

ANSP Archives: The archives at ANSP contain a let-
ter from II, A. PilsbiA- to C. C. Allen dated November
2, 1928 (Archives Collection 98). In it, PilsbiT related
that H. N. Lowe had said that "you [Allen] had a lot of
Culian snails that would be of interest, and he | Lowe]
especially siiid that Cerion alleni ought to be figured in
the Nautilus. I would \ei-v much like to have specimens
of this for the collection of the Academy and to figure

Figures 1-13. Plate 4 from The Nautilus, 1929, 42(3), Janvuiry, recaptioned. 1-6 from Pilslin's (1929a) paper on Cuban Caracolus;
7-13 from Pilshr\- 11929bl. 7. Tetrentodon {Anafreoptis) alleni |Pilsbrv, 1929], Iecto^pe ANSP 146901. 8-9. Ccrion pnueieostatum
[Pilsbrv-, 1929]. 8. I^ctotvpe ANSP ]4(S96.5. 9. PaiaJectotype ANSP 146966. 10-11. Cerion alleni [Pilsbr\', 1929]. 10. Lectotype
ANSP'l46964. 11. Paralectotype ANSP 400,3.59. 12-13. Cerion lielor [Pilsbiv, 1929], 12. ParalcctoUpe ANSP 400.360. 13. I^ec-
totype ANSP 147406.

C. B, Boyko and G. Rosenberg, 1999

Page 45

Page 46

THE NAUTILUS, Vol. 113, No. 2

also Cerion paucicostatum." These lines confirm that the
specimens that became the types of these names came
to Pilsbrv fnirn Allen, with Torre manuscript names ap-
plied to them, and not directK from Torre.

Also in the archives (Collection 845) is the manuscript
of Torre's (1929) paper on Cuban Urocoptidae, pub-
lished in the ANSP Procccdiniis. Written in Spanish, it
was translated into English and heavily edited by Pilsiiry,
who omitted some taxa. The plates and te.xt figures do
not accompany the manuscript and were probably pre-
pared by Pilsbrv. Neither version ot the manuscript
mentions anv of the four names in question. There is no
material in the archives incUcating anv connection l:)e-
tween Torre and the pubhcation of these names in Tlic
Nautilus.

Conclusions About Authorship: The archival re-
cords at ANSP strongK suggest that Pilsbn- is respon-
sible for the publication ot Cerion paucicostatum and C.
aUcui. because he requested the specimens he illustrated
from C. C. Allen. Similarly, Bermudez, not Torre, do-
nated the illustrated specimen of Urocoptis allciii to
ANSP and there is therefore no direct connection be-
tween Torre and the publication of these three species.
Torre did donate the illustrated specimen of Cerion vic-
tor to ANSP, but there is no evidence that he had any
involvement with the description of these specimens.
Pilsbn's letter to Allen suggests that he illustrated the
Cerion species as a service to readers of The Nautilus;
presumablv he had enough room on the plate to include
C victor, specimens of which he had on hand. We there-
fore regard all four names as Torre manuscript names
first made a\'ailable b\' Pilsbn (1929b), in a pre\iousl\'
unrecognized Pilsbry publication.

Because the publication was anonvmous, PilsbiTs
name should be enclosed in square brackets (ICZN Rec-
ommendation 51A) when citing the species. Torre's
name need not be mentioned, as the ICZN does not
make provision for citing authors of manuscript names,
but its mention provides easier access to the literature
relevant to each taxon. Malacologists often place authors
of manuscript names in quotation marks, but this con-
vention is also used for authors of other kinds of un-
available names. For clarity, we recommend that the rea-
son for unavailability be specified, if for some reason it
is desirable to cite such authors. For example, Cerion
alleni [Pilsbr); 1929] (ex Torre MS) is preferable to Cer-
ion alleni "Torre" [Pilsbrv, 1929].

STATUS OF TYPE MATERIAL

Because the names in question were introduced in plate
captions, without incfication of type status or of the num-
ber of specimens examined, the contents of the t)pe
series must be determined from external evidence
(ICZN Recommendation 72B). Lectotypes have previ-
ously been designated for all four species. All known
t>pe material for these species is housed at ANSP; other
material ilistributed by Torre to other collectors and mu-

seums with identical manuscript names does not have
type status as it was not examined bv Pilsbn,; nor was it
spht from ANSP type lots based on the data from ANSP
catalog books.

Urocoptis alleni [Pilsbn, 1929]

Type locahty: Pena Blanca, Sierra Anafe [near Gu-
anajav, Pinar del Rio]

Lectotype: ANSP 146901, bv subsequent designation
of Baker (1963:221); figure 7 in Pilsbry (1929b) and
herein.

Paralectotypes: ANSP 400357, 85 specimens spfit
from lectot\pe lot, which originallv had "25+ " speci-
mens according to ANSP catalog. The specimens that
Abbott (1989: 123) figured as "paratvpes" are fikelv part
of this lot, as we kiiow of no other lot of this species
(type or non-type) which contains this many specimens.

Comments: Several other lots prevaously considered
part of the type series (AMNH 154587; ANSP 167470,
220390; DMNH 135379, 147064; FMNH 75298, 92027,
103816: MCZ 247785, 79469) were collected by people
other than Bermudez and/or Allen and were not part of
the material Pilsbry examined. The species is currently
placed in the genus Tetrentodoit Pilsbi-v, 1903, and is the
tvpe species of the subgenus Anafeeoptis Jaume & Torre,
1972.

Cerion alleni [Pilsbrv, 1929]

Tvpe locality: Antilla, [Holguin Province], Oriente

Lectotype: ANSP 146964, by subsequent designation
of Baker (1963:206, as ANSP 146964a); figure 10 hi Pils-
bn- (1929b) and herein.

Paralectotypes: ANSP 400359, 3 specimens spht
from lectoty[3e lot, which originallv had 4 specimens; lot
includes specimen in figure 11 of Pilsbn' (1929b) and
herein.

Comments: We have examined USNM 346663 from
the Bahamas, laiieled b\' Bartsch with this name and the
status 'type, " and conclude that is not a tvpe specimen,
nor even this taxon, but rather a homonvmous Bartsch
manuscript name.

Cerion paucicostatum [Pilsbiy, 1929]

Type locality: Cape M;usi, [Guant;inamo Pnmnce,
Oriente]

Lectotype: ANSP 146965, by subsequent designation
of Baker (1963:207, as ANSP 146655a): figure 8 hi Pils-
bry (1929b) and herein.

Paralectotypes: ANSP 400358, 1 specimen split from
lectotvpe lot, which originalK' had 2 specimens; ANSP
146966, 2 specimens, inclucbng specimen in figure 9 of
Pilsbn (1929b) and herein.

C. B. Boyko and G. Rosenberg, 1999

Page 47

Comments: Specimens ironi Miller Hill, southern
Eleuthera Island, Bahamas labeled "paratvpes" (e.g.,
FMNH 42323, 119727, 119777) are not part' of the t\pe
series for this taxon, but are paratvpes of C paitcico-
statitm Clench, 1934 (non [Pilsbry,' 1929]) (= C. indi-
ammim Clench, 1934; see Clench,' 1957:149, 157; Rich-
ard.son. 1992:51).

Ccrion lUior [PiLslm-, 1929]

Type locality: Caleta de Ovando, [Guantanamo Prov-
ince], Oriente. [ANSP catalog adds "about 40 miles W
of Punta de Maisi on the south coast."]

Lectot>pe: ANSP 147406, by subsequent designation of
Clench and Aguayo (1952:419, by inference of holotvpe);
figure 13 in Pilsbiy (1929b) and herein.

Paralectotypes: ANSP 400360, 2 specimens; ANSP
400361, 1 specimen, figure 12 in Pilsbn- (1929b) and
herein; both lots spfit from the lectotvpe lot, which
originally had 4 specimens.

Comments: Baker (1963:207) independently
designated ANSP 147406a (Pilsbn- 1929b, fig. 12) as
lectotvpe. Clench and Aguavo's (1952) designation has
priority, making Bakers invalid, and the specimen he
designated is a paralectotvpe, now cataloged as ANSP
400361. The specimens of C. victor cited by Clench and
Aguayo (1952), MCZ 128975 and 181758, as well as
MCZ 181756, 181757, 192282, 192283 and ANSP
323229 have no original data labels and are not from the
cited tyjDe locahty; thev are therefore not considered
paralectotypes. Abbott (1989:72) figured a dozen
"paratvpes" of C. victor, but, because no recognized
paralectotvpe lot contains this manv specimens, they are
also excluded from the type series and may be part of
one of the above cited lots of excluded specimens.

ACKNOWLEDGMENTS

Thanks to John Slapcinskv (FMNH), Tim Pearce
(DMNH), Daniel Graf (MCZ), and Robert Hershler
(USNM) for pnniding data and loans oi specimens from
their respective institutions. Thanks also to Paula Mik-
kelsen (AMNH) for reviewing an early version of the
manuscript, Alan Har\ev (AMNH) for comments and
suggestions, and ]errv Harasewvch (USNM) for contrar-
ian comments that spurred us to dig deeper in our re-
search. Two anonymous reviewers offered comments
that fin"ther improved the manuscript.

LITERATURE CITED

Abbott, R. T. 1989. Compendiuiii of Landshells. Ameritaii
Malacologists, Melbourne, Florida, 240 pp.

Aguayo, C. G. and M. L. Jaume. 1950. Bibliografia malaeolo-
gioa de Dou Cados de la Torre. Agoni'a [for 1950]:119-
12S.

Baker, H. B. 1963. T\pe land snails in the Academy of Natural
Sciences ot Philadelphia part II. Land Pulmonata, exclu-
si\e ot North America nortli of Mexico. Proceedings of
the Acadenn of Natural Sciences of Philadelphia 115:191-
259.

Bartsch, P. 1950. Carlos de la Torre y de la Huerta. American
Malacological Union News Bulletin and Annual Report
[for 1950]: 1-6.

Boyko, C. B. and W. E. Sage III. 1996. Catalog of Recent tyjie
specimens in the Department ot Invertebrates, American
Museum of Natural Histon. II. Mollusca Part 1 (Classes
Aplacopliora, PoKplacophora, Gastropoda [Subclass Op-
isthobranchia], BivaKia, and Scaphopoda). American Mu-
seum Novitates 3170:1-50.

Clench, W. J. 1957. A catalogue of the Cerionidae (Mollusca-
Pnlmonata). Bulletin of the Museum of Comparative Zo-
olog>- 116(2): 121-169.

Clench, W. J. and C. G. Aguavo. 1952. The scalariniiiii .species
complex {Uinbonis) in the genus Ccrion. Occasional Pa-
pers on Mollusks I(17):413^40.

Clench, W. J. and R. D. Turner 1962. New names introduced
by H. A. Pilsbry- in the Mollusca and Crustacea. Academy
of Natural Sciences of Philadelphia Special Pubhcation 4:
1-218.

[ICZN] International Commission of Zoological Nomenclature.
1985. International Code for Zoological Nomenclature,
third edition. Intemational Tnist for Zoological Nomen-
clature, London, 338 pp.

Jacobson, M. K. [I949-I957]. Foreign Land Shells [and] Unit-
ed States Land Shells. Unpublished manuscript. American
Museum of Natural History, New York, 286 pp.

lainiie, M. L, and A. de la Torre. 1972. Catalogo de la fauna
Ciibana — XXIX-XXXVI — Los Urocoptidae de Cuba
(Mollusca — Pulmonata) (nos. 1-8). Circulares del Museo
V Biblioteca de Zoologia de la Habaiia 13(.351 ):1526-1649.
[Reprinted in 1976 in Ciencias Biologicas senes 4(53):1-
122 pp.]

La Rocijue, A. 1963. Author Index to The Nautilus \olumes
3-75 and its predecessor The Conchologists' Exchange
Volumes 1 and 2. Columbus, 279 pp.

Pilsbry, H. A. 1929a. New Cuban species of Caracolus The
Nautilus 42(3):78-S0. pi. 4, figs, 1-6, pi. 5, figs. 2, 8-11.

[Pilsbry, H. A. 1929b]. [Urocoptis nlleni. Cerion pmicicostatum,
C nUcni. C. victor]. The Nautilus 42(3):pl. 4, figs. 7-13.

Pilsbn, H. A. 1929c. Urocoptis alleiii Torre (Nautilus, Jan.,
1929, pi. 4, fig. 7), The Nautilus 42:141.

Pilsbiy, H. A. 1929d. Stmhes on West Indian mollusks, II: The
locomotion of Urocoptidae and descriptions of new fonns.
Proceedings of the Academy of Natural Sciences of Phil-
adelphia 81:449-467, pis. 12-15.

Richard.son, C. L. 1991. Urocoptidae: catalog of species. Tiyon-
ia 22:1-245.

Richardson, C. L. 1992. Cerionidae: catalog ot species. Tiyonia
25:1-121.

Torre, C. de la. 1929. New Cuban Urocoptidae. Proceedings
of the Academy of Natural Sciences of Philadelphia 81:
44:5^47, pi. 12, figs. 5-8, pi. 13, figs. 14-18, pi. 15, figs.
14-15, 8 text figs.

Welch, d'.\. 1934. New Cuban Land Shells from Oriente and
Camaguey Provinces. The Nautilus 47:104-108, pi, 11.

THE NAUTILUS 113(2):48-55, 1999

Page 48

Anatomy and Systematics o{ Ant icorhula fluviat ills
(H. Adams, 1860) (Bivalvia: Lyonsiidae) from
the Amazon Basin, Brazil and Peru

Luiz Ricardo L. Simone

Museu de Zoologia da Universidade

de Sao Paulo
CaLxa Postal 42694
04299-970 Sao Paulo. BRAZIL
lrsimone@usp.br

ABSTRACT

Anticorbula fluvintilis (H. Adams. 1860) is an enigmatic fresh-
water bi\aKe not closelv related to the usual groups of fresh-
water bivaK'es. Due to its rarit\' and conse(juent lack of spec-
imens, the validit\' of the genus and species, their occurrence
in the freshwater habitat, and even their existence, have been
questioned. Other species of closely similar taxa are common
in estuarine areas of northern and western South America. The
familial allocation of Anticorbula has also been problematic.
Specimens of A. fluviatilis (with soft parts) collected in the
Amazon Basin including material from the t\pe localits' were
examined, and the validity of the genus and species confirmed.
A detailed description of the anatomv is provided as the foun-
dation for future s\stematic comparisons. Diagnostic characters
include possibly vestigial lithodesma, small foot with bvssus,
mantle lobes almost entirely fused (with pedal and pallial ap-
ertures only), lack of siphons, very large posterior retractor
muscles of foot, stomach with ducts to digestive dix'erticula only
on right side, and stvle sac separated from intestine The pres-
ence of these characters suggests allocation within the famiK
Lvonsiidae. but this familial allocation should reniam provi-
sional until a better definition of the Lyonsiidae is made avail-
able.

Keij words: Lvonsiidae. Antia'rbiihi fluviatilis. Amazon basm.
Brazil. Peni. svstematics. morphology.

INTRODUCTION

The Amazon Basin is a region of incomparable hiodi-
versit)'; despite this, its freshwater and terrestrial mol-
lusks are verj' poorly known. Unusual organisms are of-
ten found, including die species re-described herein.

Himclla fluviatilis, an unusual freshwater mussel col-
lected in the Maraiion River, Peru, was described by H.
Adams in 1860. The genus name Himella was pre-oc-
cupied (Dallas, 1852, Insecta), and a new name was pro-
posed, Anticorbula Dall, 1898, as subgenus of the ma-
rine genus Corbula Bruguiere, 1797. Morrison (1943)
described the genus Guianadcsma for an estuarine spe-
cies, G. simiosum, from Guyana, which is verv similar to

Anticorbula fluviatilis. The genera Anticorbula and
Guianadcsma were considered synonyms of Ostomija
Conrad. 1874 by at least Keen (1969), and Vaught
(1989). However,' Nuttall (1990) pointed out that Osto-
mi/a was a distinct ta.\on and considered Guianadcsma
as valid, an opiTiion later shared b\' Leistikow and Jans-
sen (1997). Nuttall (1990) also suggested the possible
synonymy between Guianadcsma and Anticorbula. This
latter author considered it as nomcn dubium based on
lack of data on tvpes and substantiated locahts' data (p.
319). On the other hand. Nuttall did not explore further
the differences or similarities between these two taxa.

Although the Maranon Rix'er. type locaUty of A. flu-
viatilis, is very cUstant and far removed inland from the
eastern Amazon coasthne, most reported occurrences
situate this and related species on estuarine and coastal
areas (Altena, 1969, 1971; Morrison. 1993; Nuttall,
1990). These reports naturally bring into question
whether these organisms actually occur in tnie fresh-
water habitats. On the other hand. Adams did not show
any doubts about the provenance of A. fluviatilis, and
Pilsbrv' (1944) examined additional material from eastern
Peru: Essequibo and CmhiI rivers, and even from Ma-
ranon River itself. He also figtned some specimens of A.
fluviatilis for the first time. Also addressing this ques-
tion, Leistikow and Janssen (1997) published their find-
ing of specimens identified as Guianadcsma sinuosum
from Manaus, Brazil, a site located miilstream on the
Amazon Basin.

The family allocation of the above mentioned genera
has also been inconsistent; some authors allocate them
to Lyonsiidae (Pandoroidea) (Keen, 1969; Altena, 1971),
other authors to CorbuLidae (Myoidea) (Pilsbry, 1944;
Nuttall, 1990; Leistikow & Janssen, 1997).

E.xcept for the schematic figure and a brief description
of the soft parts of G. sinuosa (Morrison, 1943), no other
anatomical data on South American freshwater and es-
tuarine Konsiids has been published. The abo\^e men-
tioned references dealt onh' with conchological and en-
vironmental data.

L. R. L. Simone, 1999

Page 49

The type specimens of A. fluviatilis have not been
located (Altena, 1971:82: Nuttall, 199():29(); F. Naggs,
pers. comni.), which precludes the resolution of the sys-
tematic problems.

A lot with preserxed specimens was collected bv a
team of ichthyologists from the Museu de Zoologia in
the Trombetas Ri\er, Para, Brazil (Amazon Basin), a
freshwater localitx' well removed tr(jm the coast. The
identification of these specimens as A. fluviatilis was at-
tained upon examination of the hterature and compari-
son with material from the t\pe localitv. The present
detailed study of their moii")holog\ should help clarit\
the above mentioned conceptual problems and estabhsh
the identity of the species. It should also provide the
foundation for further comparative studies of specimens
from other (mainly estuarine) areas.

MATERIAL AND METHODS

The specimens were initially preserved in 4% formalin
and transferred to 70% ethanol. The dissection was con-
ducted under a stereomicroscope, with the specimens
immersed in water. Serial sections were made using
standard histological techniques and stained with Mal-
lory. All drawings were made with the aid of a camera
lucida. The intestinal loops and other visceral stnictures
were examined through dissections and serial sections.
The SEM figures were obtained at the "Laboratorio de
Microscopia Eletronica do Instituto de Biociencias da
USP".

The specimen presen'ation was not good enough to
allow for a detailed examination of ciha; these are not
showni in detail in the serial section figures (Figures 20-
22).

The following abbreviations are used in the figure cap-
tions: aa, anterior adductor muscle; an, anus: au, auricle:
by, byssus; dd, ducts to digestive di\'erticula; dli, dorsal
hood; as, esophagus; ex, excurrent aperture; fa, foot ap-
erture; ft, foot; ia, intestinal aperture: id, inner demi-
branch; in, intestine; is, incurrent aperture: Ig, ligament;
li, lithodesma-like projection of mantle: 11, left mantle
lobe; mb, mantle border; mo, mouth; mu, mantle ventral
union; od, outer demibranch; pa, posterior adduct(jr
muscle; pe, periostracum; pi, pallial line; pp, palps: pr,
posterior protractor muscle of foot; ps, pallial siTius; ra,
anterior retractor muscle of foot; ri, hinge; rp, posterior
retractor muscle of foot; rt, rectum; r\. right vaKe; s;i.
sorting area; sp, style sac aperture; ss, style sac; st, stom-
ach; sy, crystalline style; te, testis; um, umbo; ve, ventri-
cle; vg, visceral ganglia; vm, visceral mass.

The following institutional ;ibbre\iations are used in
the text: ANSP, Acadenn' of Natural Sciences of Phila-
delphia. Philadelphia; BMSM, The Biiiley- Matthews
Shell Museum, Sanibel, Florida: MZSP, Museu de Zool-
ogia da Universidade de Sao Paulo, Sao Paulo, Brazil.

SYSTEMATIC DESCRIPTIONS

Genus Anticorbula Dall, 1898

Himelh H. Adams, 1860:20.3 (non Dallas, 1S52); t\pe species
(by monot\p\): H fiuviatilis H Adams, I860; pre-occu-
pied name (NuttalK 1990:.319).

Corbula {Anticorbula} Dall, 1898:8.39, iionwn novum.
Aloiilis (Anticorbula). — Morretes, 1949:47 (in part),
Anticorbula.— Keen. 1969:847; Vaught, 1989:140 (in both ref-
erences as sviionvni of Osionuja Conrad, 1874).

Remarks: Anticorbula is the oldest valid name and
should be used for this species. The similarity between
the type species of the monotypic genera Anticorbula
and Guianadcsma is clear; thus these 2 genus names
may be synonyms. However, the s\iionymy will be re-
solved only after a detailed anatomical analysis of G. .sf'/i-
uo.'ium (t\pe oi Guianadcsma).

Anticorbula fluviatilis (H. Adams, 1860)
(Figures 1-22)

Himella fluviatilis H. Adams, 1860:203 [Maranon Ri\er, Pern].
Corbula (Anticorbula) fluviatilis. — Dall, 1898:839.
0,s(()((i(y«^(iii;flfi7/s.— Pilsbrv, 1944:148 (fig. 1 -I- pi. 11, figs.

42-44)
Aloidis (Anticorbula) ftuiiatilis. — Morretes, 1949:47.
Anticorbida fluiiatilis. — Altena, 1971:82.
Guianadesiiui sinuosuin. — Leistikow and Janssen, 1997:17-20

(figs. 1-3) (non Morrison, 1943).

De.scription: Shell (Figures 1-9, 19): Of medium
size (up to 25 mm), somewhat elliptical, inequivalve, ir-
regular (Figures 2, 3), ventral edge sigmoid (Figures 6,
19). Umbones inlaid, located on anterior third of hinge,
generally eroded. Periostracum semi-transparent,
brown, covering hgament (Figure 19). Color fight beige,
with browm areas on posterior slope (Figure 1). Anterior
region roundeil, posterior region somewhat straight,
oblique. Sculpture lacking except for concentric growth
lines and undulations. Hinge edentulous, with a broad
tooth in each valve posterior to umbones, each one bear-
ing a longitudinal furrow containing most of ligament
(Figures 4, 5, 9, 19). These broad teeth as\'mmetrical
(Figure 19). Left valve tooth somewhat parallel to valve
margin and close to median fine: right valve tooth
oblique, with posterior region relatively chstant from me-
dian line and separated from the vahe border by a
smooth area, which gradually approaches from median
fine to become similar to its analogue. Ligament partly
exposed, anterior extremity only a narrow thread. Pos-
teriorl\- to lunbo ligament gnidualK liecomes thick and
broad at ;ibout V3 of its length. Posterior half of this
broad region hollow, opened posteriori)', filled bv a con-
ic, sharp projection of mantle (Figure 10), which sud-
denly becomes a narrow thread in its posterior h;ilf In-
ner surface of each valve smooth, not nacreous (Figures
4, 5). Two anterior scars (Figures 4, 5. 9), a small, cir-
cular, dorsal of anterior foot retractor muscle, and other
large, reniform, ventral to anterior adductor muscle. P;d-
lial line simple, with a broader anterior region; a shallow
sinus is present behveen this broad region and adductor
muscle scar (Figure 9). A large and somewhat triangular
posterior scar composed by two roughly equal (in cross-
section) muscles, one dorsal to posterior foot retractor
muscle, other ventral to posterior adductor muscle, Ven-

Page 50

THE NAUTILUS, Vol. 113, No. 2

Figures 1-8. Shell of Anticorhiila fluviatilis. 1. Left vaKe showing most frequent color pattern; 2, 3. Two specimens showing
\ariation in outHne; 4. Right valve, inner \iew; 5. Left valve, inner view; 6-8. SEM micrographs of 1 specimen. 6. Ventral view;
7. Left view; 8. Dorsal view. Scale lines = 2 mm.

tral region of protractor muscle of foot small, triangular.
Further details on shell morphologv' were given by Leis-
tikow and Janssen (1997:17-18, figs. 1-2).

Mantle: Pale cream in color, in some specimens with
dark brown spots in imibonal region. Mantle lobe bor-
ders almo.st entire fused (Figures 11, 12, 20), a narrow,
funnel-like, conic pedal aperture (Figure 12). Incurrent
aperture surrounded by two rows of stubby papillae
(Figures 12, 15); each papilla white in color, dark browii

minute spots on base, rounded tip. E.xcurrent aperture
narrow, edged by thick borders without papillae (Figure
15). Lobes union evolves inner folds (Figure 20),

Foot ami muscles: Foot small, sessOe in antero-ventral
half of visceral mass (Figures 12, 13). Byssal furrow oc-
cupying most of foot ventral region; anterior half a nar-
row, deep furrow (Figure 13); posterior half broad,
opened, bearing two large bvss;il bundles (Figures 12, 13,
15). Anterior bvssal bundle composed bv several ribbon-

L. R. L. Simone, 1999

Page 51

Figures 9-12. Anatomy oi Anticorbula fltiviatilis. 9. Shell, left valve, inner view, showing muscle scars and hinge; 10. Ligament
extracted, left view, a quarter was artificially cut to show inner mantle projection, possibly a vestige of lithodesma; 11. Hole specimen,
left view with left valve extracted, outer surface of mantle exposed; 12. Mantle cavity, left view, infra-branchial portion of mantle
extracted by an incision surrounding insertion of ctenidiiun, visceral mass, adductor nmscles and in middle region of ventral mantle
fusion. Scale lines = 1 mm.

like, flattened fibers; posterior buncUe broad and massive.
Anterior and posterior adductor muscles somewhat equal
in size (Figure 11). Anterior adductor muscle located very
anteriorly and ventrally. A pair of large visceral ganglia
attached to its ventral surface of posterior adductor mus-
cle (Figure 18). Pair of anterior foot retractor muscles
small, slender, inserted in anterior region of foot. Pair of

posterior foot retractor muscles large (Figure 16), flat-
tened at origin (longer antero-posteriorly), just dorsiil and
anterior to posterior adductor muscle, each situated very
close to other member of pair, separated only by a narrow
space where rectum nms. Posterior retractors graduaUy
become narrow and cylindrical, inserted in foot muscles
just dorsally to byssal gland.

Page 52

THE NAUTILUS, Vol. 113, No, 2

Figures 13-16. Anatomy ot Anticorbula Jiuoiatilis. 13. Mantle cavity, ventral view, ctenidiuni part deflected, mantle e.\tracted;
14. Detail of region of left palp and adjacent structures, left view, outer heniipalp deflected to show its inner folds; 15. Detail of
incurrent and excurrent apertures (incurrent inferior), left, slight posterior view; 16. Digestive tubes and part of muscular and
circulatory systems seen as the animal was transparent, left view, left wall of stomach e.xtracted. Scale lines = 0..5 mm.

Mantle cavittj organs. Inner surface of mantle smooth,
without additional stnicture.s (Figure 11), mantle edge
thick. Ctenidium long, about % of length of mantle cav-
ity and about half of dorso-ventral distance (Figures 12,
13); filaments very narrow, producing a uniform surface.
Outer demibranch not covering inner demibranch. Out-
er demibranch divided into two regions (Figure 12): dor-
sal V3 with obhque filaments, ventral % with filaments
that become perpendicular to free edge of ctenidiuni

Anterior edge of outer demibranch fused with mantle.
Inner demibranch simple, occupving little more than
half of voknne of outer demibranch, onlv peqiendicular
filaments present. Anterior edge fused with mantle and
antero-ventral angle partially covered by palps. Food
grooves present on free edges of both demibranchs (Fig-
ure 20). Profile of ctenidiinn filaments is shown in Fig-
ure 22.

Digestive si/stcm: Palps somewhat small, antero-pos-

L. R. L. Simone, 1999

Page 53

Figures 17-18. Anatomy oi Anticorbiilii ftuiiatilis. 17. Stom-
ach and adjacent region, dorsal view, anterior part of gastric
dorsal wall extracted, posterior part deflected to right jointed
with the intestine loop which crosses this area; 18. Detail of
posterior addnctor mnscle, left, slight posterior view, left lobe
of mantle extracted to show anus and visceral ganglia. Scales
= 0.5 mm.

teriorlv elongated, free extremitv tapering off to a round-
ed tip (figures 12-14). Outer surface of palps smooth.
Inner surface of palps with several broad, oblique folds;
profile of folds showni in Figure 21. Each fold with
rounded e.xtremities, ending at some distance from palp
edge, with smooth inner margin (Figure 14). Inner folds
evanesce near mouth (Figure 13). Mouth large, with
smooth inner surface. Esophagus wide, short, flattened
dorso-ventrally, curved, not attached to, separated from

adductor muscle (Figure 16), inner siuface with several,
uniform, longitudinal, low folds (Figure 17). Stomach
(Figures 16, 17) spacious, located just posterior to shell
umbones. Esophageal folds end suddenly at same height
along esophagus. Two small orifices of digestive di\'ertic-
ula present on left anterior region of stomach. Right an-
terior region characterized bv valve-like, small septum
that represent anterior Umit of a dorsal hood. Dorsal
hood short, broad, with rounded distal extremity, inter-
nally with several low, longitudinal folds. A constriction
present in stomach region opposite to dorsal hood. Pos-
terior region of stomach with two orifices (Figures 16,
17): aperture of style sac, larger, on left side; intestine
origin, smaller, on right side. Style sac long, straight, sep-
arated from intestine (Figures 16, 20); posterior extrem-
ity rounded, bulging in posterior-ventral extremity of vis-
ceral mass (Figures 13, 16). Intestine (Figures 16, 17)
narrow, without inner folds; intestine runs anteriorly,
surrounding right ventral surface of stomach. After
forming a loop near esophagus intestine is directed pos-
teriork, brodering dorsal surface of foot; near right pos-
terior region of style sac intestine is chrected antero-dor-
sally to postero-dorsal region of stomach; in this region,
a loop touches dorsal gastric wall on stvle sac and intes-
tine apertures. This loop iims dorsally through ventricle,
between posterior foot retractor muscle and postero-
dorsal surface of posterior adductor muscle. Anus rep-
resented by a small orifice without papilla or glands (Fig-
ures 16, 18).

Visceral soc: Digestive diverticula almost completely
restricted to space around stomach, in mid-dorsal region
of animal. Remainder of visceral sac filled by gonochor-
istic gonad (Figure 20), which surrounds digestive tubes,
and is separated into several elongate, somewhat large
acina. Transversal muscles very small, present mainly in
ventral, peri-pedal regions.

Habitat: Found in river with mild current, about 5 m
ill depth, attached by byssus into furrows of dead tree
bark.

Material examined: BRAZIL: MZSP 28788, 3 spec-
imens. Para; Trombetas River, 1°41'51"S 55°51'8"W,
Lundberg et al. col, 24 Oct. 1994; MZSP 28787, .5 spec-
imens, Xingu River Porto de Moz, 5 specimens, 9 Aug.
1968; MZSP 28789, 250 shells, and BMSM 1007, 3
shells, Alenquer River, Ponta de Pedregulho, Lago
Grande. 21 Sep. 1969, col. EPA no. 690073-4; PERU:
ANSP 125525, 2 shells, Maranon River. Charles M.
Wheatley Collection, (material examined by Pilsbry,
1944).

DISCUSSION

The problematic allocation of the genus Anticorbula at
the family level is mostly due to the poor systematic
definitions of the relevant bivalve families; apparently
there is no sound information (good diagnoses based on
.svnapomoqihies) on each of the following families that

Page 54

THE NAUTILUS, Vol. 113, No. 2

Figures 19-22. Shell and serial sections of Antieorbiila fltiviafilis. 19. Shell, detail of anterior half, ventral view, valves partially
opened to show hinge region. Scale Une = 2 mm; 20. Transversal (frontal) section in approximately middle region of animal. Scale
line = 1 mm; 21. detail of middle region of outer hemipalp, inner folds cut transversally; 22. Transversal section of ctenidium
filaments at midhne, outer demibranch. Scale lines (21-22) = 0.1 mm. Histological sections (20-22): Mallorv stain, 5 (jim.

would allow for the clear cut, indisputable allocation of
Anticorhuh to any oi'them. As commented in the intro-
duction, vanous authors have allocated this genus in the
families Corbulidae (Myoidea) and Lyonsiidae (Pando-
roidea), as well as some affinity with the Myidae (Myoi-
dea), Hiatelloidea and Thraciidae (Pandoroidea) can also
be demonstrated.

The familial allocation oi Anticorbula to is not only a
problem of nomenclatural nature. Not belonging to the
most common freshwater groups such as Unionoida,
Dreissenoidea and Corbiculoidea, Anticurbula seems to

represent an independent invasion of the freshwater en-
vironment.

In present study, Auticorbula is provisioniilly allocated
in the family Lyonsiidae. Such allocation is based on con-
chological characters (outline, thick periostracum cov-
ering inclusive the hinge, almost edentulous hinge, a
projection of the mantle within ligament which can be
a vestige of the hthodesma). On the other hand, anatom-
ical studies on lyonsiids (Yonge, 1952; Thomas, 1993)
have described animals with two ducts to the tligesti\e
diverticula, fusion of the style sac with the intestine, and

L. R. L. Simone, 1999

Page 55

a siiial] pair of posterior foot retractor muscles, all char-
acters that clitler from those in Anticorbula.

Aniicorbula is similar to members of Corbiculidae
(Yonge, 1947) in having poor developed siphons and only
one duct to disiestive diverticula. However, Anticorbula
differs in hinge characters, in having a reduced foot with
byssus, the presence of posterior foot retractor muscles
and a st\le sac separated from intestine.

Similarities with members of Thraciidae are overall
shell shape, presence of a lithodesma in hgament and
fusion of mantle edge (Morton, 1995). However, Anti-
corbula differs m;iinlv in lacking developed siphons, in
having onlv one duct to digestive diverticula, a separa-
tion between style sac and intestine and by the byssus.

Anticorbula is similar to members of Mvidae in having
fusion of mantle edge and, at least of Cnjptonuja cali-
fornica (Conrad) (ct. Yonge, 1951) in having not so de-
veloped siphons and separation of st)le sac from intes-
tine. It differs mainly in having a byssus, the reduction
of the foot and the absence of hinge teeth.

Based on data from the (Yonge, 1971), Anticorbula is
similar to members of Hiatellidae in shell shape, in hav-
ing thick periostracum, reduction of hinge teeth and of
foot. In contrast, Anticorbula lacks siphons, has only one
duct to digestive diverticula and separated style sac and
intestine.

The present lyonsiid attribution to Anticorbula is,
then, provisional. All of the above mentioned families
have estiiarine representatives and could perfectly in-
clude Anticorbula and its related genera Guianadcsma
and Ostomija.

The shell of A. fluviatilis is very similar to that of
Guianadcsma sinuosum (Morrison, 1943; Nuttall, 1990).
The morphology of the soft parts of G. sinuosum is veiy
superficially kTiown (Morrison, 1943:50-51, fig. 1), and
is also similar to inner moq^hology of A. fluviatilis. Both
species were not svnonvmvzed herein mainly due their
different habitats (A fluviatilis. tnie freshwater; G. sin-
uosum, low salinity, estuarine), and also due to the lack
of a detailed anatomical study of this latter species.

ACKNOWLEDGMENTS

Specials thanks to Dr Jose H. Leal, BMSM, and Dr.
Kevin S. Cummings, Illinois Natural Historv Survey, for
important suggestions antl criticisms of the manuscript.
To Dr. Guido Pastorino, Argentina, for providing refer-
ences. To Dr. George M. Davis and Dr. Garv' Rosenberg,
Academy of Philadelphia, for loaning the specimens
from type localitv examined bv Pilsbr)'. To Fred Naggs,
The Natural Histt)rv- Museum, London, for searching tor
the types oi A. fluviatilis. To Dr. Luiz C. F. Alvarenga,
Dr. Ceha N. Ricci (Museu Nacional, Univer.sidade Fed-
eral do Rio de Janeiro) and Dr. Maria C. D. Mansur
(Pontificia Universidade Catolica do Rio Grande do Sul)
for comments on the initial stages of this project. This
research was partly supported by a Research Grant # 96/

6756-2 of Fundayao de Amparo a Pesquisa do Estado
de Sao Paulo (FAPESP).

LITERATURE CITED

Adams, H. 1S60. Description of a new genus of fresliwater
bivafve Mollusca, belonging to tlie family Corbulidae,
from the collection of Hugli Cuming, Esq. Proceedings of
tlie Zoological Society of London 28:203.

Altena, C. O. van Regteren. 1969. The marine Mollu.sca of
Surinanie (Dutch Guiana), Holocene and Recent. Part 1:
general introduction. Zoologische Verhandelingen 101:1-
48, 4 pis.

Altena, C. O. van Regteren. 1971. The marine Mollusca of
Suriname (Dutch Guiana) Holocene and Recent. Part II:
BivaKia and Scapliopoda Zoologische Verhandelingen
i 19:1-100, 10 pis., 1 map.

Dall, W, H. 1898. Contributions to the Tertiary Fauna of Flor-
ida. Transactions of the Wagner Free Institute of Science
of Philadelphia 3:571-947. ^

Keen, A. M. 1969. Family Lyonsiidae Fischer, 1887. In: Moore,
R.C. (ed.) Treatise on Invertebrate Paleontology-, Part N,
\'olume 2 (of 3), Mollusca 6, Bivalvia, pp. N845-N847.

Leistikovv, A. and R. Janssen. 1997. A record of tlie bivalve
Guianadesma sinuosum Morrison from the central Ama-
zon basin (Bivalvia: Corbulidae). Basteria 61:17-22.

Morretes, F. L. 1949. Ensaio de catalogo dos moluscos do
Brasil. Arquivos do Museu Paranaense 7:1-216.

Morrison, (. P. E. 1943. A new tvpe of fresh water clam from
British Guiana. The Nautilus 57:46-52, pi 8.

Morton, B. 1995. The ecology and functional moqihologv of
Trigonothracia jinxingae (Bivalvia: Anomalodesmata:
Thracioidea) from Xiamen, China. Journal of Zoologv' 237:
445-468.

Nuttall, C. P. 1990. A review of the Tertiarv non-marine niol-
luscan faunas of the Pebasian and other inland basins of
north-west South America. Bulletin of the British Muse-
um of Natural Historv (Geologv) 45:16.5-371.

Pilsbrv, H. A. 1944. Moiluscan fossils from tlie Rio Pachitae
and vicinity in eastern Peni. Proceedings of tfie Academy
of Natural Sciences of Philadelphia 96:137-153, pis. 9-11.

Tliomas. K. A. 1993. The functional moqiliologv' of the diges-
tive system of Lyonsia htjalina Conrad. 1831 (Bivalvia: An-
omalodesmata: Pandoroidea) |ounial of Moiinscan Stud-
ies .59:17.5-186.

V'aught, K. C. 1989. A classification of tlie living Mollusca.
American Malacologists, Inc., Meltiounie. Florida, 189
pp.

Yonge, C M. 1947. On the habits and adaptations of Aloidis
(Corhula) gibha Journal of the Marine Biological Asso-
ciation of the United Kingdom 26:358-376.

Yonge, C. M. 1951. Studies on Pacific coast moUusks I. On the
structure and adaptations of Cnjptomt/a califomica (Con-
rad). Universitv' of California Publications in Zoolo^v' 55:
395-^20.

Yonge, C. M. 1952. Studies on Pacific coast mollusks V. Struc-
ture and adaptation in Entodesnui saxicola (Baird) and
Mi/tilinuriii nuttallii Conrad with a discussion on e\'olu-
tion within the faniilv Lyonsiidae (Eulaniellibranchia).
Universitv of California Publications in Zoologv' 55:439-
449.

Yonge, C. M. 1971. On functional moqihology and adaptive
radiation in tlie bivalve superfaniily Saxicavacea (Hiatclla
{ = Saxicava), Saxicavella. Pnnomi/n. Pfinope. Cytiodaria).
Malacologia 11:1^14.

THE NAUTILUS 11.3(2):56-63, 1999

Paee 56

Egg Capsules in the Genus Adelomelon (Caenogastropoda:
Volutidae) from the Atlantic Coast of South America

Pablo E. Penehaszadeh

Museo Argentino de Ciencias

Naturales-CONICET
A\'. Angel Gallardo 470
1405 Buenos ,\ires. ARGENTINA

and
INTECMAR

Uni\ersidad Simon Bolnar
1080 Caracas, VTINEZUELA

Patricia Miloslavich

Departamento de Estudios

Anibientales
Universidad Simon Boluar
1080 Caracas, \ENEZUELA

Paulo Marcio S. Costa

Departamento de Zoologia, Institute

de Biologia
Universidade Federal de Rio de

Janeiro
21941-570 Rio de Janeiro, BRAZIL

Mario Lasta

Institute) National de Investigacion y

DesarroUo Pesquero
Mar del Plata, ARGENTINA

ABSTRACT

In the past centurv, few additions ha\e been made to the earlv
descnptions of egg capsules of South American \olutes, and
unfortunately many of these earh' descriptions ha\e pro\en to
be incorrect. Herein, we describe the egg capsule of the largest
South American \oIute, Adelomelon beckii (Broderip, 18.36),
and redescribe the egg capsule of A. ancilla (Lightfoot, 1786),
two species that undergo direct de\elopinent. Adelomelon bec-
kii produces single, conspicuous, large, globose, and hemi-
spheric egg capsules that measure about .50 mm in basal di-
ameter and 35 mm in height; egg capsules of A. beckii are
found attached to scallop shells. The base of the egg capsule
is round with a narrow (3 mm) margin. The number of em-
bryos per egg capsule is 7-9. The internal yolume of the egg
capsule is 30—35 nil. All studied material was at pre-hatching
or hatching stages and no nurse eggs were present. Crawl-awa\"
juyeniles have shells measuring between 16.0-18.6 mm length.
The egg capsule of A. ancilla is o\al and flat, with a diameter
of 27—46 mm. Egg capsules are never coyered bv a calcareous
layer. They are generally attached to scallop shells. The number
of embr\'os per capsule is 2-8 and no nurse eggs are present
The eggs are about 65 |jim in diameter, the smallest among
South American xolutids. and are surrounded b\- a \er\- dense
fluid. The intemal volume of egg capsules is 2.5--4.0 ml. Hatch-
lings are released as crawling ju\eniles with shells measuring
11.7-12.7 mm. Abridged information on spawn patterns in
members of the Xolutidae from the southwest Atlantic, Carib-

bean, Subantarctic, Antarctic, eastern Atlantic, Indo-Pacific,
and Australia is pro\ided.

Kei/uord.s: Caenogastropods, \'olutidae, egg capsules, de\el-
opment, reproduction, geographic homogeneit)-, southern At-
lantic.

INTRODUCTION

The egg capsules of relatheK^ few South American \'ol-
utids have been described. Egg capsules of Adelomelon
hrasiliana (Lamarck, 1811) were first described by
dOrbignv (1846) and have been studied by De Mahieu
et al (1974) and Penehaszadeh and De Mahieu (1976).
It is the largest knowii unattached gastropod egg cap-
sule, with an intemal volume of up to 140 ml (Figures
1-2). Other species of South American volutes for which
egg capsules have been described are Zidona dufrcsnci
(Donovan, 1823), Odoulocijinhiola maficllanica (Gmelin,
1791) (see Penehaszadeh anil De Mahieu, 1976), Vohita
musica Linnaeus, 1758 (see Clench and Turner, 1970),
and Vohita virc.sccns Lightfoot, 1786 (see Handel, 1976).
Herein we describe the egg capsules of Adelomelon bec-
kii (Broderip, 18.36) ami redescribe the egg capsules of
Adelomelon ancilla (Lightoot, 1786), which have been

p. E. Penchaszadeh et al. 1999

Page 57

Figures 1-2. Atli'loinclon brasiliann (Lamarck. 1811) from
Mar del Plata, Argentina. Egg capsvile.s at pre-hatching stage
with live embrvos. Scale bar = 10 mm.

confused several times in the literature, mostly with
those of Odontoci/mbiola ma<iellanica.

A reevaluation of volutid egg capsule descriptions for
southwestern Atlantic volutids confirms that some earK
observations (e.g., Duhaut-Cilly in Lesson, 1840; Stre-
bel, 1906) are incorrect (Carcelles, 1944; Penchaszadeh
and De Mahieu, 1976). Therefore, this .studv aims to
properly describe the up to now erroneouslv identified
egg capsules of A. ancilla as well as to provide new in-

fcnination regarding the egg capsules, development, and
hatching mode of A. ancilla and A. bcckii.

MATERIAL AND METHODS

Egg capsules, hatchlings, and adult specimens of A. an-
cillo were collected by otter trawl. This tspe of gear is
regularlv used in the fisheries of the scallop Zt/<s_oclilamiis
pata<i(»iica (King and Broderip, l'S32) (for details on col-
lecting methods see Lasta and Iribame, 1997). Egg cap-
sules were invariably attached to empty scallop shells.

Egg capsules and snails were fi.xed in 7% formalin
buffered in seawater, and preserved in 70 % ethanol, and
examined under microscope.

RESULTS

Adelomelon beckii
(Figures 3-11, Table 1)

Description of egg cap.sule: The spawm consists of a
single egg capsule attached to hard substrate, usually the
external surface of an empty scallop shell. The large egg
capsule is globose and hemispherical. It measures about
50 mm basal diameter and 35 mm height. The base is
round, with a narrow margin (3 mm). No external cal-
careous layer is present. All embryos were at late de-
veloping stages. Egg capsules contain some albuminous
material and calciiun carbonate particles suspended
within the intracapsular li(juid. The numlier of embryos
per capsule is 7-9. Nine embrvos at early shell calcifi-
cation stage from a single egg capsule measure 13.5-18.5
mm in shell length. Shell size at hatching was 16.0-18.6
mm in length. The internal \-olmne of the egg capsule
is 30-35 ml, and the total volume of the pre-hatching
embryos 3.7 ml (n= 9). No nurse eggs were observed.
No exit plug or escape aperture is present in the capsule.
Hatching snails crawl away from the egg capsule as ju-
venile snails. Smooth surface, three whorls, and a lan-
ceolate calcarella characterize the juvenile shell at hatch-
ing (Table 1).

Material e.xamined: MACN 33SS4, 4 egg capsules
and 3 live-collected adults, off Cabo Frio, Rio de Janeiro
State, Brazil, 60-80 m, trawler, Feb. 1996; MACN
12900, protoconchs of juvenile and adult shells.

Addomclon inicilla
(Figures 12-21, Table 1).

Description of egg capsule: The egg capsule of A.
ancilla is oxal and Hat, with a basal minor axis measuring
25-44 mm and basal major axis measuring 27^6 mm.
The egg capsule wall lacks a calcareous layer. The num-
ber of eggs per capsule is 2-8. Nurse eggs are absent.
Eggs measure 60-71 |xm diameter and are embedded
in a very dense intracapsular liquid containing granulat-
ed particles. This gives an opaque appearance to the cap-
sule. The intracapsular liquid loses its opacity- during em-
bryonic development and growth. Egg capsule volume

Page 58

THE NAUTILUS, Vol. 113, No. 2

Figures 3-8. Adelomelon bcckii (Broderip, 1836) from off Cabo Frio, Brazil. 3. Lateral \iew of an egg cap.sule. 4. Emhrv'os
from a single egg capsule at a late stage of development. 5. Pre-hatching embryos. Scale bar = 10 mm. 6. Protoconch of an
adult shell from Mar del Plata, Argentina, MACN 12900. 7. Protoconch of an adult shell from Mar del Plata, Argentina (H.
Caldini Collection, Buenos Aires, Argentina). Scale bar = 6 mm. 8. Protoconch of a young specimen from off Cabo Frio, Brazil.
Scale bar = 6 mm.

is 2.5-4.0 ml. The total volume of the hatching juveniles
was as much as 1.1 ml measured for 8 emhrvos. Hatch-
ing takes place when juveniles crawl away from the egg
capsule. Juvenile shells at hatching measure 11.7-12.7
mm length, have a well-defined calcarella, and 2.5
whorls (Table 1).

Material examined: MACN .33882, 162 egg cap-
sules, and MACN 33883, 68 hatchlings, 180 miles off
mouth of Rio de la Plata, Buenos Aires Province, Ar-
gentina, trawler Erin, 90-105 m, Sep. -Nov. 1995, otter
trawl, egg capsules invariably attached to empty scallop
shells; MACN 16797, 2 egg capsules. Undine, 1925;
MACN unnumbered, 2 egg capsules, 39°30'S, 56°20'W,
Undine, 1925; MACN unnumbered, 1 egg capsule.

38°25-S, 56°20'W, Undine, 1925. The three last records
were examined and identified by Carcelles (1944) as A.
ancilla.

DISCUSSION

The first known description of the spawn of a species of
AdcJomclon was pnnided bv d'Orbignv (1846; p. 424) for
A. hrasiliana (Lamarck, 1811). He figured and recorded
the great abundance egg capsules in the Bay of San Bias,
in the Proxince of Buenos Aires, Argentina, during the
month of Fe])iaiar\-. He indicated that the diameter ot
the unattached egg capsule was 80-100 nmi, with 15-
20 embryos per egg capsule. Dall (1889, p. 312) reported

p. E. Penchaszadeh ct al, 1999

Page 59

Figures 9-11. Adelonielon beckii (Broderip, 1836) from off
Cabo Frio, Brazil. 9. Egg capsule. Scale bar = 20 mm. 10.
Recently hatched juvenile shells. Scale bar = 10 mm. 11. De-
tail of the protoconch of an adult specimen. Scale bar = 10
mm.

that the spawn of A. hrasiliana consisted of the most
extraordinarv" egg capsule in the Albatross collection.
Dall's description follows:

"This ovicapsule is oblate-spheroidal in shape, a view
from above giving a perfect circular outline, while from
the side the profile is a symmetrical oval. It is vellowish
in color but nearly transparent, thin, with a smooth, pol-
ished surface like that of wet gelatine, and possesses con-

siderable rigidity" (Dall, 1889). He also observed that:
"It was filled with a Hnid, probably not very different
from sea-water, and contained a single bubble of air,
which, by its hghtness remaining in the doTUe of the
capsule, just about counterbalanced the weight; so that,
without rising to the surface, the capsule would float in
the sea at a moderate depth". In view of our findings,
we suggest that this latter statement is incorrect. We
examined hundreds of egg capsules of A. briisiliaua im-
mediately after collection. Some contained air bubbles;
however, air infiltration clearly took place after removal
from seawater. If the capsule wall is cracked, liquid is
lost and iiir enters the egg capsule. Furthermore, in na-
ture, the egg capsules do not float or drift in mid-water.
The egg capsules of A. hrasiliana are a common by-catch
of shrimp trawlers that fish along the bottom, over which
the unattached egg capsules slowly drift. Unfortunatelv,
Dall's (1889) proposition about air inside the capsules
was accepted by Marche-Marchad (1968b), who hypoth-
esized about a possible South American origin for West
African volutes, with transport from east to west facili-
tated bv the alleged "pelagic transport" of egg capsules
of Adclomcloii. Bondarev (1995) has also adopted this
inaccurate assumption.

To add to the confusion surrounding egg capsules of
South American volutes, Clench and Turner (1964) re-
produced a figure by d'Orbigny (1846), which incorrectly
illustrates the egg capsule of Zidona dufresnei as being
one of A. hrasiliana. Furthermore, Knudsen (1993) er-
roneously stated that the egg capsule of A. hrasiliana
has a flattened base. De Mahieu et al. (1974) and Pen-
chaszadeh and De Mahieu (1976) measured and ana-
lyzed hundreds of egg capsules of A. hrasiliana from the
Mar del Plata region, off Argentina. The spherical egg
capsules had diameters of 40-80 mm, and an internal
volume of up to 140 ml. The number of embryos found
per egg capsule was 9-33 (Carcelles (1944) had reported
5-15 embrvos per capsule). The uncleaved egg mea-
sured 240 |xm diameter, and no nurse eggs were present.
Apparently, embiTOS fed on albumen and amino acids
present in the intracapsular fluid and internal wall of the
egg capsule.

In the past, several different egg capsules were be-
lieved to belong to A. ancilla. Carcelles (1944) errone-
ously assigned spherical, bottom-attached egg capsules
covered with a calcareous wall (characteristic of Odon-
tocijmbiola inagellanica, see figures 11-12) to A. ancilla
(MACN 16797). Penchaszadeh and De Mahieu (1976)
repeated the mistake made by Carcelles in his descrip-
tion. The egg capsules studied by Carcelles possibly be-
long to O. ina^cllanica. which lives in deeper waters off
Rio de la Plata. The egg capsules of O. mafiellanica stud-
ied bv Penchaszadeh and De Maliieu (1976) came from
Gulf San Jose and Gulf Nuevo, in northern Patagonia,
Argentina, 814 m depth. Geographic variation could be
relevant to the differences in numbers of hatchlings per
egg capsule found by Carcelles (n= 19-22) and Pen-
chaszadeh and De Mahieu (n = 6-10). This variation
could be related to differences in female size, geograph-

Page 60

THE NAUTILUS, Vol. 113, No. 2

Table 1. Summarv of the reproductive attributes of Af/c/o/iK'/oo bcrkii and Aclcloiucloit anciUa. Numbers in parentheses indicate
range.

Attribute

Adeloinelon beckii

Adelomelon anciUa

Shape and mode of attachment
of egg capsule

Diameter of egg capsule (nnn)

Globose
Attached to hard substrate

Base

48.8 ± 2.2
(46-51)
n = 4

Internal volume of egg capsule

30-35

(ml)

Number of enibr>'os per capsule

7-9

Egg diameter ((jlui)

Unknown!

Shell length at hatching (mm)

17.2 ± 1.0

(16.0-18.6)

n = 7

Hatching mode

Crawling juvenile (direct

development)

Number of shell whorls at

3

hatching

Height

34.3 ± 1.5

(32-35)

Flat oval
Attached to hard substrate

Base (ma.\.)

40.3 ± 3.4
(27-46)

Base (min.)

37.4 ± 4.2
(25-44)

n = 41
2.5-4.0

5.5 ± 1.8

(2-8)

n = 20

60-71

12.1 ± 0.4

(11.4-12.7)

n = 8

Crawling juvenile (direct

development)

2.5

Height
6.0 ± 1.3

(4-9)
n = 17

ic variation, or the e.\isteiice of a so far undetected cr^iJ-
tic species. However, the egg capsules of all material of
O. ma^ellanica examined consisted of an attached spher-
ical case covered by a white external calcareous la\er.
Adelomelon anciUa has an attached, \en' flat, expanded
egg capsule ^Aith no calcareous wall.

By re-evaluating old descriptions of South American
volutids, we are able now to recognize that the egg cap-
sules attributed bv Strebel (1906, figs. 10, 52, and 52a)
to Adelomelon anciUa (as Valuta ancilla), are indeed the
egg capsules of that species. However, Strebel also in-
correctly assigned egg capsules of several other species,
of different sizes and shapes, to this latter species. The
early anecdotal description made bv Captiiin Duhaut-
Cilly of the egg capsule of Odontociimbiola ma^cUanica
from the Magellan Strait (and repeated by Lesson
(1840)), "a round membrane, slightly convex, very sim-
ilar to a watch cover glass", could verv well represent
the egg capsule herein described for A. ancilla. The egg
capsules of A. ancilla are indeed more similar to the egg
capsules described for Zidona dufrcsnei than to those of

O. magellanica (see Penchaszadeh and De Mahieu,
1976).

The material assigned by Dall (1889) to O. nmgellan-
ica does not agree with the shape or size of the egg
capsules of this species as we know it. The capsules ex-
amined by Dall lacked the external calcareous layer. Ac-
cording to the shape, size, and number of embryos (2-
4), it is plausible that Dall (1944) had referred in this
case to the egg capsules of Zidona dufrcsnei (as de-
scribed by d'Orbign)', 1S46 and Penchaszadeh and De
Mahieu, 1976). The egg capsules that Arnaud (1978) as-
cribed to Hai'povoluta charcoti resembles those of Zi-
dona dufrcsnei, and not those of O. magellanica. as Ar-
naud suggested, probably following Dall's (1889) erro-
neous identification of the species.

Reproductive traits of South American volutes tor
which the reproduction has been studied (Table 2) sep-
arate veiv clearlv these species from the West African
and the Indo-Pacific members of the iamilv (Marche-
Marchad, 1968a, 1980; Smith, 1910; Tokioka,' 1962; Cot-
ton, 1937, 1944; Knudsen, 1993). Most South American

Figures 12-14. Aclcloinelon ancilla (Lightfoot, 1836). 12. Lateral \iew of an egg capsule detached from the substrate. Scale bar

= 10 mm. 1.3-14. Views through the transparent base of the egg capsule. Scale bar = 10 mm.

Figures 15-17. Adelomelon ancilla (Lightfoot, 1836) from off Rio de La Plata. Argentina. 15-16. Shell of a recently hatched

ju\enile. Scale bar = 5 mm. 17. Protoconch of an adult shell. Scale bar = 6 mm.

Figures 18-19. Volutid egg capsules. IS. (A) Capsules of Odonfoci/mbiola magellanica (Gmelin. 1791) and 18. (B) Adelomelon

ancilla (Lightfoot, 1836) attached to a valve of Zi/gochlami/s pataaonicn from oft Ri'o de la Plata, Argentina. Scale bar = 25 mm.

19. Detail of the capsule of O. magellanica. co\ered by a calcareous layer Scale bar = 20 mm.

p. E. Penchaszadeh ct ai. 1999

Page 61

Page 62

THE NAUTILUS, Vol. 113, No. 2

Table 2. Summan' oi information on the egg capsules in the family Volutidae.

Southwest Atlantic and

Caribbean, Subantarctic and

Antarctic

Eastern Atlantic

Indo- West Pacific and
Australia

Adelomelon, Odontocijmbiola. Zidona.
Harpovohita. Provocator, Vohiin imn-
sica )

Penchaszadeh and de Mahieu (1976),
Amaud and Van Mol (1979), Clench
& Turner (1964), present study

Spawn is a single egg capsule; generally
attached by flat base to hard sub-
strates; lens-shaped, hemispherical, or
spherical

Beh\'een 2-.3.3 eggs per capsule, \aria-
tions intra and interspecies

Unclea\ed egg diameter bet\A'een 60-
.300 |xm depending on species

No nurse eggs (°). Supplementary food
for the embrsos present as albumen
in intracapsular liquid and inner cap-
sule membrane

Hatching as crawling jm'eniles (direct
deyelopment)

Cijmbium

Marche-Marchad (1968a, 1980)

Spawn is a single spherical egg capsule
with a \'ery thin wall. It is incubated
on the foot of the female

Between 4-46 eggs per capsule, intra-
and interspecific variation occur

Diameter of the dexeloping egg arouTid
100 jxm

About 100,000 nurse eggs per egg cap-
sule; albumen present in intracapsular
hquid

Hatching as crawling juveniles or large
veliconch

Melo

Smith (1910), Tokioka (1962), Cotton
(1937, 1944), Knudsen (1993)

Spawn roughly cylindrical (pineapple-
shaped), consisting of about 10() indi-
\idual egg capsules spirally arranged,
central space present

Single embr\o per egg capsule

No information available

No nurse eggs reported. Apparently al-
bmnen present in intracapsular liquid

Hatching as crawling ju\'eniles (direct
de\elopment)

(°) If nurse eggs are confinned in Vohifa virescens. as stated by Bandel (1976). this would be an exception for this group

volutes have flat, hemispherical, or spherical attached
egg capsules (with the exception of A. hrasiliana. wliich
has a non-attached, spherical capsule), with 2-.3.3 eggs
per capsule, egg diameter usually of 90-.300 |jLm (60-71
|xm in A ancilla), and direct development. In addition
to volk, alliumin, amino acids, and carbohydrates are
consumed bv the embrvos. No nurse eggs are present.
South American members of the subfamilv Volutinae ap-
parendy represent an exception to the above (e.g.. Val-
uta virescens, Bandel, 1976).

When the reproductive strategies of our stiid\' animals
are compared to those described for volutids from West
Africa and the Indo-Pacific, several major differences
become apparent. The spawn in the genus Cijmhium
Roding, 1798 (West Africa), following the description by
Adans<.>n (1757) was at first thought to be viviparous. The
spawn of tiiis species was later described by Marche-
Marchad (ia68a, 1980) as consisting of one spherical egg
capsule with very thin walls, measuring .3^ cm diameter
and containing a very dense liquid (albumin). The de-
veloping eggs measured around 100 [Lxn and up to
100,(X)0 nurse eggs, measuring in average 1.50 (xm were
present. The egg capsules are brooded by the female in
a "brooding sac" located on the foot. Hatchhngs are re-
leased as veliconch larvae, their shell measuring 20-.50
mm (Marche-Marchad, 196Sa, 1980). The number of
developing embryos is 4-46, depending on the species
and the female size. The developmental time until
hatching may be as long as 5 months.

The Indo Pacific species Melo miltonis (Griffith and
Pidgeon, 1834), M. umbilicata Broderip in Sowerbv I,
1826, M. ducalc (Lamarck,18Il), and M. mclo (Ught-
foot, 1786) have been studied respectively bv Cotton
(1937, 1944) Knudsen (1993), Man and Middleton
(1946), Tokioka (1962), and Amio (1963). The spavm in
all these species consists of egg masses, roughlv cvfin-
drical, with a variable number of egg capsules spiraUv
arranged. The egg mass in these different species have
an empty, central space that is connected to the exterior
bv a nimiber of fontanels between the bands of egg cap-
sules. These egg masses are attached to hard substrata
by a base and may reach large sizes (e.g., 20 cm in M.
miltonis and 25 cm in M. ducalc). Onlv' a single embryo
develops inside each capsule. The egg diameter and type
of extra-emlirvonic nutrition have not been studied.
Hatchlings are released as crawhng juveniles.

ACKNOWLEDGMENTS

We greativ appreciate the manv valuable suggestions
made by Paula Mikkelsen (American Museum of Natu-
ral History) and John Wise (Houston Museum of Nat-
ural Sciences) of an earher version of the manuscript.
We also thank Ana K. Carbonini and Claudio Paredes
for laboratory assistance. This work was partly supported
by a grant from Decanato de Investigaciones v' Desar-
roUo, Universidad Simon Boh'var, Fundacion Antorchas,
Argentina, and CONICET (PIA-6139)

p. E. Penchaszadeh et al., 1999

Page 63

Figures 20-21. Adelomelon ane'dla (Lightfoot, 1836) from
off Rio de la Plata, ArgenHiia. 20. Dorsal (A) and (B) lateral
\iews of egg capsule. Scale bar = 10 mm. 21. Shell of a re-
cently hatched juvenile. Scale bar = 5 nun.

LITERATURE CITED

Adanson, M. 1757. Histoire Naturelle du Senegal. Coquillages.
Paris, 275 pp.

Allan, J. and T. C. Middleton. 1946. Obsen'ation on the egg
case and gonad of the baler shell Melo umbilicata Brod.
Victorian Naturalist, Melbourne 62:172-177,

Amio, M. 1963. A comparative embryology of marine gastro-
pods, with ecological considerations. Journal of the Shi-
monoseki University of Fisheries 12:229-.358.

Amaud, P. M. 1978. Obser\ations ecologiques et biologiques
sur le V'olutidae antarctique Hiiiyovohita cliarcoti (Lamy,
1910) (Gastropoda Prosobranchia). Haliotis 7:44—46.

Amaud, P. M. and J. -J. Van Mol. 1979. Anatomy, ecologv' and
distribution of the Volutidae and Volutomitridae (Gastro-
poda: Prosobranchia) of the southern Indian Ocean. The
Veliger 22:19-31.

Bandel, K. 1976. Spawning, development and ecology of some
higher Neogastropoda from the Caribbean Sea of Colom-
bia. The Veliger 19:176-193.

Bondarev, 1. P. 1995. Classificazione filogenetica di Volntidae
dell'Austraha (Mollusca: Gastropoda) con la descrizione di

un nuvo sottogenero e di ima nuova sottofamiglia. Natur-
ama 27(276):25-39.

Carcelles, A. 1944. Catalogo de los moluscos marinos de
Puerto Quequen. Revista del Museo de la Plata 3:233-
309, 15 pis.

Clench, W. J. and R. D. Turner 1964. The subfamilies Voluti-
nae, Zidoninae, Odontocsmbiolinae and Calliotectinae in
the Western Atlantic. Johnsonia 4:129-180.

Clench, W. J. and R. D. Turner 1970. The familv Volutidae in
the Western Atlantic, johnsonia 4:369-271.

Cotton, B. C. 1937. Eggs and Egg cases of some Southern
Australian Mollusca. Records of the Southern Australian
Museum 6:101-103.

Cotton, B. C. 1944. The egg capsule of the Southern Australian
Baler Shell Melo miltonis Gray. Records of the Southern
Australian Museum 8:143-144.

Dall, W. H. 1889. Scientific results of explorations bv U.S. Fish
Commission Steamer Albatross. 7. PreliminaiT Report on
the collection oi Mollusca and Brachiopoda obtained in
1887-1888. Proceedings of the United States National
Museum 12(773):219-362.

De Mahieu. G. C, P E. Penchaszadeh and A. Casal. 1974.
Algunos aspectos de las \'ariaciones de proteinas y ami-
noacidos libres totales del liquido intracapsular en relacion
al desarrollo embrionario en Adelomelon brasiliana (La-
marck, 1811). Cahiers de Biologic Marine 15:215-227.

d'Orbignv. A. 1846. Voyage dans I'Amerique Meridionale;
Mollusques, V. Paris, 758 pp.

Knudsen, J. 1993. Redescription of the egg mass of Melo mil-
tonis (Griffith and Pidgeon, 1834) (Mollusca, Prosobran-
chia, Volutidae). Joumal of the Malacological Societv of
Australia 14:107-112.

Lasta, M. L. and O. O. Iribanie, 1997. Southwestern Atlantic
scallop (Zi/gochlnmijs patagonica) fishery: assessment of
gear efficiency through a depletion experiment. Journal of
Shellfish Research 16:59-62.

Lesson, R. P. 1840. Observations Zoologiques. (Notice sur la
maniere dont se reproduit une espece de Volute du de-
troit de Magellan). Re\aie de Zoologie 1, Societe Cinder,
Paris: 167-169

Marche-Marchad, 1. 1968a. Un nouveau mode de developpe-
nient intracapsulaire chez les Mollusques Prosobranches
Neogastropodes: I'incubation intrapediose des Ct/niba
(Volutidae). Comptes Rendues de I'Academie des Scienc-
es, Paris 266:706-708.

Marche-Marchad, I 1968b. Remarques sur le de\eloppement
chez les Ci/mbti (Prosobranches Volutides) et I'Inpothese
de leur origine sud-americaine. Bulletin de I'lnstitut F"ran-
9ais d'Afrique Noire, T XXX, ser. A. 3:1028-1037.

Marche-Marchad, I. 1980. Sur la strategic de la reproduction
chez le genre Ctjmbiiim Roding 1798 (Gastropoda, Volu-
tidae). HalioHs 10:94.

Penchaszadeh, P. E. and G. C. De Mahieu, 1976. Reproduc-
cion de gasteropodos prosobranquios del Atlantico Sur-
occidental. Volutidae. Physis A, 35(91 ):145-153.

Smith, E. A. 1910. Note on the egg-capsules of Melo. Pro-
ceedings of the Malacological Society of London 9:4— .5.

Strebel, H. 1906, Beitrage zur Kenntnis der Molluskenfauna
der Magellan Pnnanz 4. Zoologische Jalirbucher (Syste-
matik):91-174, 7 pis.

Tokioka, T 1962. Record of a giant mass of Melo ducale (La-
marck) from the Arafura Sea, Publications of the Seto Ma-
rine Biological Laboraton- 10:21-26.

THE NAUTILUS 113(2):64-70, 1999

Page 64

Reassignment of Henry a morrisoni Bartsch, 1947 from the
Family Aclididae to the EbaUdae (Gastropoda: Heterobranchia

John B. Wise

Department of Malacologv'
Houston Museum of Natural Science
One Hermann Circle Drive
Houston, TX 77030-1799 USA
jwise@hmns.org

ABSTRACT

Henn/a inorrisoni Bartsch, 1947, originally assigned bv Bartsch
(1947) to the famil>- Aclididae, is herein placed in the family
Ebaiidae. Henrya nwrrisoni is a cryptic, minute gastropod pos-
sessing a complex jaw apparatus. This apparatus, consisting of
a single pair of elongate hooks, hook elements, and muscular
base, is characteristic of the family Ebaiidae (e.g., Ebola niti-
dissima). The infaunal H inorrisoni is a simultaneous her-
maphrodite, with sperm nioqihology typical of basal hetero-
branch gastropods. The tamiK' Ebaiidae is a sister taxon of the
Pxraniidellidae, superfamiK P\ raniidelloidea.

Key words: Anatomy, fimctional moqihology, systematics, Pyr-
amidelloidea, Heterostropha

INTRODUCTION

In 1994, Waren assigned the extant genera Ebala Grav,
1847, Hcnn/a Bartsch, 1947, Murchisonella Mcirch,
1S75, and the fossil genus Donaldina Knight, 19.3.3 to his
then newly named family Ebaiidae. Pre\iously, these
taxa were placed in AcUdidae and Pyramidellidae solely
on the basis of shell characters. Waren (1994) demon-
strated that, unlike pvramidellids, which possess a very
diagnostic buccal st\let (Maas, 1965; Wise, 1996), these
snails possess a comple.x jaw apparatus. Recent Euro-
pean species of Ebaiidae live in shallow-water lagoons,
estuaries, and seagrass beds (Waren, 1994).

Our understanding of ebahd hie histon- is limited to
a single species. Rasmussen (1944) described and illus-
trated the egg capsules, embryonic development, and
vehger larval stage of E. nitidi.ssiina (Montagu, 1803),
from specimens that he collected at Isefjord (Sealand),
Denmark in 5-7 m (see Rasmussen for list of other Eu-
ropean locahties). This species, which is usually found in
association with Zostcra beds, is very abundant during
its spawiiing season (May-June). Thorson (1946) de-
scribed the sinistral, planorbiform larval shells of E. ni-
tidissima, as well as the external moipholog)' of its ve-
liger larva. Healy (1993) examined the sperm morphol-
ogy of E niiidissima and determined that its nucleus is

elongate, strongly keeled, and almost completely pene-
trated by the axoneme/coarse fiber complex. In contrast,
members of the Pyramidellidae have a short nucleus,
with one (jr more strong helical keels and a shallow basal
invagination, which cont;iins only the centriolar deriva-
tive and proximal portion of the axoneme/coarse fiber
complex. Huber (1993) examined the neivous system of
E. nitidi.ssiina (then assigned to the familv Pyramidelli-
dae) and determined that it is significantK- different from
the other pyramideUids he investigated. The eyes of E.
nitidissima, unfike those of pvramideUids, are attached
to the cerebral ganglion by the optic nerves. These and
other differences in their respective ner\'ous systems
(Hiiber, 1993) and sperm moiphologs' (Healv, 1993) fur-
ther suppcjrt the separation of Ebahdae and Pyramidel-
lidae.

J. P. E. Morrison collected specimens of Henrya mor-
risoni from the drift line off a "mangrove swamp" on
Shell Key, off St. Petersburg, Florida in April, 1936.
Subsequently, this species was named and assigned to
the Aclichdae by Bartsch in 1947. Bartsch described the
shell of the new species as very minute, pupoid, very
slender, and milky white, with its nucleus consisting of
a single, strongly rounded, hyahne whorl forming a very
blunt apex. Postnuclear whorls were characterized as
high between summit and suture, strongly rounded, and
marked h\ conspicuous growth lines. See discussion on
Bartsch's reasons for assigning this taxon to this family
under Taxonomic Remarks below.

This study was undertaken to provide insight into the
moq^hology and systematic position of Henrya morri-

MATERIAL AND METHODS

The author (see HMNS material below) collected spec-
imens ot Hcnnja morrisoni. Samples of the top 5-15 cm
of mostly mud substratum were sieved through a 0.5
mm sieve, rinsed with seawater, and the remaining frac-
ticMi sorted under a dissecting microscope. Snails were
kept ah\e in small bowls of aerated seawater.

J. B. Wise, 1999, 1999

Page 65

Figures 1-8. Scanning electron microphotogruphs of shell and operculum of Hciin/n morrisoni. 1. Apertural \new. 2. Dorsal
\iew. Scale bars = 250 p-m. 3. Enlargement of aperture. Scale bar = 100 (jLm. 4. Apical view of protoconch. Scale bar = 30 |xm
5, 6. Lateral Niews. Scale bars = 30um. 7, 8. Operculum. 7. Attachment side. 8. Unattached side. Scale bars = 100 (jlui.

Page 66

THE NAUTILUS, Vol. 113, No. 2

Figure 9. Living Henrtja morrisoni. Scale bar = 300 |xni.
att = anterior alimentan- tract, ct = ceplialic tentacle, me =
mentum.

Morphology

Light Microscopy

a) Photography: Living snails were photographed with a
Pentax 35 mm camera mounted on a Zeiss Tessavar
dissecting microscope.

b) Dissections: Snails were dissected whole, once their
shells were cracked with forceps and shell fragments
removed. Whole animals and/or their ahmentar)' and
reproduction systems (after excision) were stained
with toluidine blue. Line drawings were produced
vmder a dissecting micnjscope equipped with draw-
ing tube.

Scanning Electron Microscopy

Opercula and shells were cleaned bv sonication, air-
dried, coated with gold-palladium, and examined under
Selectron 250 or JEOL JSM-6100 scanning electron mi-
croscopes operating at .5-10 KEV.

Museum material examinech USNM 466297, 5 shells,
[not 11, as stated by Bartsch (1947)], type locality;
USNM 466225, 1 specimen, Morrison station 14, upper
line of drift along mangrove swamp. Shell Key St. Pe-
tersburg, Florida; IIMNS 46019, 2 specimens. May,
1994; HMNS 46020,1 specimen, Feb. 1996; both lots
from junction of Pine Island and Bayou roads, off Flor-

ida State Highway 550, near Brooksville, Florida, sub-
tidal, marsh area, J. Wise coll.

SYSTEMATICS

Subclass Heterobranchia Grav, 1840

Superiamily Pyramidelloidea Gray, 1840
FamiK- Ebalidae Waren, 1994

Taxonomic remarks: Van Aartsen (1995) has argued
that the names Ebala Gray, 1847 and Ebalidae Waren,
1994 are una\'ailable and/or in\alid. He contended that
the names Anisocijcla Monterosato, 1880 and Anisocy-
chdae van Aartsen, 1995 should be used instead. Van
Aartsen also stated that "even if one considers Ebola
Grav, 1847 (not of Leach) a new genus as of November,
1847 [Gray, 1847b], it is still a junior honionvm of Ebola
Leach in Gray, October, 1847 [Gray 1847a], and there-
fore cannot be used." This however is incorrect as the
name was introduced in sviionvinv in October 1847 and
unavailable names do not enter into homonvmy (ICZN,
1985, Article 54). As outhned by Waren (1994), the ge-
nus Ebola should date from Gray (1847b) with the t\^e
species as Turbo nitidissima Montagu, 1803. Mr P. K.
Tubbs, Secretarv^ of the International Commission on
Zoological Nomenclature (ICZN) (Waren, pers. comm.),
confirmed this conclusion. Waren (1994) further dem-
onstrated that AnisocijcJa Monterosato, 1880 is a re-
placement name for Aciculina Deshayes, 1862.

Recentlw Le Renard (1998) discussed the \arious uses
of the names Ebola and Anisocijcla and concluded that
the later should be used, following van Aartsens's inter-
pretation of Grav (1847b). It is beyond the intended
scope of this paper to delve into the problems arising
from Cray's work. It should be sufficient to state that I
consider both Ebalo and Ebalidae to be valid names as
demonstrated by Waren (1994). As the Secretary of the
ICZN has endorsed this position, anvone in disagree-
ment should petition the ICZN for adoption of his or
her \iews.

In addition, various ebahd species (e.g., Ebala nitidis-
sima) have been placed in the family PvTamidellidae and
tvpically assigned to the genera Euliniella or Anisocijcla
(Rasmussen, 1944; Thorson, 1946; Wilke and van Aart-
sen, 1998). The status and systematic position of Euli-
mella has not been examined as part of this study.

Hcnri/a Bartsch, 1947

Tjpe species; Hcnrifa lienn/i Bartsch, 1947, by origi-
nal designation.

Taxonomic remarks: Bartsch (1947) erected the genus
Henn/a to which he assigned the new species hcnnji,
^oldmani, and morrisoni (respective type localities: San
Salvador, Bahama Islands; Progresso, Yucatan, Mexico;
and Florida [see above]). Essentially, he separated these
taxa using interspecific differences in shell diameter
Bartsch placed them in the family Aclididae because of

J. B. Wise, 1999, 1999

Paee 67

13

Figures 10-15. Hcnnjn morrisoni. 10. External anatomy of H. morrisoni. Scale bar = 300 |j.mi. 11. Right mantle edge and
siphon. Scale bar = 125 (Jim. 12. Dorsal \iew of mantle edge and lobe. Scale bar = 125 (xm. 13. Diagram of alimentarv' tract.
Scale bar = 300 |xm. 14, 15. Jaw apparatus. 14. Schematic view. 15. Single hook. Scale bars = 10 (j-m. aat = anterior alimentan,'
tract, at = alimentarv tract, ct = cephalic tentacle, dgl = digestive gland, hbgl = hvpobranchial gland, hk = hook, int = intestine,
j = jaw, je = jaw element, ma = mantle, mae = mantle edge, nial = mantle edge lobe, nuis = mantle edge/siphon, me = mentum,
mub = nnisciilar base, ne = neck, s = stomach. \m = visceral mass.

their shell morphology and small size (generally < 2
mm).

Hciin/a inorrisdiii Bartsch, 1947

Hc)in/a niDnisoiii Bartsch. 1947:14-15, pi, 3, fig, 1. (Holotvpe:
USNM 57.36.36 (lost), t\pe localitv", ]. P, E, Morrison's' sta-
tion 15, in drift at etlge ot a mangrove swamp. Shell Is-
land, off St, Petersburg, Florida); Abbott, 1974:1.33,

Description: Shell and upvindinn (Fifiuivs IS): Shell

elongate, transparent, thin and fragile, to 1.25 mm in
length, with 4-.5 teleoconch whorls (figures 1, 2). Teleo-
conch whorls convex, with moderately shallow sutures
and numerous, fine axial growth lines. Bodv whorl .50 to
.5.5% of shell length. UmLilicus absent. Aperture elon-
gate-ovate to round (figure .3). Sinistral, smooth proto-
conch, with lower portion sunken in first teleoct)nch
whorl (figures 4-6). Operculum horny, translucent oper-
culum, ovate and closing aperture (figures 7, 8).
Head-foot and visceral mass ( F/g(/rp.s 9, 10): Head grey
to golden brown, with anterior remon of alimentarv tract

Pase 68

THE NAUTILUS, Vol. 113, No. 2

16

17

-atp

Figures 16, 17. Hennja morrisoni. 16. Reproductive tract. Scale bar = lOOjjLiii. 17. Diagram of penis. Scale bar = 10 [Jiin. atp
= anterior tip of penis, blp = bulbous base of penis, dgl = digestive gland, fgl = female glands, gd = gonoduct, hd = hermaph-
roditic duct, sr = seminal receptacle, sv = seminal \'esicle, gd = gonoduct, mc = mantle cavity, o = egg, o\t = o\otestis, pe =
penis, vm = visceral mass.

visible by transparency as \'er\' di.stinctive, medial black
stripe extending from area just behind mentum notch
onto neck (figures 9, 10, aat). Flattened wing-hke ten-
tacles extend 90 degrees on either side of head (figure
9, ct). Tentacles not joined. Tentacular pads lacking.
Mentum deeply bifid and bilobed (figures 9, 10, me).
Tentacles, mentum, and lateral portions of head and toot
darkly pigmented. Eyes small, subepithefial, black, wide-
ly spaced, on either side of medial head stripe, with well-
developed lenses (figures 9, 10). Mouth opening medi-
ally within bilobed mentum. Propodium not bifid, an-
tero-laterally rounded. In its region behind propodium,
foot narrows initially to then widen, tapering to blunt
tip. Both mentum and propodium highly ciliated, with
particularly long propodial cilia. Mentum stretched on
substratum in region anterior to propodium in crawfing
animal. No attachment thread or pedal gland obsened.
Visceral mass (figure 10, vm) divided into digestive
gland, whicli consists of redchsh-brown, grey, or black
cells and greenish-yellow reproductive tract (mostly ovo-
testds).

Pallial cavity (Figure 10, 11, 12): Pallial cavity, long,
spacious, tapering in posterior direction, with short si-

phon (figure 11, mas). Dorsal surface of mantle darkly
pigmented. Mantle edge (figure 12, mae) thick, finely
crenulated and wide, wdth large, elongate lobe on right
forming 45°-90° v\ith mande edge (figures 10, 11, 12,
mal). Ventral and dorsal ciliated strips located on right
side of mantle cavitv, joining in posterior region. Ventral
ciliated strip originates on mantle lobe. Large, elongate,
yellow hypobranchial gland located on left side of mantle
roof. Hypobranchial gland (figure 10, hbgl) composed of
clear matrix in which large and small cells containing
yellow substance are suspended. When snail is dis-
turbed, cells release thick, pale-yellow exudate. Heart,
located at right posterior of mantle cavity, consisting of
single auricle and single ventricle. Gill absent. Kidney
not found.

Aliiiicntanj tract (Figures 13-15): Gut extending in
posterior direction as long, straight tube (figure 13, at)
from beUveen mentum lobes to stomach, which is lo-
cated within visceral mass (figure 13, s). Jaw (figures 13,
14, j) located inside buccal opening and consisting of one
pair of large hooks, each with an element attached to a
basal plate (figure 14, 15, je). Hooks inserted into mus-
cular basal plates, which provide support and operation

J, B. Wise, 1999, 1999

Page 69

of jaw. It is liere suggested that jaw is exerted on short
snout through buccal aperture during feechng. Radula
absent. Anterior portion ot gut green, posterior section
\ello\\ish-orange. Anterior gut passing through nerve
ring, within head. Small stomach anterior to junctures
of digestive gland (figure 13, dgl) and intestine (figure
13, int). Intestine forming 90° with stomach, turning and
extenthng in anterior direction parallel to gut. Rectum
and anus not observed, but anus probably opening on
right posterior portion of mantle ca\it\', on mantle floor.
Digesti\e gland packed with mostlv dark cells (a com-
bination of reil and blue cells), extending to tip of \is-
ceral mass.

Reproductive system {Figures 16, 17): Species exliibits
simultaneous hermaphroditism. Elongated ovotestis (fig-
ure 16, ovt) located within visceral mass on columellar
side, consisting of lobes where both eggs (figure 16, o)
and sperm are produced. At lower portion of ovotestis
is a short duct e.xtending anteriorly to elongate and ta-
pering seminal vesicle (figure 16, sv). Seminal vesicle
King parallel to ovotestis, joining hermaphroditic duct
(figure 16, hd), which extends from visceral mass (figure
16, vin) into mantle cavity (figure 16, mc) to gonoduct
(figure 16, gd) and female glands (figure 16, fgl). Narrow
gonoduct ends on neck on right side of head. Small,
round seminal receptacle (figure 16, sr) joins gonoduct
through short duct in region anterior to widening of gon-
oduct and female glands. Penis (figure 17, pe) with ta-
pered, round anterior tip (figure 17, atp) and long neck
that extends in posterior direction to form bulbous base
(figure 17, blp). Protnisible penis extends from body
through medial opening ventral to mentum. Vas defer-
ens not observed. Egg and larval development unknown.

CONCLUSIONS

The infaunal and diminutive snail Henri/a morrisotii in-
habits the tidal marshes of southern Florida. It was first
described from its shell onlv by Bartsch (1947) who as-
signed the species to the family Aclididae. The present
findings, when compared with those of Waren (1994;
pers. comm.) and Rasmussen (1944), indicates that this
species should be reassigned to the familv' Ebalidae War-
en, 1994. This species, like those described by Waren
(1994), have a diagnostic jaw apparatus. Its jaw, consist-
ing of a single pair of hooks, supporting elements, and
a muscular base, is similar to that of E. ititidissiiiia.
Moreover, the external anatomy (e.g., tentacle shape,
mentum configuration, anterior alimentary tract delin-
eated by black stripe, etc.) of H. morhsoni closelv re-
sembles that of E. nitidissima (Rasmussen, 1944). Con-
sequently, the allocation of this taxon to the familv Ebal-
idae is unequivocal.

ACKNOWLEDGMENTS

I would like to thank Dr. Robert Bullock for calling my
attention to these snails and for disclosing their location.

I am grateful to Dr. John Hicks and Mr. Jim Barrish of
the Texas Children's Hospital, Houston. Texas for SEM
time and darkroom expertise. I also thank Dr A. Waren
for providing his thoughts on this matter and his unpub-
lished data. Mr Richard Petit provided very useful in-
sight into the nomenclatural confusion concerning ge-
neric and familial names. The critical comments of the
anonvmous reviewers improved this manuscript. Lastlv,
none of this would have been possible without Dr. M.
Rice and her staff at the Smithsonian Marine Station at
Fort Pierce, Florida This paper is Smithsonian Marine
Station Contribution no. 477.

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Aartseii, ]. J. van. 1995. Anisoci/cla Monterosato. 1880 or Ebala
Leach in Gray, 1847; That is the question. Bollettino Ma-
lacologico .3:6.5-68.

Bartsch, P. 1947. A monognipli of tlie west Atlantic niollusks
of the family Acliditlue. Smithsonian Miscellaneons Col-
lections 106(20): 1-29, 6 pis.

Deshayes, G. P. 1862 (1859-65). Description des animaux sans
vertebres, decouvertes dans le Bassin de Paris. II. Bailli-
ere, Paris. 968 pp.

Grav, I- E. 1847a. The classification of Britisli Mollusca. Annals
and Magazine of Natural Historv' 20:267-27.3.

Grav, J. E. 1847b. A hst of the genera of recent mollusca, their
sviionyma and types. Proceedings of the Zoological Soci-
ety of London for 1847:129-219.

Healv, J. M. 1993. Comparative sperm ultrastnicture in basal
heterobranch gastropods (\'alvatoidea. Architectonicoidea,
Rissoelloidea, OmalogvToidea, Pvramidelloidea (Mollus-
ca). Zoologica Scripta 22:263-277.

Huber, G. 1993. On the central nervous system of marine Het-
erobranchia (Gastropoda). Journal of Mollnscan Studies
59:381^20.

[ICZN] International Commission on Zoological Nomencla-
ture. 1985. International code of zoological nomenclature
Third edition. International Tnist for Zoological Nomen-
clature, London, .3.38 pp.

Knight, J. B. 19.33. The gastropods of St. Louis, Missouri,
Pennsylvanian outlier V. The Trocho-Tubinidae. Journal
of Paleontology 7:30-58.

Le Renard, J. 1998. Anisocijcia Monterosato 1880 (Mollusca:
Gastropoda), un genre victime d'un artifice nomenclatural
analvse: La sviionviiiisation per restriction heterotvpique.
Cossmamiiana 5: 109-1 14.

Maas, D. 196.5. Anatomische nnd histolo0sche Untersucliun-
gen am Mundapparat der Pyramidelliden. Zeitschritt tiir
Morphologic und Okologie der Tiere 54:.566-642.

Montagu, G. 1803. Testacea Britannica — J.S. Hollis, Rom.sey.
xxvii -I- 610 pp.

Monterosato, A. Di. 1880. Conchiglie della zona tlegli abissi.
Bolletino de la Societa Malacologica Italiana 6:.50-84.

Morch, O. A. L. 1875. Sviiopsis Molluscornm marinomm In-
diarum occidentaliuni. Malakozoologische Blatter 22:
I42-1S4.

Rasmussen, E. 1944. Faunistic and biological notes on marine
invertebrates I. The eggs and larvae of Brachystomiii ri.s-
soides (Hani.), Eulimella nitidissima (Mont.), Retusa tnin-
catula (Brug.) and Emhletonia pallidn (Alder and Han-
cock), (Gastropoda marina). N'idenskablige Meddelelser

Page 70

THE NAUTILUS, Vol. 113, No. 2

fra Dansk Natuiiiistorisk Foreiiing I Kobenhaxii 107:207-
233.

Thorson, G. 1946. Reproduction and lanal development of
Danish marine bottom invertebrates, with special refer-
ence to the planctonic lai-vae in the sound (Oresund).
Meddelelser fra Komniissionen for Danmarks Fiskeri-Og
Ha\imders0gelser, Serie Plancton 4:1-523.

W'aren, A. 1994. Systematic position and vahdit\- of Efo«/n Gray,

1847 (Ebahdae Fam N., P^Tamidelloidea, Heterobran-

chia). BoUetino Malacologico 30:20.3-210.
Wilke, T. and ]. J. van Aartsen. 199S. The family Pyramidelhdae

(Heterostropha, Gastropoda) in the Black Sea. Ba.steria

62:7-24.
Wise, J. B. 1996. Moqihologv and phvlogenetic relationships

of certain pvTamidellid taxa (Heterobranchia). Malacologia

37:443-511'.

THE NAUTILUS 113(2):71-72, 1999

Page 71

Book Review

The Freshwater Mussels of Tennessee

hij Paul \V Ftiniuilcc and Aiihiir E. Bo^an. 1998. Uni-
versitv of Tennessee Press, Knowalle. xii, 1-32S, 133 pis,
130 maps.

In the arena (if s\steniatic re\isions and fauna! re-
views, freshwater nialacolog)' has always been behind the
marine malacological community for several decades or
more. The pioneerina; studies of Ortmann in Pennsyl-
vania were fitfulK followed bv similar works on Missouri,
Indiana, and a paltw tew other states. It is only very
recently that monographic-scope works on freshwater
mussels have finally come into their own. This book
brings freshwater malacologv into the same league as
marine malacologv.

One of the quirks of freshwater malacology is its in-
sistence to cover not a meaningful biogeographic region
such as the Holarctic, or the Ohio River basin, but the
meaningless arbitran^ unit of The State, no matter
whether a large state (Texas) or a small one (Vermont)
is concerned. This cycle seems unalterable, as nearly all
of the most recent works have been at the state level,
and nearh' all planned ones will be as well. Parmalee
and Bogan's book is no exception; it deals with Tennes-
see. But here it fares better than most because of the
nature of the state's boundaries. Within Tennessee is a
portion of the Mississippi Ri\er, with all of its big river
and bayou taxa. More irnportantlv Tennessee is located
predominantly in the Tennessee and Cumberland River
basins — those wonderfullv diverse svstems that have el-
ements of the Ohio River fauna plus numerous endem-
ics. Finallv, tucked in the southeastern comer is the
Conasauga River, a lone portion of the Mobile River sys-
tem, with its very unique fauna. As such, Tennessee has
(or had) within its borders nearly half of all of the rec-
ognized unionoid ta.xa in North America. For this reason,
this book is useful far outside of the boundaries of the
State of Tennessee.

There is a lot to like about this book. As with many
malacological works that emerged on the market in re-
cent times, it has a colorful, appealing format catering
to more than just professional malacologists. Intrcjduc-
tor\' chapters range from adequate to excellent. The sec-
tion on ecology is rather brief, and is concerned mainly
with causes for mussel decline rather than the ecological
requirements of the animals. However, the section on
aboriginal exploitation is excellent and represents an as-
pect of these animals often neglected in other works. A
concise history of the classification of freshwater mussels
provides a useful basis to understanding the svsteniatics
of the group. Readers ahead) familiar with Dall,
Bartsch, and Pilsbry are introduced to those other giants
of American malacolog\-: Ortmann. Call, and Lea. The

section on diversity is an excellent backdrop to the book,
bringing together information on the hydrologv', topog-
raphy, and biogeographv of the Tennessee river svstems.
Most of the book is de\'oted to species accounts, which
are mercifully arranged in alphabetical order of scientific
names. Each species is represented by a small color plate
that depicts both the inside and outside of the shell and
that illustrates sexual dimorphism as well. Each species
has its own dot distrilnition map with a smaller map
indicating the total range of the species. Plates and maps
are conxeniently arranged with the text to each species,
rather than gathered in the back. The authors have la-
boriousK' assembled priman' .s\iionymies that are backed
by an extensive literature cited section. For each species,
sections are included for distribution, description of the
shell, life histoiv and ecolog\, and its legal status. Intro-
duced Corbicida and Drcisscna are covered as well.

For its size, errors are lew. Perhaps mv main criticism
is with their use of type localities. For example, under
ArcUlens confrago.sii.s (Sav, 1829) they give "Bavou
Teche. " But Says original localities were the Wabash
River (later emended bv Say to the "Fox River") and
"New Orleans" (emended to "Bayou Teche"). By choos-
ing Bayou Teche as "type locality," Parmalee and Bogan
have restricted the tvjje localits' — but did they mean to
do so? On sexeral occasions the authors ignore the type
designations (and therefore the type locality designa-
tions) made h\ other workers. For Pti/chobranchiis fa.s-
ciolaris (Rafinesque, 1820), the authors give the t\pe lo-
cality' as the Muskingum Ri\er, although the lectotype
selected by Johnson and Baker (1973) is from the Ken-
tuck"v River. Whether the authors have reason to doubt
or ignore these designations is not stated.

Although the authors describe each species, what is
missing is a diagnosis that would allow separation of a
species from its lookalikes. For example, several closely
related taxa of Plciirobcma that have never been ade-
quately differentiated are fotmd in the Conasauga River.
Yet the authors only connuent that "it is nearly impos-
sible to identify and separate many individuals." This is
no help to the reader. The authors could ha\e greatly
increased the worth of this book bv gi\ing the reader
some simple guidelines lor differentiating each species.

The svnonvniies are invaluable, but occasionalK' lack
justification. Uniu nobilis Conrad, 1854, has recently
been resurrected as a valid species. \et Parmalee and
Bogan synonvniize it [in part] with Quadnda apicidata
(Say, 1829). In view of the renewed intere.st in this spe-
cies, some mention should have been made as to why
they felt this course was correct. Similar unsupported
.synonymies are found under Ellipfio arctata (Conrad,
1834) and Lampsdis altilis (Conrad, 1834), which prob-
abl\ represent species complexes. Under Vdlosa iris
(Lea, 1829) thev sviionvmize without comment Unio si-

Page

THE NAUTILUS, Vol. 113, No. 2

mils Lea, 1838, which was recently resurrected and
placed in Venustaconcha by Gordon (1995). Other S)n-
onymies may leave the reader confused. Under Lamp-
silis ahnipta (Sav, 1831) we find the name Unio orhi-
ctilatus Hildreth, 1S28, which would seem to ha\'e pri-
ority over Unio abniptits Say, 1831. But no explanation
is given why Hildreth s name is not used (Hildreths
specimen was actually Obovaria subrottinda (Rafin-
esque, 1820), and the svnonvmv should read "Unio or-
biculatiis ot authors, not Hildreth, 1828").

Despite these rather specific criticisms, this book rep-
resents the state-of-the-art in freshwater malacological
guides. Government agencies and consenation groups
will embrace the maps and plates, and malacologists will
continue to argue over the synonymies. This promises to
be a landmark book in its field.

LITERATURE CITED

Gordon. M. E. 1995. Veiuistaconchii ■iiina (Lea), an overlooked
freshwater mussel (BivaKia: Unionoidea) from the Cum-
berland River Basin of Central Tennessee. The Nautilus
108(3):55-60.

Johnson, R. L and H. B. Baker 1973. The hpes of Unionacea
(Mollusca; Bi\'alvia) in the Academy of Natural Sciences
of Philadelphia. Proceedings of the Academy of Natural
Sciences of Philadelphia 125:14.5-186.

G. Thomas Watters

Ohio Biological Survey
Ohio State University
1315 Kinnear Road
Columbus, OH 4.3212 USA
g\vatters@postbox. acs.ohio-state.edu

Notices

OPINIONS PUBLISHED IN THE BULLETIN OF ZOOLOGICAL NOMENCLATURE

The following opinions were pubhshed on 31 March 1999 in Vol. .56, Part 1, of the Bulletin of Zoological Nomeclature.
Copies of these Opinions can be obtained free of charge from the Executive Secretarv International Commission on
Zoological Nomenclature, c/o The Natural History Museum, London, Cromwell Road, London SW7 5BD. U.K.
(iczn@nhm.ac.uk).

OPINION 1913. Pila Roding, 1798 and Pomacca Periy, 1910 (Mollusca, Gastropoda): placed on the Official List, and
Ampullariidae Gray, 1824; confirmed as the nomenclaturally valid synonym of Ph.idae Preston, 1915.

OPINION 1914. Belemnotheutis Pearce, 1842, Geopeltis Regteren Altena, 1949, Geotciithis Miinster, 1843, Jeletz-
h/tcuthis Dovle, 1990, Loligoscpia Quenstedt, 1839, Parahclopcltis Naef, 1921, Paraplcsioteiitlm Naef, 1921 (Mol-
lusca. Coleoidea): conserved, and the specific name of Beleninotcuthis (sic) montejiorci Buckman, 1880: consen'ed.

In Memoiiani

Sally Diana Kaicher

Februan- 19, 1922

March 29, 1999

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THE NAUTILUS

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OCT 1 b jggg \

Volume 113, Number 3
October 7, 1999
ISSN 0028-1344

A quarterly devoted
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T H E €9 N A U JU. L U S

CONTENTS

•ianogfapriic inewuiion

OCT 1 8 1999

WOOOU MOie, MA. Oi<543

Volume 113, Numbers
October 7, 1999
ISSN 0028-1344

D. L. Geiger

Description of Hahotis fatui new species (Gastropoda:
Vetigastropocla) from the tropical western Pacific ....

73

Serge Gofas

The West African Rissoidae (Gastropoda: Rissooidea) and
their similarities to some European species

78

Book Review

102

Notices fmii

Notices

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THE NAUTILUS 113(3):73-77, 1999

Page 73

Description of HaUotis fatui new species (Gastropoda:
Vetigastropoda) from the tropical western Pacific

D. L. Geiger

Department of Biological Sciences
University' of Southern California
Los Angeles, CA 90089-0371 USA
dgeiger@usc.edu

ABSTRACT

A new abalone species from the central Indo-Pacific is de-
scribed. The species has been known for almost two decades
and resulted in the introduction of a nomen nudum. The name
is validated here as Halioti.s fatui Geiger (e.x--Rehder MS). The
species is described from shell, radular, epipodial, and hvpo-
branchial gland characters. It is most similar to and occurs in
part of the range (Tonga, Solomon Islands, V'anuatu, and Ma-
rianas Islands) of the highly variable Haliotis varia Linnaeus,
1758. It differs from this latter species, however, in its shell
and epipodial characters.

Key words: Abalone, Gastropoda, HalioHdae, Haliotis fatui
new species, Indo-Pacific.

INTRODUCTION

Geiger (1998a) recently discus.sed all 200 species-level
taxa and the 17 genus-le\'e! t;Lxa in the family Hahotidae
Rafinesque, 1815. Most temperate species are fairly well
known; some tropical species, however, pose problems
in respect to their alpha t;Lxonomv. Geiger (1998b) dis-
cussed the controversial taxon H. clathrata Lichtenstein,
1794 {non Reeve, 1846), for which Geiger and Stewart
(1998) formulated Case 3036 now pending with the In-
ternational Commission on Zoological Nomenclature.
Stewart and Geiger (1999) designated a lectotvpe for H
crebri.'iculpta Sowerby, 1914, with a discussion on H.
clathrata Reeve, 1846. Geiger and Groves (in press) re-
view the fossil taxa including the few records from trop-
ical regions. The study of tropical abalone is continued
here with the introduction of a new species that has
been knowii for almost two decades. A nomcn nudum
had been introduced twice for this species under the
epithet "fatui Rehder" (Anon., 1981) and "fatui Rheder
[sic]. 1981?" (Ubaldi, 1993). Herein I clarify the system-
atics of this species and vahdate the name.

MATERIAL AND METHODS

Depository collections: AMNH, American Museum ol
Natural History, New York; ANSP, Academy of Natural

Sciences, Philadelphia; MHNG, Museum d'Histoire Na-
turelle, Geneve; USNM, National Museum ot Natural
History, Smithsonian Institution, Washington, D.C.

Studi/ of animals: Radula and epipodium were pre-
pared as detailed in Geiger (1996) and Stewart and
Geiger (1999). The terminology of radula parts of Hick-
man (1981) and Geiger (1996) is adopted.

SYSTEMATICS

Order Vetigastropoda SaKdni-Plawen, 1980
Family Haliotidae Rafinesque. 1815
Genus Haliotis Linnaeus, 1758

Remarks: As discussed by Geiger (1996, 199Sa), there
is currentK' no justification to subcb\ide the genus Hal-
iotis scnsu lato into subgenera. Accordingly, the species
described here is placed in that large genus. Further
study of the family may eventually identify distinct clades
worthy of ta-xonomic recognition, but this stage has not
yet been reached.

Haliotis fatui Geiger new species

(Figures 1-7, 9-16)

H varia subspecies — Kaicher (19S1: card no. 2902).
H. s-picjdata fatui Rehder — Anon, (1981 ;7) {nomen nudum).
H. fatui Rheder [.sic], 1981?— Ubaldi (199.3:11 .3-1) (nomcn nu-
dum).

Description (diagnostic characters underlined):

Shell mechum size (to 6 cm), medium weight, some-
what arched, sub-circular to ov;il-oblong; strong, kiiobby
bumps on dorsal surface of shell, usually in few, distinct,
spiral rows, occasionally also in prosocline rays. Spiral
threads strong, coarseK' granulated. Periphery between
row of holes and columella in smooth transition to dorsal
surface. Holes, small, round, slightly raised. Color mud-
dy green, brown, yellow brown. Color pattern irregular
to blotchy, no spiral elements, no prosocline rays. Col-
umella of mechiun wdth. Nacre light steel-gray. Muscle
scar absent in small specimens, present in some large
specimens, composed of fine, irregularly concentric ac-

Page 74

THE NAUTILUS, Vol. 113, No. 3

Figures 1-6. Shells ot HaUotis fatu, new species. 1, 2. Holot>pe, USNM 486708 46.6 mm, M.ng Island. 3 4- Parat>pe, USNM
487953, 29.5 .nm, Tonga. 5, 6. Paratope, MHNG 21226, 42..3 mm, Vanuatu. Note the vanabiUt>- m the extent of the bumps on he
dorsal surface of the shell, and the equally variable strength of the spiral cords. The tremata are generally rather small for the
genus, but note the differences between 1 and 5 as compared to 3.

D. L. Geiger, 1999

Page 75

Figures 7-9. Soft parts of Hnliotis fatui new species, and H.
varia Linnaeus, 1758. 7, 8. Epipodia with ilorsal side toward
top of page. Semi-schematic, reconstructed drawings from
camera hicida tracings. 7. Haliotis fatui, lioloty|ie, USNM
486708. Scale bar = 5 mm. 8. Haliotis varia. DLG unnum-
bered, Sengiggi, Lompok, Indonesia, leg. T. Baer shell length
= 27 mm. Scale bar = 1 nun. 9. Hyjiobranchial gland of H.
fatui. holotype, USNM 486708. Viewed from anterior to right
side. Scale bar = .5 mm. r = rectum

cretion lines (pathological feature?). Spire visible in ven-
tral view.

Epipodium (Figure 7) of medium width, dor.so-ven-
trally symmetrical, with undulating mid-epipodial fold.
Face covered with low, warty tubercles, larger elements
alternating dorso-ventrally. Fringe with irregular lobes,
secondary short projections, denser on large elements,
few between them. No hand-shaped structures, no dor-
sal or ventral tentacles.

Hyj^obranchial gland simple (Figure 9), neither sec-
ondaiy lamellae of the northeastern Pacific species, nor
the thick lamellae of H. asinina (Geiger, unpubl. data).

Radula (Figure 10) shghtlv asymmetrical, skewed to
left side. Rachidian tooth (Figure 11) somewhat narrow-

er than first lateral tooth; cutting edge not thickened,
postero-basal projection absent. Lateral tooth 1 (Figure

11) with straight primaiy ridge (pr), secondary ridge (sr)
much smaller than primary. Lateral teeth 3-5 (Figure

12) without denticles. Marginal teeth (Figures 13-15)
symmetrically denticulated.

Type material: Holoty^De, USNM 48670S, shell with
animal, radula mounted on stub, desiccated and mount-
ed piece of epipodium: Mang Island, N Marianas, 1945,
leg. Banner, 46.6 mm (Figures 1-2, 7, 9-15). Paraty^ies,
AMNH 81878, 1 shell, Santa Ana Lsland, Solomon Is-
lands, Whitney South Seas E.xpedition 1929-1930, 58.9
mm; AMNH 81899, 1 shell, Santa Ana Island, Solomon
Islands, Whitney South Seas Expedition 1929-1930, Jan.
1930, 53 mm; ANSP 284325, 1 shell, Kumbun Island,
SW New Britain, ex. coll. W. E. Jamison, Nov 1962, 40
mm; MHNG 21226, 1 shell, Vanuatu, leg. de Villoutreys,
42.3 mm (Figures 5-6); USNM 702975, 7 shells, NE
Tofua Island, Ha'apai Group, Tonga, April 1971, on coral
rocks off reef edge, leg. John H. Fatu, 29.3 mm, 36.4
mm, 40.4 mm, 42.2 mm, 46.9 mm, 49.2 mm, 52.7mm;
USNM 702512, 9 shells, reef off Tofua Island, Tonga,
April 1971, ex-L. J. Lancaster, 34 mm, 37.2 mm, 37.4
mm, 44.0 mm, 43.5 mm, 48.2 mm, 50.7 mm, 58.0 mm;
50.2 mm; USNM 487953, 1 shell, Agrigan Island, N Ma-
rianas, 29.5 mm (Figures 4-5); USNM unnumbered, 1
shell, Tonga, 53.1 mm; USNM unnumbered (ex-San Di-
ego Societv- of Natural HistoiT No. 32719), 2 shells, Ton-
ga Island, Vavau, leg.H. N. Lowe, 1924, 48.9 mm, 46.5
mm.

Type locality: Mang Island, Northern Marianas.

Distribution (Figure 16): Most specimens have been
found on Tonga, with additional specimens stemming
from the Solomon Islands, Vanuatu, and the Marianas
Islands. An overall western-central Indo-Pacific distri-
bution pattern seems to be emerging. From the little
information available, the species seems to live in shal-
low coral reef habitats, which is typical for tropical ab-
alone species (Geiger, 1996; Stevyart & Geiger, 1999).

Etymology: The name honors the collector of se\'eral
specimens from Tonga, John H. Fatu. The proxisional
name that has been associated with this species is vaU-
dated, following the original intent of the late Dr. H. A.
Rehderof the'uSNM.'

Comparative remarks: The only species similar to H.
fatui is H. varia Linnaeus, 1758. Haliotis varia is an ex-
tremely variable species and it is likeK' that some spec-
imens will not be separable from shell morphology
alone. The comparison provided here is between a typ-
ical H. fatui and a typical H. varia, i.e., the well-known
green form of the latter species from the Philippine Is-
lands. In H. varia the shell is more arched, usually heavi-
er, and the knobs are more weakly developed than in H.
fatui. The apertural lip is more ouhvardlv cur\'ed in H.
varia. The holes in H. fatui tend to be smaller and more
mnnerous, althouirh this last character is extremely var-

Page 76

THE NAUTILUS, Vol. 113, No. 3

Figures 10-15. Radula of llaliotis fatui new species, lioloUpe, USNM 486708 The enstalline precipitations are preparational
artifacts of unknown origin, 10. \'iew of entire radula from anterior third of rilibon. Note slight asvinnietrv. Scale bar = 1 mm. 11.
Rachidian and lateral tooth 1. pr: pnman- ridge, sr: secondary ridge. Scale bar = 200 \i.m. 12. Lateral teeth .3-.5 without denticles.
Scale bar = 200 jjim. 13. Face of entire left row of marginal teeth in posterior view. Larger and stronger teeth to the right are
from a more inner position of the preceding row than the more slender teeth to the left. Scale bar = 200 (jim. 14, 15. Cusps of
inner marginal teeth. Note symmetrical denticles. 14. Dorsal view. Scale bar = 100 (xm. 15. Ventral view. Scale bar = 100 |xni.

iable (Geiger and Groves, in press) within species. The
coloration of H. vaiia consists typically of some oblique,
prosocUne, jagged flammules, in green tones, whereas in
H. fatui, the distribution of pigment is rather blotchv
with green and brown specimens being equalK repre-
sented.

The epipodium of H. fatui is devoid of dorsal (ir ven-
tral tentacles (Figure 7), whereas these are prominent
in H. varia (Figure 8). Both species share alternating
larger elements, but this character is also shared with a
number of other abalone species. The presence of hand-
shaped elements in H. varia, as seen in H. unilatcralis
Lamarck, 1822, and H clathrata Reeve (Geiger, 1996;

Stewart and Geiger, 1999), distinguishes H. varia from
H. fatui. The radula of the two species are very similar
to each other, as well as to other tropical species such
as H. unilatcralis (see Geiger, 1996), H. clathrata Reeve
(see Stewart & Geiger, 1999), and H. asinina Linnaeus,
1758, and H. ovina Gmelin,1791 (see Chitramvong ct
al, 1998). Chitramvong ei al. (1998) noted strong dif-
ferences for lateral teeth 1 and 2 based on their fig. 3C.
However, their fig. 3C does not agree with their figs 3A,
3B, and 3D, which show a very different and very typical
lateral tooth 1. Overall, these results corroborate indi-
cations that radular characters better define taxa above
the species level (Geiger, unpublished data).

D. L. Geiger, 1999

Page 77

^?

%

*v* --^ 1

' ... V-

Figure 16. Map showing distribution of Haliotis fatui new
species.

Remarks: Kaicher (1981: card no. 2902) illustrated
specimens of this species. Anon. (1981), and Ubaldi
(1993), both mentioned the taxon as nomina niida as
discussed by Geiger (1998a: note 71). The mention by
Anon. (1981:7) of "Haliotis spicidata Reeve, 1846" ac-
tually refers to H. aepiculata Reeve, 1846, one of the
many synonyms of H. varia (see Geiger, 1998a). Al-
though most material available originates from Tonga,
the Marianas Islands specimen is chosen as the holotvpe,
because it is the sole specimen with the animal. The
distribution of H. varia circumscribes that ot H. fatui:
hence, I do not consider H. fatui a subspecies ot H. varia
(of. Kaicher, 1981), because sympatric subspecies by def-
inition do not exist.

The specimen from USNM 702512 measuring 50.2
mm had been separated from the main lot by Rehder
for designation as primary type; a handwritten label "Ho-
lotype" accompanied the specimen. This original inten-
tion of Rehder is here overridden.

ACKNOWLEDGMENTS

Thanks to Rave Germon, M. G. Harasewvch, and Alan
Kabat (USNM), Yves Finet (MHNG), Paula Mikkelsen
(AMNH) for accommodation at the respective institu-
tions and loan of specimens. The study of the animals

was made possible by grants from the Hawaiian Mala-
cological Sf)ciety, the Western Societv of Malacologists,
and the Lehner Gray Fund for Marine Research
(AMNH). The contribution benefited from discussions
with Mark Jones, Buzz Owen, and Kit Stewart. Jim Mc-
Lean anil two anonymous reviewers helped improve the
manuscript.

LITERATURE CITED

Anonvinous. 1981. Shell ot the month: The abalones ororniers
(Haliotis). The Mollusk, Greater Miami Shell Club. Inc.,
19(3):1, 7.

Chitramvong, Y. P., E. S. Upatham, .M. Kruatrachue. P. Sobhon
and y. Limthong. 1998. Scanning electron microscope
.stndv ot radulae in Haliotis fl.s(/i(>iflLinnaeus, 1758 and
Haliotis ovina Gmelin, 1791 (Gastropoda: Haliotidae).
Jonrnaj of Shellfish Research 17:755-759.

Geiger, D. 1996. Haliotids in the Red Sea, with neotype des-
ignation for Haliotis unilateralis Lamarck, 1822 (Gastro-
poda: Prosobranchia). Revue Suisse de Zoologie 103:339-
354.

Geiger, D. L. 1998a. Recent genera and species of the family
Haliotidae Rafinesque, 1815 (Gastropoda: Vetigastropo-
da). The Nautilus 111:8.5-116.

Geiger, D. L. 1998b. Note on theidentiW of Haliotis clathrata
Lichtenstein, 1794 (not Reeve, 1846). MoUuscan Research
19:157-159.

Geiger, D. L. and L. T. Gro\es. In press. Review of fossil ab-
alone (Gastropoda: V'etigastropoda: Hahotidae) with com-
parison to Recent species. Journal of Paleontology'.

Geiger, D. L and K. A. Stewart. 1998. Case 3036." Haliotis
clathrata Ree\e, 1846 inon Lichtenstein, 1794) and H.
elcoans Plnlippi, 1844 (Molkisca, Gastropoda): proposed
consenation of the specific names. Bulletin of Zoological
Nomenclature 55:209-21 1 .

Hickman, C. S. 1981. Evolution and function of asvmmetrv in
the archaeogastropod radula. The Veliger 23:189-194.

Kaicher, D. 1981. Card Catalogue of World Shells. Pack 28,
Haliotis. Author's edition, St. Petersburg, Florida.

Stewart, K. A. and D, L. Geiger. 1999. Designation of lectotvpe
for Haliotis crchrisculpta Sowerbs. 1914, with a discussion
of H. clathrata Ree\e, 1846 inon Lichtenstein, 1794), The
Veliger 42:85-96.

Ubaldi, R. 1993. Atlas of the li\-ing abalone shells of the world
[First part with introductorv notes and four species treat-
ed]. Associazione Malacologica Inteniazionale, Rome [ir-
regular pagination].

THE NAUTILUS 113(3):78-101, 1999

Pa^e 78

The West African Rissoidae (Gastropoda: Rissooidea) and their
similarities to some European species

Serge Gofas'

Museum National d'Histoire Naturelle
Laboratoire de Biologie des

In\ertebres marins et Malacologie
55 me Buffon, F-75005 Paris.

FRANCE

ABSTRACT

The family Rissoidae, which is represented b\ man\' species in
temperate European and Macaronesian shore and shelf fauna,
is found to be also well represented in West Africa. Eighteen
species are reported, of which 10 (7 Alvania, 2 Crisilla, 1 Ze-
bina) are here described as new. Only 2 species are shared
with the Western European fauna, and all the temperate lin-
eages hnked to photophile algae or marine phanerogams (i.e.,
the genus Rissoa. and some lineages in Alvania) are missing in
West Africa. Four (3 Alvania and 1 Crisilla) ot the shelf species
with planktotrophic lar\ae are closeh' related to a planktotroph-
ic species of \\'estem Europe, and tliis is tentati\'el\' interpreted
as the result of breakup of formerly broader latitudinal ranges.

Key words: ^^este^l Africa, new species, species diversitv.

INTRODUCTION

The Rissoidae are a prominent part of the httoral, shelf
and upper bathval molkiscan faunas in the Mediterra-
nean, temperate Western Europe and in the Atlantic ar-
chipelagoes: Canary Islands (Moolenbeek and Faber,
1987; Moolenbeek and Hoenselaar, 1989), Madeira
(Watson, 1873), Azores (Dautzenberg, 1889; Gofas,
1990). Pallary (1920) reported twenty species along the
coast of Morocco, south of Cape Spartel, but the family
is believed to become abruptK' impoverished further to
the south. Nickles (1950) stated that there are 6 species
in 3 genera for the whole of West Africa. Indi\idual spe-
cies have later been described (Moolenbeek and Piers-
ma, 1990; Rolan and Fernandes, 1990) but the general
perception is still one of very low number of species.

The holdings in the Mu.seum National d'Histoire Na-
turelle of Paris (MNHN) include a large number of lots
from West Africa, which shows that this perception is
essentially a result of low sampHng efforts for small con-
tinental shell species in the past. Eighteen species are
represented, of which 10 are new and will be described
herein, and 4 were described in the last decade.

' Current address: Departamento de Biologia Animal, Um\er-
sidad de Malaga, E-29071 Malaga, Spain, sgofas@unia.es

MATERIAL AND METHODS

All the material examined is deposited in MNHN except
where otherwise stated. Listings of material examined
are arranged from North to South.

The main sources of MNHN material are:

Mal'rit.ani.\: shore collecting bv P. Bouchet in 1983,
and dredgings by WV "N'Diago" bv B. Richer de
Forges in 1981;

Seneg.al: collections made by I. Marche-Marchad be-
tween 1953 and 1956, mostly bv dredging from shore
to ca. 250 m depth; some material collected by M. Pin
or by K. Leung-Tak since 1980:

I\'ORV Coast: dredging on the continental shelf by P. Le
Loeuff;

Angola: collecting h\ the author (dredging, shore col-
lecting and snorkling) from 1981 to 1987, jointly widi
the late F. Fernandes.

The remainder of the West African coast has been very
poorly sampled for micromollusks, so that apparent dis-
junctions in the ranges are not supported by absence in
a homogeneous sampling effort.

Some of the Angolan material was sieved in seawater
and partly sorted in situ, which allowed for preparation
of drawings of the hxing animals. This proved particu-
larly useful for observation of color patterns and dehcate
stnictures such as pallial and metapodial tentacles, which
Ponder (1985) scored as \en' informative taxonomic
characters. Details about origin of the specimens ex-
amined for descriptions of externid nioqihology are giv-
en in the "material examined " section of each species.

Specimens were measured with precision of 0.05 mm.
Measurements of largest and smallest adult specimens
or shells in each lot are given with material examined.
Specimens selected for scanning electron microscopy
were sonicated, and some washed in a 10% solution of
sodium-lauPil sulphate.

Particular attention has been given to protoconchs as
taxonomic characters at the species level. Supraspecific
classification follows Ponder (1985), except for change

S. Gof'as, 1999

Paee 79

ol rank oi Chsilla troni subgenus to genus, lollowing
Bouchet and Waien (1993).

List of ahbreviations and text conxentions:

coll.: collected bv

shell(s): collected dead

specimen(s): collected alive

sta.: station number

AMS: Australian Musemn, Svdnev

MNCN: Museo Nacional de Ciencias Naturales,

Madrid
MNHN: Museum national d'Histoire naturelle,

Paris
UAN: Universidade Agostinlio Neto, Luanda

ZMA: Zoologisch Museum, Amsterdam

SYSTEMATICS

Family Rissoidae Gray, 1847
Subfamily Rissoinae Gray, 1847
Genus Piisilliiw Monterosato, 1884

T\pe species: Ris.soa piisilhi Philippi, 18.36, by monotyj^y (pre-
occupied by Ris.soa piisilla Grateloup, 1S2S, renamed fl(.$-
soa pliilippi Aradas and Maggiore, 1844).

Pusillina inconspiciia (Alder, 1844)
(Figures 1-8)

Rissoa inconspicua Alder, 1844: 32.3, pi. 8 figs. 6-7.

Description: Shell conical, adults 1.45x0.9.5 to
2.75X1.7 mm. Protoconch of 3 rather convex whorls;
protoconch 1 less than 1 whorl, smooth; protoconch 2
with a spiral thread running next to the suture on ab-
apical part of whorls, and with very sinuous, indistinct
growth lines on last lanal whorl. Teleoconch of 2.5 to 3
whorls; spire whorls slightly convex, regularly increasing
in size, sculptured with sinuf)us axial ribs variable in
number (10 to 24 on body whorl) and development, and
faint spiral cords between these ribs: body whorl with
axial ribs interrupted abniptlv at periphen', which leaves
penumbilical area sm(M)th. Aperture pNiilorm; Outer lip
orthochne, smooth inner surface, thickened externally at
some distance from edge by a vailx of variable strength
developed mostly on adapical part, then thinning out to
a cutting edge. Inner lip thin, bordering a narrow um-
bihcus. Shell color buif, with square blotches abapicalK
along suture; protoconch often tinged with purple or
dark browri.

Head-toot with superficial black and vellow pigmen-
tation; black marks on upper midline ol propodium and
on anterior part of opercular lobes, on more intensely
pigmented specimens also on sides of propodium and
over posterior part of head. Tentacles cylindrical, with
an axial series of yellow blotches; eyes in a small liulge
at base of each tentacle. Snout bilobed, with a yellowish
buccal mass \isible by transparency. Yellow granular
masses behind each eye; other more irregular superficial
yellow marks on snout, propodium, opercular lobes (be-

hind black area) and sometimes on periphery of foot
sole. A deep vellow or orange bar \isible b\' transparency
beneath operculum. Single metapodial tentacle rather
slender. Sole of foot colorless or with few peripheral yel-
low marks, with posterior pedal gland apparent around
a small longitudinal slit. Right paUial tentacle small, col-
orless with a yellow blotch at its base; left pallial tentacle
not detected, a velkjw blotch present at its expected lo-
cation.

Type locality: Northumberland coast (NE Great Brit-
ain), in deepish waters among corallines".

Material examined: Western Europe: Malaga
(Spain), in coarse sand 20-40 m, 100 shells (1.35 X 0.85
to 2..55 X 1.5 nun), coll. Gofas, Sept. 1990; Sagres (Por-
tugal), on coarse sand near cliffs 17-23 m, 11 specimens
(1.3 X 0.85 to 1.8 X 1,1 mm), mission Algai-ve, May
1988; and several hundred specimens from throughout
the range. Mauritania: 3 miles W of Kiaoue (20°02'N,
16°22'W), 1 .shell (1.85 X 1.25 mm), coll. Marche-Mar-
chad. May 1965; Port Etienne (now Nouadhibou,
20°20'N, 'l6°22'W), 10 m, 64 shells (1.5X0.95 to
2.25X1.35 mm), coll. Marche-Marchad, May 1965; Baie
de I'Etoile, 20 .shells (1.8X1.15 to 2.6x1.7.5 mm), coll.
Bouchet, 1983. Senegal (coll. Marche-Marchad except
where othe:-vvise noted): Baie de Goree 10 m, 90 spec-
imens (1.55X0.95 to 2.15X1.4 mm); Baie de Goree 30
m (1.4X0.9 to 1.9X1.25 mm); Baie de Goree 3.3-34 m,

6 shells; Off Goree, sta. 55-6-7A, 145-170 m, 3 shells;
Baie de Goree 25 m, S. of "Tacoma" shipwreck, 9 shells
(1. 50X0.95 to 2.00X1.20 mm); South of Goree 110-112
m, 16 shells (1.4X0.9 to 2.3X1.75 mm); Dakar area 97-
98 m, .30 shells (1..55xl.()5 to 1.8X1.2 mm); Dakar area,

7 shells (2.15X1.35 to 2.45x1.5 mm); Dakar area 78 m,
sta. 55-6-3D (14°19'N, 17°23'W), 5 shells (1.75X1.15
to 1.9X1.5 mm); SW of Cap Manuel 250 m, 1 shell
(1.9X1.3 mm); Cap de Naze, in stomach of Alnfcits
pimctatus, 2 specimens (2.25X1.45 to 2.35X1.45 mm);
Off Saloum, sta. .55-;3-9B, 80 shells (1.4X0.9 to
1.85X1.25); Off Saloum, 40 shells (1.5X0.95 to 2.4X1.4
mm); Saloum, mangroves, sta. 8, 25 specimens (2.0X1.0
to 2.75X1.7 mm), coll. Bouchet, 1973. Gabon: Conga,
Bay of Corisco, 32 shells (1.4X0.95 to 2,15X1.2 mm).
Congo: Pointe Noire (Plage Mondaine), 4 shells, coll.
von Cosel, 1985. Angola (coll. Gofas): Ambrizete fight-
house, 25 shells; S. of Ambrizete, Bango, 90 shells; Barra
do Dande, 13 shells (1.25X0.85 to 1.6X0.95 mm); Praia
Sao Tiago, 3 shells; Cacuaco intertidal; 3 specimens and
13 shells; Cacuaco .5-10 m, 2 specimens and 3 shells;
Etambar, rocks near beach, 18 specimens and 20 shells;
Corimba, 20 m on bioclastic gravel, 5 specimens and 400
shells (1.25X0.9 to 2.05X 1.3 mm; fiving animal observed
and drawni); Bay of Mussulo km 17, on algae at low tide,
120 .specimens (1.45X1.05 to 1.95X1..35 mm); Bay of
Mussulo km 31, 1.5 specimens (living animal obsei"ved
and drawn); Off Mussulo 90 m, 16 shells (subfossil). Off
Mussulo 120 m, 50 shells (subfossil); Cape Palmeirinhas
(Buraco) on rocks 2 m, 2 specimens (living animal ob-
sened antl tlrawm); Santo Antonio near Benguela, 4

Page 80

THE NAUTILUS, Vol. 113. No. 3

Figure 1. Pii.silL

isulinn tncon.

spiciia (Alder, 1844). Living animal from off Corimba (Angola), shell length 1.7 mm.

shells; Lucira (Sta Marta) 40 m, 1 shell; Lucira (Bisson-
ga) on gravel of coralline algae, 2 shells; Chapeu Ar-
1 specimen and 3 shells (living animal observed

mado,

and drawai); Sao
tidal, 1 shell.

Nicolau, 5 shells; Praia Amelia, inter-

Habitat: Most common on suhtiilal hioclastic coarse
sands or gravels, in 10-60 m depth. Also on httoral algae
from shore to subhttoral sites, and on deeper muddv
bottoms on the continental shelf.

Distribution: From northern Norway to Angola, and
the Mechterranean. Unverified record from the Azores
(Nordsieck, 1972).

Remarks: This species is characterized bv a multispi-
ral protf)conch (see \^'aren, 1996: 1042) with a small pro-
toconch 1 of less than 1 whorl, and a protoconch 2 of
sUghtly more than 2 whorls, suggesting that a long plank-
totrophic stage is present. Nevertheless, the protoconch/
teleoconch transition is nsualK' extrenieK ilifficult to de-
tect, even under scanning electron microscopv. It is in-
dicated by a change over less than 1/4 of a whorl, from
the extremely sinuous growth lines that correspond to
the notches that accommodate the velar lobes of the
lar\'a, to nearly axial growth lines formed after the velum
is lost.

The range of bathynnetric, geographic and moqiho-
logical variation among West African PusiU'mo suggests
that more than 1 species may be present. Individuals
vary in shell morpholog)' and color pattern (ribs strong
to fine to nearly absent; apex tinged or not with puiple,
subopercular bar orange or yellow, yellow spots extend-
ing or not onto the tentacles and foot). The habitat range
is also too broad for what is usual in a single species: in

algal wash in the lagoon of Bay of Mussulo, but also on
muddv bottoms on the outer part of the continental
shelf. Nevertheless, there are no stable combinations of
characters and thus no convincing gromids for recogni-
tion of more than 1 species at anv particular site. Mem-
bers of the population found in lagoonal environments
of Mussulo have multispiral protoconchs and do not dif-
fer in this respect from open sea populations. Thus,
there is no differentiation comparable to that of the Eu-
ropean Pusillina liiwohita ( Midland, 1832) and P. incon-
spiciia, the former clearly separated from the latter by
1 less protoconch whorl.

Genus Sctia H. and A. Adams. 1852

Tvpe species: Rissoa pulchcrrima Jeffreys, 1848, subsequent
designation bv Kobelt (1878).

Sctia nomca Moolenbeek and Piersma, 1990
(Figures 9-10)

Sctia iioiiicd Moolenbeek and Piers

1990: ,31-,3.3, figs. 1-4.

Description: Shell conical, adults 1.25X0.85 to
1.55X0.95 mm. Protoconch of 1.3 to 1.5 whorls, regu-
larly convex, with 9-10 strong spiral cords. Teleoconch
of 2.5 whorls; spire whorls verv' convex, regularlv increas-
ing in size, sculptured with faint, uneven spiral cords (9-
12 on penultimate whorl); bodv whorl rounded with spi-
ral lines continued all over. Apertiu'e rounded, continu-
ous; Outer lip orthocline, smooth inner surface, hardly
thickened externallv. Inner hp thin, bordering a narrow
umbilicus. Shell color buff, with 2 internipted, very
slightlv darker bands mnnning along adapical and ab-

S. Gofas, 1999

Pa^e 81

Figures 2-8. Pusilhna inconspicua (Akler, 1S44), 2. Specimen troni Sagres (Portugal), sliell length 1.65 mm. 3. Protoconch of

the same specimen. Scale bar = 100 jjini. 4. Specimen from off Saloum (Senegal), shell length 2.1 mm. 5. Protoconch of the same

specimen. Scale bar = 100 |jLm. 6-7. Specimens from Bav of Mussulo (Angola), shell lengths 1.6 and 2.1 mm. 8. Protoconch of

the specimen in fig. 7. Scale bar = 100 (xm.

Figures 9-10. Sctia nomen Moolenbeek and Piersnia, 1990. 9. Specimen from Baie tie I'Etoile (Manritanie), shell length 1..3

mm. 10. Protoconch of another specimen from the same localitv. Scale bar = 100 jjim.

Figure II. Obtiisellri intcrsccta (Wood, 1857). Specimen from off Ilha de Luantla (Angola), shell length 0.9 mm.

apical 1/3 of the spire whorls, and a third hand on peri-
umbihcal area of body whorl.

Type material (not seen): Holotype ZMA Moll. n°
■3. 89. 018 and 68 paratvpes from t\pe locahty.

Tjpe locality: Serini (Mauritania, 19°34'N, 16°25'W)
low tide level.

Material examined: Mauritania: Baie de I'Etoile, in

Pase S2

THE NAUTILUS, Vol. 113, No. 3

tertidal, ca. 400 specimens (1.15X0.85 to 1.55X0.95
mm), coll. P. Bouchet 1983.

Habitat: On large tidal flats.

Distribution: This species is knowii onlv from a small
stretch ot the Mauritania coast.

Remarks: The generic allocation is not inllv supported
but the species is herein treated in its original binominal
combination until characters of soft parts are made axail-
able. The general shape of the shell, presence of spiral
sculpture, and intertidal habitat are tvpical of Riulolpho-
setia Monterosato, 1917 (t\pe species Tnuicatclla fii.sca
Philippi, 1836; subsequent designation by Kobelt, 1878),
a taxon which Ponder ( 1985) placed in the synonymy of
Setia. Protoconch sculpture with strong spiral cords is a
character shared with other species currentK placed in
Alvania (e.g., Alvania pairula Jeffreys, illustrated in Go-
fas and Waren, 1982; Alvania watsoni (Watson), Ponder,
1985, p. 151 fig. 102D); Setia pidchcrriina does have
spiral microsculpture on the protoconch, but one of vei-y
fine threads that look separated from the surface (Pon-
der, 1985, p. 134 fig. 85G; Gofos, 1990, fig. 33). The
teleoconch sculpture is also shared with some species of
Alvania (e.g., A. paiviila, A. punctnra) and not with So-
fia pu

mlchenima, which shows only growth lines.

Genus Alvania Risso, 1826

T\pe species: Alvania europaea Risso, 1826, subsequent des-
ignation by Nevill, 1885.

Alvania africana new species
(Figures 12-19)

Description: Shell high conical, sfightly cvTtoconoid,
solid, adults 1.9X0.95 nun to 3.0X1.6 mm. Protoconch
of 2.2 to 2.3 convex whorls, regularly conical; surface of
larval whorls (except for narrow subsutural band) with
spiral lines of small granules, which merge into 3—4 con-
tinuous threads on abapical part of last laival whorl
Teleoconch of 2.3 to 4.3 whorls; spire whorls convex,
with strong, widely spaced a.xial folds (ca. 15-25 on pen-
ultimate whorl) superimposed by strong spiral cords; 2
cords on first teleoconch whorl, 5 (rarelv 4) on penulti-
mate whorl; increasing in strength from subsutural cord
to abapical one, this latter overhanging adjacent suture.
Body whorl rounded, shghtly constricted, with 9-10 spi-
ral cords and a.xial folds gradualK- fading toward peri-
umbilical area. Outer lip oprsthocline, externally thick-
ened by very strong rim strongK differentiated from ad-
jacent external shell surface, then thinning out into del-
icate edge; superimposed by spiral cords that continue
onto edge of lip; internally with 7-8 strong denticles.
Inner fip slightly thickened, appressed on imperforate
columella.

Shell color tawny with 2 broad, blurry reddish browm
bands on body whorl, one subsutural and another on
abapical part of body whorl; bands better defined inside

Figure 12. Alvania africana new species. Living animal (par-
ahpe) from off Ilha de Luanda (Angola), shell length 2.3 mm.

aperture where they terminate at edge of outer lip. Col-
umella generally tinged with same reddish browm; some
specimens entireK- reddish browii, others pale and lack-
ing bands.

Head-foot colorless except for yellowish granular mas-
ses behind each eye, and faint brownish hue ;it tip of
snout. Tentacles parallel sided, e\es in small bulge at
base of each tentacle. Snout markedly bilobed, with pink
buccal mass visible by transparency. Three metapodial
tentacles, rather small and short, clustered tightK. Sole
of foot colorless, with pedal gland inconspicuous. Right
and left pallial tentacles rather conspicuous, with parallel
sides, colorless.

Type material: Holotype (specimen 2.3X1.3 mm)
and paratope (specimen 2.3X1.3 mm; living animal ob-
served and drawn) from type localitv (MNHN). Paratv-
pes (9 specimens 2.3X 1.2 to 2.9X 1.45 mm; living animal
observed and drawai) from off Ilha de Luaiula, 40-60 m
(MNHN). Paratvpes (2 specimens 2.35X1.25 to
2.7X1.45 mm and' 10 shells 1.9X1 to 2.7X1.4 mm) from
off Mussulo, 90-100 m (AMS). Paratvpes (2 specimens
2.4X1.3 to 2.45X2.6 mm and 10 shells 2.0X1.2 to
2.75X1.45 mm) from off Mussulo, 9t)-l()0 m (UAN).

Type locality: Off Ilha de Luanda, Angola, shell grav-
el, 75-80 m.

Material examined: Mauritania (coll. Richer de
Forges): "Ndiago' ,sta. 317 (19°12'N, 16°4()'\\'). 47 m,l
shetl (apex broken); "N'Diago" sta. 321 {19°18'N,
16°47'W), 40 m, 1 shell. Senegal (coll. Marche-Mar-
chad): Off Saloum, 50 m, 4 shells (2.35X1.2 to 3.15X1.5
mm); Baie de Goree, 1 shell (1.9X0.95 mm); Off Goree

S. Gofas, 1999

Page 83

50 m, sta. 55-7-5A. 1 shell (2.35X1.25); Off Goree 95
111,1 shell (2.1X1.05 mm). Cape Verde Islands: Porto In-
gles, Maio island, "Sylvana" sta 136, 1 shell (listed as A.
cancellata by Lamy, 1923). Angola (coll. Gofas): Tyi^e
material and off Ambrizete 45 m, 40 shells, (1.9X().95
to 2.75X1.4 mm); Off Ambrizete 80 m, 60 shells
(2.05X0.95 to 2.7X1.25 mm); Off Ilha de Luanda 40-
60 m, 50 shells (2.2X 1.2 to 2.9X 1.45 mm); Corimba 10-
20 m, 5 specimens (2.75X1.5 to 3.0X1.6 mm); Off Mus-
sulo 90-100 m, from fishermen, 3 specimens and 50
shells (2.35X1.25 to 2.75X1.45 mm); Cabo Ledo 40 m,
1 specimen (2.85X1,45 mm); Santa Maria 8-10 m, 1
specimen (3.0X1.5 mm); Bay of Lucira (Cesar) 10 m in
maerl, 5 shells (2.35X1.25 to 2.95X1.55 mm); Santa
Marta 40 m, 6 shells (2.35x1.2 to 3.0X1.55 mm); Praia
Amelia 40-60 m. 200 shells (2.15X1.2 to 2.65X1.45
mm); Porto Alexandre, 2 specimens (2.25X1.3 mm; liv-
ing animal observed and drawn).

Habitat: On hard bottoms of rocks, coarse shell gravel
or (in Santa Maria and Lucira, Pro\'ince of Benguela,
Angola), concretions of calcareous algae, 10-60 m.

Distribution: Mauritania and Senegal; northern and
southern Angola. The record from Cape Verde Islands
needs confirmation.

Remarks: This species mostlv resembles the Euro-
pean species Alvania heani (Hanley, 1844), having same
general outhne, type of teleoconch sculpture, multispiral
protoconch, and colorless animal. The main differences
are: (1) the protoconch is broader, with Hatter spire an-
gle, the spiral cords near abapical suture of last proto-
conch whorl are stronger, (2) teleoconch in average
seems to be more slender, (3) the suprasutural cord on
all spire whorls is overhangs the suture more conspicu-
ously (juveniles look somewhat carinate), and (4) adults
are smaller. The 2 species are not sympatric, but A. be-
am is present with a apparently consistent range of varia-
tion from the British isles to southern Morocco, then
separated hy ca. 1000 km from the knovsni West African
range of A. africana.

Alvania maiioi new species
(Figures 20-24)

Description: Shell high conical, solid, adults 1.9X1.2
mm to 3.15X1.9 mm. Protoconch of 2.2 to 2.5 convex
whorls, regularly conical; surface of larval whorls (except
a narrow subsutural band) with spiral lines of small gran-
ules, which merge into 3—4 quite continuous threads on
abapical part of last lai"val whorl; latter sometimes bro-
ken into slightly oblique segments arranged along spiral
line. Teleoconch of 2.7 to 3.9 whorls; spire whorls con-
vex, with strong, widelv spaced iLxial folds (ca. 16-20 on
penultimate whorl) and strong spiral cords; intersections
spinose; 2 cords on first teleoc(jnch whorl, 3 cords of
equal size on penultimate whorl; adapical cord at some
distance from suture forming shouldered whorl. Body
whorl rounded, v\qth 9-10 strong spiral cords and LLxial

folds graduallv fading toward periumbilical area where
the\' are onl\' expressed as beads on spiral cords. Outer
lip slightlv opisthochne, externall) tliickened by strong
rim strongly differentiated from adjacent external shell
surface, then thinning out into delicate edge; superim-
posed bv spiral cords that continue without fading onto
edge of lip; internallv with 6-7 denticles. Inner lip slight-
ly thickened, appressed on imperforate columella.

Shell color whitish, with extremely tenuous brownish
tinge sometimes on bodv whorl adjacent to apertural
rim; commonlv incrusted with loistv stains on li\ing spec-
imens.

Head-foot colorless, with elongate opaque white gran-
ular masses behind each eye, and faint browniish hue at
tip of snout. Tentacles parallel sided, slender; eves in
small bulge at base of each tentacle. Snout markedly
bilobed, with pink buccal mass visible by transparency.
(No notes on metapodial tentacles). Sole of foot color-
less, with pedal gland inconspicuous. Right and left pal-
hal tentacles rather conspicuous, with parallel sides, col-
orless.

Type material: Holotype (specimen 2.75X1.6 mm)
and 5 paratypes (shell 2.5X1.55 to 2.8X1.65 mm) from
the t\pe localitv- (MNHN). Paratype (specimen 2.7X1.65
mm, li\ing animal obser^'ed and drawTi) from off Ilha de
Luanda, "75-80 m (MNHN). Paratvpes (10 shells
2.0X1.3 to 2.75X1.65 mm) from off Mussulo 90-100 m
(AMS). Parat\pes (10 shells 2.25X1.1.45 to 2.75X1.7
mm) from off Mussulo, 90-100 m (UAN).

Type locality: Off Ilha de Luanda (Angola), shell grav-
el', 40-60 m.

Material examined: Mauritania: "Ndiago" sta. 316
(19°12'N, 16°37'W), 34 m, 1 specimen, coll. Richer de
Forges 1981. Senegal (coll. Marche-Marchad unless oth-
erwise noted): Bale de Goree 12 m, 3 specimens
(2.5X1.5 to 2.8X1.7 mm), coll. Bouchet; SW of Made-
leines 455 m, 1 shell (2.25X1.4 mm); SW of Goree 150-
250 m, 2 juvenile shells; Baie de Goree, S. of "Tacoma"
.shipwreck, 25 m, 4 .shells (1.6X1.1 to 2.2X1.4 mm); Baie
de Goree 30 m, 1 juvenile shell ; SW of Madeleines 50
m, 3 shells (2.4X1.4 to 2.6X1.5 mm); Off Saloum 50 m,
18 shells (2.1X1.3 to 2.55X1.5 mm). Ivorv Coast: "Ra-
fale" sta. 6 (5°01.5'N, 3°23.5'W), 70 m, i shell and 2
fragments. Angola (coll. Gofas unless othenvise noted):
Type material and off Ambrizete 45 m, 5 shells
(2.75X1.6 mm to 2.9X1.65 mm); Off Ambrizete 80 m,

15 shells (1.9X1.25 to 2.65X1.6 mm); Off Corimba 10-
20 m. 10 shells (2.7X1.7 to 3.15X1.9 mm); Off Ilha de
Luanda 40-60 m, 40 .shells (2X1.2 to 2.8X1.6 mm); Off
Ilha de Luanda 120 m, 14 shells (2.25X1.4 to 2.7X1.7
mm); Off Mussulo 90-100 m, from fishermen, 60 shells
(2.25X1.4 to 2.9X1.75 mm); Santa Maria 5-10 m, 1
.shell (3.25X1.85 mm); Santa Marta 40 m, 10 .shells
(2.6X1.6 to 2.9X1.95 mm); Sao Nicolau, shore, 22 .shells
(2.3X1.5 to 3.0X1,85 mm); Off Praia Amelia 40-60 m,

16 specimens + 85 shells (2.7X1.5 to 3.1X1.8 mm); Off
Praia Amelia, dredged 15-35 m, 5 juvenile and 1 adult

Page 84

THE NAUTILUS, Vol. 113, No. 3

Figures 13-19. Alvnnia africana new species, troni Angola. 13-14. Holotrvpe, from off Ilha de Luanda, shell length 2. .3 mm.
15. Protoconch of the hulot)pe. Scale bar = 100 ixm. 16. Detail on a teleoconch whorl of the holotvpe, showing the reticulate
pattern of the periostracum. Scale bar = 100 |xm. 17. Specimen from off Coriniba, shell length 2.75 mm. 18. Protoconch of the
same specimen Scale bar = 100 (j.m. 19. Shell from off Amhiizete, shell length 2. .3 mm

shell (2.0X1.2 mm), coll. Mi.s.sion Gnivel May 1910
(Dautzenberg collection. Institut Roval des Sciences Na-
turelles, BrtLxelles; identified by Dautzenberg as Alvania
cancellata (da Costa, 1778)).

Habitat: On hard bottoms of rocks, coarse shell gra\el
or (in the pro\'ince of Benguela), concretions of calcar-
eous algae, 10-60 m; often found with A africana.

Distribution: Mauritania and Senegal; hon Coast;
northern and southern Angola.

Etjinolog}-: This species is named after Mario ^Albano
dos Santos, a friend from Mo^amedes, who owns there
a crab fisher\^ and has hosted many of the collecting trips
to Southern Angola.

Remarks: This species resembles Alvania cancellata
(da Costa. 1778) v\ith which it shares coarse sculpture

and strong nodes on outer lip at the end of the abapical
spiral cords. It is distinguished hv being smaller and
more slender and having a different constniction of the
most aliapical cord (the one nmning ne.xt to columella)
(){ bod\ whorl, narrow and Ijeaded in A. inarioi, and onlv
a massive concretion in A. cancellata.

The citation by Dautzenberg (1913) of A. cancellata
is based on specimens ot A. marioi (material examined
at IftSN, Bnissels). Nickles (19.50) figured tnie A can-
cellata (presmnably a specimen ol European origin) but
included the species in his guide on the basis of "being
found in Angola" (i.e., based on the record bv Daut-
zenberg, 1913, as no further data on Angola were avail-
able in 1950).

Alvania venu.s (d'Orbignv, 1852), a well known fossil
species from the Miocene of Aquitaine Basin, S\\'
France, resemliles A. marioi. with which it shares die out-

S. Gofas, 1999

Page 85

Figure 20. Alvnnia inarioi new species. Living animal (par-
atvpe) from off Ilha de Luanda (Angola), shell length 2.7 mm.

line and size, the kind of sculpture and the shouldered
whorls with a cord at some distance from the suture. It
is nevertheless distinguished by having a third cord ap-
pearing on early teleoconch whorls

This species may be confused with the Mediterranean
A. hispidula Monterosato, 1S84 (figures 25-26), with
which it shares similar proportions, pattern of reticula-
tion, and protoconch. One of the consistent differences
resides in the architecture of the subsutural area, shoul-
dered in A. marioi, with a strong cord at some distance
from the suture; sloping, with a small subsutural cord in
A. hispidula. The abapical part of the body whorl is also
produced differently: 2 shaip, neatk' beaded cords sep-
arated by a deep depression and terminating with
marked indentations on the outer lip and then a simple
cord along the umbilical chink in A. inarioi; in A. his-
pidtda the 2 cords are more roughlv irregularlv beaded,
not so sharp, and weakly marked over the outer lip; the
last small cords along the umbilical chink tend to be
duphcated. The relationships of A. hispidula to coarse
forms ("A. calathus Forbes and Hanley") of the A. hcaiii
complex have yet to be documented. Alvania hiiia
(Monterosato, 1884), a species with a restricted distri-
bution in the southeastern part of the Western Medi-
terranean, resembles this species in the architecture of
the subsutural area, and in size and proportions. It dif-
fers in having a paucispiral protoconch iind onK 2 cords
on the penultimate whorl.

Alvania flexdis new species
(Figures 27-32)

Description: Shell high conical, cyrtoconoid, moder-
ately solid, adults 1.0X0.7 mm to 2.0X1.2 mm. Proto-
conch of 2 to 2.2 convex whorls, first one sUghtly de-
pressed; surface of lanal whorls (except for narrow sub-
sutural band) with spiral hues made of closely set or con-
tiguous small granules. Teleoconch of 2 to 3 whorls, with
sculpture of thin, shaip, widelv spaced spiral cords su-
perimposed on thin, sinuous axial ribs. Spire whorls have
1 cord running along adapical suture, second cord quite
close to subsutural one, and suprasutural cord on abapical
side, overhanging suture at some distance; spir;il cords are
poorlv de\'eloped in beUv'een. One or 2 additional spir;il
cords gradu;Jly appear in that internal on penultimate
whorl, so that there are about 10 evenly spaced spiral
cords on bod\ whorl. Axial ribs verv graduaJlv fade out
on abapical part of bodv whorl. Ribs on bod\' whorl 1.5-
30. Outer lip opisthocline, widi thin edge and thickened
externally at distance of edge by broad rim, superimposed
by spiral cords; smooth and not thickened intem;illy. In-
ner lip thin, bordering tiny umbilical chink.

Shell color tawny with 2 very faint brownish bands,
one over the 2 subsutural cords, other approximately at
anterior 1/3 of body whorl, ending in a more intense
blotch near edge of aperture.

Type material: Holotype (shell 1.6X1.05 mm) and 5
paratvpes (shells 1.45x0.95 to 1.85x1.1 mm) from tvpe
loealit\- (MNHN). Parat\pes (10 shells 1.2X0.75' to
1.9X1.15 mm) from off Mussulo, 90-100 m (AMS). Par-
atvpes (10 shells 1.25X0.8 to 1.95X1.15 mm) from off
Mussulo, 90-100 m (UAN).

Type locality: Off Ilha de Luanda (Angola), shell grav-
el, 40-fiO HI.'

Material examined: Senegal (coll. Marche-Marchad):
Off Saloum 50 m. 9 shells (1.05X0.7 to 1.5X0.9 mm);
Dakar area 97-98 m, 1 shell (1.6X1 mm). Equatorial
Guinea; (1°40'N, 9°25'E), 150 m, 1 shell (1.3X0.8 mm).
Angola (coll. Gofas); Type material and off Ambrizete 45
m,'l8 shells (1.05X0.7 to 1.35x0.9 mm); Off Ambrizete
80 m, 38 .shells (1.05X0.75 to 1.75X1 mm); Off Illia de
Luanda 40-60 m, 20 .shells (0.95X0.7 to 1.85X1.1 mm);
Off Ilha de Luandal20 m, 40 shells (1.55x0.9
tol.05X0.65 mm); Off Mussulo 90-100 m, 80 shells
(1.1X0.75 to 2.0X1.2 mm) ; Pnda Amelia 40-60 m, 4
shells (1.4X0.9 to 1.6X 1.0 mm); Off Ponta Albina 40 m,
1 shell (1.2X0.75 mm).

Habitat: On nniddv bottoms of otiter shelf

Distribution: Mauritania and Senegal; Gulf of Guin-
ea; Northern and Southern Angola.

Etymology: The specific name alludes to the shell
sculpture with sinuous axial ribs.

Remarks: This species somewhat resembles the Eu-
ropean species Alvania testae (Aradas and Maggiore,

Page 86

THE NAUTILUS, Vol. 113, No. 3

Figures 21-24. Ahania marloi new species. 21. Shell from oft Anihrizete (Angola), shell length 2 9 iiini 22. Holotype, from off
Ilha (Ic Luanda (Angola), shell length 2.7.5 mm 23. Protoconch of the holotyi^e Scale bar = 100 |im 24. Specimen from off
Salonin (Senegal), shell length 2. .5 mm.

Figures 2.5-26. Aliaiiin liispirhil/i Monterosalo, 1884. 25. SpecinK'n Iroiii oil San N'ilo (Sicily), coll. Montcrosalo, shi-II Icn0h
.3.25 mm. 26. Protoconch ol the sairie specimen Scale hur = 100 (Jim.

1844) l)ecaii.sc' of its sinuous rihs and vcw opisthoclinc
outer lip. It also occupies a siniilar liahilat on mmkKK oil
shore holtoins. It dilfcrs in lia\'ing ( I ) thinner and smaller
shell, and (2) larger protoconch that lornis a more olitnse
apical angle, lacking ohlifjne threads on adapieal pari ol
the whorls (see Bouchct and WareTi, 1993: ng. 1387, lor
an illustration of the protoconch of A. tcslac).

The MioceTie lossil sjiecies Alvtiiiia niuliui (>'ossniaini
and Peyrot, 1918, Ironi the A(|uitaine Basin, is \'ei-\ sim-
ilar i,s size, general outline, and hahilat nn the niiidcK
outer shelf. It shares with A. Jlcxilis the liK-oconeli
sculpture with 2 stronger spiral cords, respectively ada]i-
ically and ahapically from tlie suture. The protoconch is
nevertheless nnich smaller, with 1 les.s whorl,

Aluinia retina new species
(Figures 33-3.5)

Description: Slu'll high conical, solid, adnlls
2.0.5X1. .3.'5 mm to 2.7x|.fi mm. Protoconch with I.I

1.5 convex whorls, globose; whorls apparently smooth,
with Tuinute and scarce gramik's visible onlv under high
magnification (scamiing electron microscopv). Teleo-
coneli with 2.9-3.2 whorls; spiri' whorls conve.\, with sin-
uous axial folds (ca. 18-22 on penultimate whorl) ;m(l
sjiiral colds much narrower than inlersp;iees; intersec-
lions bhmt; most i-ords indistinct on first teleoiDUch
whorl, except for I subsulural and I suprasutural which
will remain stronger throughout subsequent whorls;
;idapical one (juite separated from sninrc Penultimate
wlioil with fi-7 spiral cords Hods whorl rounded, with
Ki 18 s]Mral cords and folds gradually lading towards
pcrnnnbilical area; snbsutural cord stronger, others
(•(|ii,il in size. Fine mieroseulpture of spiral thri'ads on
leleoconeh whorls. Outer lip slightly opisthoclinc, exter-
nalK thickened by strong rim that is strongK' differen-
lialcd from adjacent extiMiial shell surface; superim-
posed 1)\- spiral cords which coiiliiiue, \rr\ atleiiiiated,
on its outer surhice; inner sinlaee siiioiilli. Inner lip

S. Gofas, 1999

Page 87

Figures 27-32. Alvdnia jlcxilis iii-w sjx-ncs, troiii Angohi, 27-28. Shell (liololypej fioiii oil lllia df Luanda, shell leij'^h \\i nun,
29. Protoeonchofthe holotype. Scale bar = 100 |xni. UO-.'Jl. Shells Iron] off Anilirizefe. shell leiiijth 1.3 and 1 .8 nun, .'}2. Protoeoneh
of ihe speeirneii fig. 31, Scale t)ar = 100 \i.in.

Fijiurc's .■{.■J-.3.5. Alvania ret:,iiui new .species, lioni Seneffil, .TJ. Shell (hololype) from oil SVV oi Cup Manuel, 250 rn, shell lenad,
2,7 mm. .'54. Protoconch of the liolotyjie. Scale har - 100 |xin, .'5.5. Delail ol die microsculplure on a teleoconch wliod of die
holotype. Scale bar = 100 (J,m.

slif^htly thickened, leaving a distinct iniiiiijical chink.
Slieil white.

Type material: llulotyix- (.shell 2.7X1,6 mm) and 5
paratypes (shelLs 2. 05X1. .35 to 2.45 XL. 55 mm) from
type kjcality, eoU, Marche-Marchad (MNHN).

Type locality: SW of Cap Manuel (Senegal), 250 m.

Habitat: On niiiddv liolfoins c)l iijipcr slope.

Distribution: Known on!\- from Ispc locality.

Etymology: The specific name alludes to the pattern
of snhsiiturai knobs, which resemlile a (jueen's crown.

Remarks: 'fliis species shares with A. jiexilis the .sculp-

Page 88

THE NAUTILUS, Vol. 113, No. 3

ture of sinuous, shaip a\ial ribs, spiral cords that become
stronger abapicallv from suture, and outer lip smooth
internally. It is clearly difterentiated by its larger size,
outer lip thickened externally, spiral microsculpture, and
protoconch with 1 less whorl. It does not resemble anv
of the knowm Western European species.

Alvania c^ofasi (Rolan and Femandes, 1990)
(Figures 36-39)

Manzonin gofasi Rolan and Femandes, 1990, p. 6.'3-64, pi. 1,
figs. 1^3.

Description: Shell high conical, solid, adults 1.5X1
mm to 2.1X1.3 mm. Protoconch with 2 convex whorls,
first one slightly flattened on top; surface of larval whorls
with pair of spiral lines, formed by slightly oblique seg-
ments, at some distance from adapical suture, and clus-
ter of other 2-3 such Unes near abapical suture. Teleo-
conch with 2.2-2.8 whorls; spire whorls very convex,
with strong, widely spaced axial folds (ca. 12-14 on pen-
ultimate whorl) and strong spiral cords; intersections
pointed; 3 cords on all spire whorls; adapical cord situ-
ated at some distance from suture forming shoulder Spi-
ral microsculpture of fine striae (ca. 40 between a pair
of cords) visible under SEM, evenly set on cords and
interspaces, and superimposed on axial folds. Bodv*
whorl rounded, with 6 strong spiral cords and faint sev-
enth cord just bordering umbilical chink; axial folds
gradually fading toward periumliilical area; fifth cord
separated from sixth one b\ slightK broader and deeper
interspace, sixth cord smooth. Outer lip verv slightlv
opisthochne, externally thickened by very strong rim that
is strongly differentiated from adjacent external shell
surface, terminating with clearly differentiated flat sur-
face on apertural plane; superimposed externallv by spi-
ral cords that terminate abruptly at edge of that surface;
inner surface of aperture smooth, clearly chfferentiated
from apertural edge. Inner lip slightlv thickened, nar-
rowing towards small uniiiilical chink.

Shell color whitish, with p;ile browmish liand on mid-
dle part of spire whorls and another one on periumbilical
area, not continued on aperture.

Type material: Holotvpe (not seen) in MNCN, Ma-
drid.

Type locality: Off Ilha de Luanda (Angola), 50-100

Material examined: Senegal (coll. Marche-VIarchad):
Off Goree 95 m, I specimen + 1 shell (1.8X1.2 to
2.1X1.3 mm); SVV of Cap Manuel, 2 shells. Angola (coll.
Gofas): Ambrizete 45 m, 7 shells (1.45X0.95 to
1.75X1.1 mm); Ambrizete 80 m, 3 shells (1.7X1.05 mm
to 1.4X0.9 mm); Off Ilha de Luanda 40-60 m, 9 shells
(1.5X0.95 to 2X1.2 mm); Off Ilha de Luanda 120 m, 9
shells (1.75X1.15 mm to 1.45X1 mm); Off Mussulo
(Macoco) 50-70 m, 4 shells (1.75X1.15 to 1.9X1.25
mm); Off Mussulo 90-100 m, 2 specimens + 20 shells
(1.6X1 mm to 2X1.25 mm).

Habitat: On rock^v areas of the outer shelf

Di.stribution: Only kniowni from Senegal and from
Northern Angola.

Remarks: This species is very similar to the European
A. zcilamlica (Montagu, 1815) and shares v\ith it the
sculpture, microsculpture, and the pecuBar construction
of the aperture v\dth a flat, beveled edge. This latter
structure is not present in any other European of West
African rissoid and seems unique. I agree with Mool-
enbeek and Faber (1987) that this species should not be
allocated to the genus Manzonia Brusina, 1870 (type
species by original designation: Turbo costatus Adams,
1797 = T. crassus Kanmacher, 1798), and that Manzonia
should include only species having pitted spiral micro-
sculpture.

Alvania fariai (Rolan and Femandes, 1990)
(Figures 40-42)

Manzonin fariai Rolan and Femandes, 1990: 64—65, pi. 1 figs.
4-6.

Description: Shell conical, moderatelv sohd, adults
2.15X1.6 mm to 2.55x1.85 mm. Protoconch conical
with 2-2.2 convex, laterally compressed whorls; proto-
conch whorls with 7-8 verv fine spiral threads that tend
to break into small, slightlv oblique segments. Teleo-
conch with 2.2-2.8 whorls, with sculpture of blunt spiral
cords superimposed on broad, sinuous axial ribs; 4 cords
on first teleoconch whorl, 5 on penultimate, 12-13 on
bod\ whorl. Axial ribs (ca. 15-16 on body whorl) grad-
ually fading on periumbihcal area, where spiral cords are
broader and flatter. Outer Bp very sUghtlv opisthochne
and sinuous, thickened externally by prominent rim,
raised to form sort of crest in its adapical part; super-
imposed b\ spiral cords which terminate abiiiptlv at
edge of the inner surface; most abapical spiral cords of
body whorl also extending on outer Hp; inner surface
smooth and not thickened, clearly differentiated from
outer surface of apertural rim. Inner lip rather narrow,
bordering tiny umbifical chink. Shell color white.

Tjpe material: Holot\pe (not seen) in MNCN, Ma-
drid.

Type locality: Off Ilha de Luanda (Angola), 100 m.

Material examined: Senegal (coll. Marche-Marchad):
SW of Cap Manuel 250 m, 12 shells (2.15x1.65 to
2.55x1. ,S5 mm); SW of Goree 150-250 m, 1 shell
(2.25X1.65 mm). Off Goree sta. 56-1-lOA, 150-200 m,
1 shell (2.5X1.85 mm). Angola (coll. Gofas): Off Am-
brizete 60 m, 1 shell + 2 fragments (2.2X1.6 mm); Off
Ambrizete 80 m, 5 shells (2.2X1.65 to 2.45X1.75 mm);
Off Ilha de Luanda 40-60 m, 1 shell (2.35X1.65 mm);
Off Ilha de Luanda 120 m, 3 shells (2.3X1.7 to 2.35 X 1.7
mm); Off Mussulo (Macoco) 50-70 m, 2 shells
(2.25X1.7 to 2.35x1.7 mm).

Habitat: On the outer shelf and upper slope, tvpe of
bottom unknown.

S. Gofas, 1999

Paee 89

Figures 36-39. Akaiiia ^ojasi (R(il;in and Fenuuides, 1990), from Angola. 36-37. Shells from off Ambrizete, shell length 1.7
nun and 1.55 mm, 38. Protoconch of the specimen fig, 37, Scale bar = 100 (jliii. 39. Detail of the microsculpture on a teleoconch
whorl ot the same specimen. Scale bar = 50 fjim.

Figures 40^2. Alvdiiia fniiai (Rolan and Femandes, 1990), from Angola, 40. Shell h-oiii off Ambrizete, shell lengtli 2,2 mm,
41. Protoconch of the specimen fig. 40. Scale bar = 100 jjim 42. Detail of the micro.scnlpture on a teleoconch whorl of the same
specimen. Scale bar = 20 (jlhi.

Distribution: Senegal and northern Angola.

Remarks: This species is not closely related to anv
other extant species of the Rissoidae, hut resembles an
undescribed species from deep-water Oiigocene depos-
its of SW France (Lozouet, pars, comm.), with which it
shares the peculiar arrangement of the outer lip and the
rather cylindrical protoconch whorls. It does not have
the punctate spiral sculpture of Manzonia.

Alvania marcluuli new species
(Figures 43—47)

Description: Shell higli to verv high conical, solid,
adults 2.25x1.25 mm to 2.95X1.5 mm. Protoconch with

l.'3-1.5 convex whorls, globose; with 7 spiral Hat cords,
as broad as interspaces, formed b\- minute granules that
can be distinguishetl onlv under high magnification.
Teleoconch with .3. .3—1.2 whorls; spire whorls convex,
with strong, widely spaced, nearK" vertical axial folds (ca.
12-14 on penultimate whorl) and strong spiral cords,
narrower than interspaces; intersections pointed; 2 cords
on first and second teleoconch whorl, third one inserted
between suture and most adapical of preexisting cords
on second teleoconch whorl, progressivelv increasing in
strength. BocK' wliorl rounded, v\ith 8 spiral cords and
axial folds gradually fading towards periumbilical area;
spiral cords most developed near periphery 2 cords bor-
dering inner lip faint and coalescent. Teleoconch whorls
with fine spiral microsculpture of wrinkled threads. Out-

Page 90

THE NAUTILUS, Vol. 113, No. 3

S. Gofas, 1999

Page 91

er lip slightly opisthocline, externally thickened h\ veiy
strong rim stn)ngl\' differentiated from adjacent external
shell sinface; superimposed In' spiral corils that contin-
ue, verv attenuated, onto edge of lip; inner surface
smooth. Inner hp slightlv thickened, loosely appressed
on columella. Shell color whitish.

Type material: Holotype (shell 2.7X1.4 mm) and 9
parat\pes (shells 2.2X1. .35 to 2.95X1.45 mm) from tvpe
locahh-, coll. Marche-Marchad. 20 Feb. 1956 (MNHN).
Parat^pes (3 shells 2.5X1.4 to 3. 0X1. ,55 mm) from t>qie
locality (AMS).

Type locality: SW of Cap Manuel, Dakar area (Sen-
egal), 80-250 m.

Material examined: Morocco: Off Sich Ifni, "Cineca
3" sta. B33 (29°23'N, 10°50'W), 132 m, 34 shells
(2.25X1.3 to 2.S5X1.45 mm), coll. Glemarec 1972.
Mauritania: -Nchago" sta. 01 (20°1S'N, 17°40'W), 185
m, 1 shell (2.5X1.5 mm), coll. Richer de Forges 1981.
Senegal (coll. Marche-Marchad): Ty]3e material and Da-
kar area 170-200 m, 1 shell (2.5x1.5 mm); S. of Goree
110-112 m, 12 shells (2.25X1.25 to 2.75X1.45 mm); Off
Goree 150-200 m, sta. 56-1-lOA, 12 shells (2.4X1.,35
to 2.9X1.5 mm); Dakar (no details) 1 specimen
(2.25X1.25 mm).

Habitat: On deeper part of shelf and upper part ot
slope, bottom type unknown.

Distribution: From the Saharan coast to Senegal.

Etymology: The species is named after Igor Marche-
Marchad, who has collected most of the Senegalese ma-
terial studied herein and contributed much to the knowl-
edge of West African mollusks.

Remarks: This species resembles the Mediterranean
species A. sitbareolata (Monterosato, 1869) (figures 48-
49 herein, specimen collected near the type localit)')-
The main differences are that A. marchadi is larger, the
spiral microsculptin^e is better defined, there is an ad-
ditional cord on the broad shoulder of bodv whorl, the
granules on protoconch are more clearly organized to
form spiral cords, and it is whitish instead of brf)wn or
banded.

The length-width ratio is apparently suiprisingly vari-

able, even within the same population, but this is not
correlated with an\' (ither character which woukl suggest
presence of more than 1 species.

Ahania richcri new species
(Figures 50-52)

Description: Shell high conical, solid, adults 2.1X1.2
mm to 2.7X1,55 mm. Protoconch with 1.3-1,5 convex
whorls, globose; surface of larval whorls with minute
granules that can be distinguished onl\' under high mag-
nification; some of them looselv aligned spiralis' or
obliquelv. Teleoconch with 3.3-4.0 whorls; spire wh(jrls
convex, with widely spaced, nearly vertical axial folds (ca.
15 on penultiniate whorl) and spiral cords narrower than
interspaces; intersections bluntK pointed; 2 cords on
first and second teleoconch whoil, ailapical one quite
separated from suture; adapical edge of whorls along su-
ture somewhat swollen. Bodv whorl rounded, with 6 spi-
ral cords and axial folds gradualK' fading towards peri-
umbilical area; spiral cords near periphen most de\el-
oped, 2 liordering inner lip quite taint and coalescent.
Teleoconch whorls with extremely fine spiral microsculp-
ture of granulose threads. Outer lip slightK opisthocline,
externally thickened by \en' strong rim stronglv differ-
entiated from adjacent external shell surface; superini-
posed by spiral cords that continue, ver\ attenuated, on
its outer surface; inner surface smooth. Inner lip slightK
thickened, lea\ing a distinct umbilical chink. Shell color
whitish.

Type material: Ilolot\pe (shell 2.7()X 1.45 nnn) and 8
paratypes (shells 2.1X1.2 to 2.7X1.45 mm) from the
tyjDe localitv, coll. Marche-Marchad, 20 Feb. 1956. Par-
at\pes (3 shells 2.20X1.25 to 2.7X1.4 mm) from type
locality (AMS).

Type locality: SW of Cap Manuel, Dakar area (Sen-
egal), 80-250 m.

Material examined: Mauritania: "Nthago" sta. 243
(17°48'N, 16°32'W), 200 m, 1 shell (2.4X1.35 mm), coll.
Richer de Forges 1981. Senegal: Tvpe material and S.
of Goree 110-112 m, 3 shells (2.25X1.3 to 2.35X1.35
mm), coll. Marche-Marchad; South of Dakar (14°35'N,

Figures 43-47. Ahania marchadi new species. 43. Shell (holotvpe) from off SW of Cap Manuel (Senegal), shell lengtli 2,7 mm.

44. Shell (paratvpe), from the type localits', shell length 2.5 mm. 45. Protoconch of the paratope fig, 44. Scale bar = 100 |j.iii- 46.

Detail of the microscuipture on a teleoconch whorl of the same specimen Scale bar = .50 |xm. 47. Shell from off Sidi Itni (Morocco),

shell length 2.7 mm.

Figures 48^9. Akiinin suharcolata (Monterosato, 1869). 48. Specimen from off Linosa (SiciK cliannel), shell leii'^tli 1,9 nun,

49. Protoconch of the same specimen. Scale bar = 100 |jLm.

Figures .50-.52. Ahania richeri new species. 50. Holot\pe from off SW of Cap Manuel (Senegal), shell length 2.7 nnn. 51.

Protoconch of the holotype. Scale bar = 100 |xm. 52. Detail of the microscuipture on a teleoconcli whorl of the holotype. Scale

bar = .50 [i.m.

Figures .5.3-.56. Alvania coseli new species. 53. Shell (holotvpe) from off Dakar (Senegal), shell length 2.75 mm. 54. Parahpe

Inim the t\pe localitv, length 2. .3 nun. 55. Protoconch of the parat\pe fi^, 54, Scale bar = 100 jjim. 56. Detail of the microscuipture

on a teleoconch wliorl ot the same specimen. Scale bar = .50 (jlui.

Page 92

THE NAUTILUS, Vol. 113, No. 3

17°35'W), 120 ni, 1 shell (2.6X1.35 mm), coll. Leung
Tak.

Habitat: On deeper part of shelf and upper part of
slope, t\pe of bottom unknown.

Distribution: Mauritania and Senegal.

Remarks: This species is extremely similar to and lives
sympatrically with A. marcliadi, a species with which it
is easily confused. It is distinguished by absence of linear
arrangement of minute granules on protoconch sin-face,
vyhich appears smooth under stereomicroscope. The
teleoconch ot A. richcri can be tlifferentiated b)- the
more distinct umbihcal chink and by lack of an addi-
tional cord on shoulder of last whorls. It is also similar
to A. siibarcolata. Iiut differs in its larger size, stouter
profile with a swollen subsutural area, and blunt teleo-
conch sculpture.

The species is named after Bertrand Richer de Forg-
es, marine biologist at ORSTOM (the French goyern-
ment's overseas research agency) and collector ot most
of the Mauritanian material studied here.

Alvanio coscli new species
(Figures 53-56)

Type material: Holot>pe (shell 2.75X1.5 nun) and 7
paratypes (shells 2.3X1.4 to 2.8X1.55 mm), coll.
Marche-Marchad. Paratypes (3 shells, 2.1X1.2 to
2.25X1.3 mm) from off Goree 95 m, coll. Marche-Mar-
chad (AMS).

Type locality: Dakar area, 95 m.

Material examined: Morocco: Off Sidi Ifni, "Cineca"
sta. B33 (29°23'N, 10''50'W), 132 m, 3 shells (2.1X1.1
to 2.25X1.1 mm), coll. Glemarec 1972. Mauritania:
"N'Diago" sta. 118 (18°36", 16°31'W), 96 m, 1 shell
(2.35X1.3 mm); -N'Diago" sta. 119 (18°36', 16°28'W),
70 m, 4 shells (2.45X1..3 to 2.7x1.4 mm), coll. Richer
de Forges 1981. Senegal (coll. Marche-Marchad): Type
material and S of Goree 110-112 m, 14 shells (2.1X1.25
to 2.6X1.4 mm); Off Goree 50 m, 1 shell (2.2X1.3 mm);
Dakar area (14°32', 17°25'W), 50 m, 1 shell (2.25x1.3
mm); Off Goree, sta. 56-l-l()A, 150-200 m, 1 shell
(2.5X1.35 mm).

Description: Shell high, conical, sohd, adults 2.1 X 1.2
mm to 2.75X1.55 mm. Protoconch with 1.3-1.5 con\ex
whorls, globose; surface with 7 spiral flat cords, as broad
as interspaces, formed by minute granules visible onlv
under high magnification. Teleoconch with 4 whorls:
spire whorls convex, with widely spaced, nearly vertical
axial folds (ca. 16-18 on penultimate whorl) and spiral
cords narrower than interspaces, superimposed on axial
folds; 4 cords on first and second teleoconch whorl,
adapical one bordering suture. Body whorl rounded,
with 12-13 quite even spiral cords and axial folds grad-
ually fading toward periumbilical area. Teleoconch
whorls with extremely fine spiral microsculpture of v\'ide-

ly spaced granulose threads. Outer lip slightly opistho-
chne, externally thickened by very strong rim strongly
differentiated from adjacent external shell surface; su-
perimposed by spiral cords, which continue, very atten-
uated, on its outer surface: inner surface smooth. Inner
lip slightl) thickened, leaving distinct umbilical chink.
Shell white.

Habitat: On deeper part of shelf and upper part of
slope, type of bottom unknown, often found together
with A. marchadi and A. liclicii.

Distribution: From the coast of Sahara to Senegal.

Etymology: The species is named after Rudo \'on Co-
sel, as a triiiute to his contributions to West African mal-
acolog\.

Remarks: This species shares with A. siibarcolata, A.
marchadi, and A. richcri the general outline, \ertical ax-
ial ribs, teleoconch microsculpture and the \'ery pecuhar
microsculpture of closely arranged granules on the pro-
toconch. Based on these shell characters, these species
are provisionally considered to belong to a monophvletic
group. Alvania coscU is distinguished from these species
by having about tsvice as man\' spiral cords.

Genus Crisilla Monterosato. 1917

T\pe species: Turbo scmistrintus Montagu, 1808, b\- nionotvpy.

Crisilla transitoria new species
(Figures 57-61)

Description: Shell conical, slightly cvrtoconoid, mod-
erately solid, adults 1.40X0.85 'mm' to 2.25X1. .35 mm.
Protoconch with 2.3-2.7 whorls, first one depressed, giv-
ing spire slightly cyrtoconoid profile; with spiral lines of
tiny granules, well-defined on first whorl, becoming
looselv arranged on second whorl, again well-defined on
last lar\'al whorl. Teleoconch with 2.2-3.0 whorls; spire
whorls shghtK con\ex, with moderateK" dexeloped, tight-
ly arranged axial folds (ca. 35 on penultimate whorl) and
spiral cords; 2 subsutural and suprasutural cords stron-
ger; remaining (2 on first teleoconch whorl, 5 on pen-
ultimate) weak and not superimposed on axial folds.
Body whorl rounded, slightly constricted, with lS-20
spiral cords and axial folds gradualK fading toward peri-
umbilical area. Outer lip orthocline, smooth inner sur-
face, slightly thickened externally at some distance from
shaip edge. Inner lip thin, bordering tiny umbihcal
chink. Shell color buff, with 1 row of large, square sub-
sutural brown patches; 1 row of smaller suprasutural
brown patches, which continues on body whorl, and spi-
ral rows of smaller patches on periumbilical area.

Head-foot semitiansparent, whitish with triangular,
opaque yellow patches behind eyes, and internipted yel-
low bars on tentacles. Foot with distinct anterior pedal
gland and cluster of 3 small metapodial tentacles; 2
small, colorless pallial tentacles.

S. Gofas, 1999

Paee 93

Figure 57. Cri.silla transitoria new species. Living animal (paratype) from Palnieinnhas (Angola), shell length 1.8 mm.

Type material: Holotype (specimen l.Sxl.l mm)
and 5 parahpes (specimen 1.65X1.1 to 2.1X1.3 mm)
from t\pe localit\- (MNHN). Paratypes (6 specimens,
1.7X1.05 to 1.8X1.1 mm; lixing animal obsei-ved and
drawn) from Palmeirinhas, Buraco, rocks in 2-3 m. Par-
atypes (5 specimens 1.65X 1.1 to 1.9X1.2 mm) from type
locality (AMS). Paratypes (5 specimens 1.65X1.05 to
1.95X1.2 mm) from type localit)- (UAN).

Type locality: Corimba, province ol Luanda (Angola,
8°50'S, 13°09'E), on shell gravel. 10-20 m.

Material examined: Senegal (coll. Marche-Marchad):
Off Saloum 50 m, 3 shells (1.75X1.15 to 2.0X1.2 mm);
Baie de Goree, S. of "Tacoma" shipwreck, 25 m, 1 juv
shell; SW of Cap Manuel 50 m, 1 .shell (1.85X1.15 mm);
Casamance (12°46,9'N, 17°29,9'W), 1 shell (1.6X1.15
mm). Sao Tome: Morro Peixe, 1 shell (1.85X1.1 mm);
Mutamba near Neves, 1 shell (1.85X1.15 mm). Congo:
Pointe Noire, Plage mondaine N of lighthouse, 5 shells
(1.4X0.9 to 1.55X0.95 mm): Pointe Noire, Plage Or-
stom dredged 5-7 m, 1 specimen (1.8X1.15 mm), coll.
von Cosel, Nov.-Dec. 1985. Angola (coll. Gofas): Tvpe
material and off Ambrizete 45 m, 14 shells (1.4X0.95 to
1.5X1.0 mm): Off Ambrizete 80 m, 20 shells (1.35X0.95
to 1.6X1.0 mm); Ambrizete Lighthouse, shore, 8 shells
(1.5X0.9 to 1.9X1.2 mm); Barra do Dande, intertidal, 2
specimens + 3 shells (1.8X1.15 to 2.25X1.35 mm); Ilha
de Luanda, dredged 40-60 m, 15 specimens + 20 shells
(1.4X0.95 to 1.8X1.15 mm); Cabo Ledo, 10-40 m, rocks
from tangle nets, 100+ specimens (1 dr.), mostlv juve-

nile (1.75X1.15 mm to 1.50X1.00 mm); Off Mussulo
90-100 m, 15 specimens + 40 shells (1.3X0.9 to
2.2X1.25 mm); Chapeu Armado, intertidal, 30 speci-
mens + 3 shells (1.5X0.85 to 1.65x0.95 mm); Bai'a das
Pipas, intertidal, 1 shell (1.85X1.15 mm); Sao Nicolau,
shore, 400 shells (1.5x0.9 to 2.0X1.2 mm); Praia Ame-
lia, intertidal, 5 specimens (1.5x0.9 to 1.75X1 mm; liv-
ing animal observed and drawn); Praia Amelia 2-5 m,
23 specimens (1.4X0,9 to 1.65X0.95 mm); Praia Amelia
40-60 m, 1 specimen + 15 shells (1.1X0.75 to 1.5X1
mm); S. of Ponta Albina, 6 shells (1.7X1.1 nnn to
1.35x1 mm); Bai'a dos Tigres, low tide, 5 specimens +
3 ,shells (1.35X0.9 to 1.5x0.95 mm).

Habitat: Intertidal to outer shelf. On rock's' or shell
gravel bottom, in crevices percolated bv seawater, or in
coarse sediment.

Distribution: Senc"al; from Congo to southernmost
Angola: Sao Tome.

Etymology: The specific epithet alludes to the general
aspect of the shell, intermediate between typical Crisilla
and Alvania.

Remarks: This species shares a number of characters
with the tvpe species of Crisilla, C. scmistriata (Mon-
tagu, 1808). The two species show the same microsculp-
ture on the multispiral protoconch (see Bouchet and
Waren, 1993: fig. 1535, for a good illustration), same
color pattern of square blotches, and stronger subsutunil
cords and furrows differentiated from the remainder of

Pa^e 94

THE NAUTILUS, Vol. 113. No. 3

Figures 58-61. Crisilla transitoha new species, from Angola. 58. Parahpe, from Palmeirinhas, shell length 1.8 mm. 59. Holohpe,
from Corimba, shell length 1.8 mm. 60. Paratvpe, from Corimba, shell length 1.75 mm. 61. Protoconch of the parahpe fig. 60.
Scale bar = 100 |xm.

Figures 62-63. Crisilla fallax new species. 62. Holot\pe. from off Goree, (Senegal), shell length 2.0 mm, 63. Protoconch of a
paratope. Scale bar = 100 |j.m.

whorl. It also lives in the same Idnil of haliitat, in oxide-
lined cavities under rocks that are not filled with sedi-
ment and through which seawater percolates. Crisilla
transitoria is distinguished b\- a more cvrtoconoid pro-
file, a lower spire, more definite axial fcjlds, and presence
of yellow stains on tentacles (colorless in C. sonisiriata).

Crisilla fallax new species
(Figures 62-63)

Description: Shell conical, moderately solid, adults
1.6X1.15 mm to 2.25X1.4 mm. Protoconch with 1.3-
1.5 whorls, convex. v\ith sculpture of minute granules
arranged along spiral fines. Teleoconch with 3.0-3.2
whorls; spire whorls slightly convex, with moderately de-
veloped, tightly packed a.xial folds (ca. 35 on penultimate
whorl) and spiral cords; 2 subsutural and 1 suprasutural

cords stronger; remaining cords (2 on first teleoconch
whorl, 5 on penultimate) weak and not superimposed on
a.xial folds. Body whorl rounded, with 18-20 spiral cords
and axial folds gradually fading towards periumbilical
area. Outer lip orthocline. smooth inner smiace. hardly
thickened extemalK" at some distance from edge. \v'hich
is cutting. Inner lip thin, bordering tin\' umbilical chink.
Shell color buff, with 1 row of large, square subsutural
brown patches, 1 row of smaller suprasutiual brown
patches, which continues on body whorl, aiul spiral rows
of smaller patches on periumbilical area.

T\pe material: Holotype (specimen 2.0X1.2 mm)
and 14 parat^pes (2 specimens 1.9X1.25 to 2.1X1.25
mm and 12 shells 1.6X1.15 to 2.05X1.25 mm) from type
locality-, coll. Marche-Marchad (MNHN).

T^pe locality: S of Goree. Senegal, 110-112 m.

S. Gofas. 1999

Page 95

Figure 64. Benthonellania gofasi Lozouet, 1990, Imng spec-
imen from off Miissulo (Angola), .shell leniith 2.0 mm.

Material examined: Senegal: T\pe material and Da-
kar area, 95 ni, 1 .shell (2.25X1.4 mm), coll. Marche-
Marchad.

Habitat: Outer shell, t\pe of bottom unknowai.

Distribution: Known only from near type localitv; off
Senegal.

Etymology: The species name alludes to possible con-
fusion with the other West African Crisilhi.

Remarks: This species is veiy similar to C. transitoiia,
sharing with this latter the teleoconch sculpture and col-
or; they differ, however, hv the spire profile, which is
higher and more regular!)' conical in C. tiansitoiia, not
cyrtoconoid, and by the number of teleoconch whorls,
one more in C. tronsitoria. The essential diagnostic char-
acter is the paucispiral protoconch, which hints at one
more case of a pair of similar species, with one species
supposedly undergoing planktotrophic development, the
other one undergoing non-plankiotrophic de\ek)pment.

Genus Benthonellania Lozouet, 1990

Tvpe species: Bcnthonelliniia '^ofasi Lozonet, 1990. by original
designation.

Benthonellania gofasi Lozouet, 1990
(Figures 64-68)

BcnthoHcUanm gofasi Lozouet. 1990. p, :314-.318, 322-.324.

Description: Shell elongate, conical, slightly cyrtoco-
noid, rather thin, adults 2X1.2 to 2.8X1.6 mm. Proto-

conch with 2.1-2.4 con\ex whorls, with irregular spiral
threads from which short ol>li([ue lines branch-off, form-
ing loosely reticulate pattern; last protoconch whorl with
broad smooth spiral zones. Teleoconch with .3-3.5
whorls; spire whorls rather flat with swollen subsutural
rim and strong, slightly sinuous a\ial folds (ca. 20 on first
teleoconch v\'horl, 10-12 on penultimate whorl) termi-
nating adapically by bulge on subsutural rim. Body whorl
rounded, with a.\ial folds fading toward periumbilical
area, and gradualK replaced there bv blunt spiral rims.
Outer lip not thickened, orthocline. Inner lip thin, leav-
ing small umbilical chink.

Head-foot semitransparent, whitish with tinv opaque
white flecks, and poorlv defined a.\ial opaque white bar
on tentacles. Distinct neck lobe on right side behveen
head and opercular lobe, not present on left side. Snout
rather small and cylindrical, with yellow buccal mass vis-
ible by transparency. Foot with broad and flat metapo-
dium and ven' conspicuous, opaque white posterior ped-
al gland antl 1 cluster of small white flecks on propodium
sole, and another such cluster on metapodiuni; no me-
tapodial tentacles. Pallial tentacles slender, colorless,
with parallel sides, right one quite conspicuous, left one
very small.

Type material: llolot^pe (specimen 2.15X1.2 mm)
and 20 paratypes (MNHN); 2 paratypes (AMS).

Type locality: SVV of Cap Manuel, Senegal, 250 m.

Material examined: Mauritania (coll. Richer de
Forges 1981): "N'Diago" .sta. 222 (17°42'N, 16°34'W),
200 m, 1 specimen + 1 .shefl (2.2X 1.25 to 2.5X 1.4 mm);
"N'diago" sta. 243 (17°4S' N, 16°32'W), 200 m, 12 shells
(2.0X1.25 to 2.3X1.35 mm); "N'Diago" sta. 244
(17°54'N, 16°32'W), 1 specimen; "Ernst ^Haeckel" sta.
CH26S, (18°46'N, 16°51'W), 610 m, 7 shells (1.55X1.0
to 1.7X1.05 mm). Senegal: Off M'Bour, 246 m. 130
specimens (2.2X1.35 to 2.8X1.6 mm), coH. Pin; Dakar
area sta. 55-6-3D (14°19'N, 17°23'W), 78 m, 4 speci-
mens, coll. Marche-Marchad; Off Goree 80-250 m, 9
shells, coll. Marche-Marchad; South of Dakar sta. L4
(14°35'N, 17°35'W), 120 m, 4 shells, (1.9X1.2 to
2.1X1.25 mm), cofl. Leung Tak. Angola: Off Ilha de Lu-
anda, 120 m, 140 specimens (mostly juveniles, adults
1.65X1.0 to 2.05X1.25 mm), cofl. Gofas.

Habitat: On mudd\ bottoms of lower shelf and upper
slope.

Distribution: From off Essaouira, Southern Morocco
(Monaco sta. 1116 citeil bv Bouchet and Waren, 1993)
to Angola. Bouchet and Waren (1993) mentioned its oc-
currence off Madeira, but did not refer to an\ record in
particular

Genus Ohtusella Cossmann, 1921 ex-Monterosato ms.
(new name for Cingiilina Monterosato, 1884,
preoccupied bv Cingiilina H. and \. Adams, 1860)

T\pe species: Rissoa ohttisa Cantraine, 1842 (preoccupied h\
R ohfiisti Brown. 1841; = Risson intcrsccta Wood, 18.57).

Page 96

THE NAUTILUS, Vol. 113, No. 3

Figures 65-68. Benthoncllania gofasi Lozouet, 1990. 65. Holotvpe, from off Cap Manuel (Senegal), shell length 2.15 mm. 66-
67. Specimen irom oft Mussulo (Angola), same as fig. 64, shell length 2.0 mm. 68. Protoconch of a juvenile specimen from the
same locality-. Scale bar = 100 (jim.

Ohtusclla intersecta (Wood, 1857)
(Figure 11)

Rissoa obtum Cantraine, 1842: 34S (preoccupied).

Risson solutn (not of Philippi, 1844) sensu Forbes and Hanle\',

1850: 131, pi. 75 fig. .3-4.
Rissoa intersecta Wood S.\'.. 1857: 318 (based on Rissoa sohita

sensu Forbes and Hanley 1850).
Putilla (Obtiisella) cantrainei Nordsieck, 1972, replacement

name for R. obtusn Cantraine, 1842).

Tjpe locality: "Briti.sh".

Material examined: Western Europe and Morocco:
several hundred specimens from throughout the range.
Senegal: Off Saloum 50 ni. 4 shells (0.9X0.7 to 1.15x6.8
mm), coll. Marche-Marchad. Angola (coll. Gofas): Am-
brizete 45 m, 6 shells (0.6X0.5 to 0.8X0.6 mm); Am-
brizete 80 m, 40 shells (0.8X0.6 to 1.1X0.7 mm); Off
Ilha de Luanda, dredged 40-60 m, 5 specimens + 44
shells (0.6X0.5 to 0.85X0.7 mm); Off Ilha de Luanda
120 m, 40 shells (0.6X0.5 to 0.8X0.65 mm); Off Mus-
sulo 90 m, 150 shells (0.75x0.6 to 1.05X0.75 mm); Off
Mussulo 50-70 m (Macoco), 40 shells (0.70X0.60 to
0.90X0.70 mm). Off Praia Amelia, 40-60 m, 20 shells
(0.85X0.7 to 1.0X0.75 mm).

Description: Shell conical, somewhat tyrtoconoid;
adults 0.8X0.6 to 1.15X0.8 mm. Protoconch of 2 whorls,
apical one depressed, regularly convex with spiral thread
adapically next to suture. Teleoconch of 1.7 to 2 whorls;
spire whorls quite convex, regularly increasing in size,
sculptured with faint, uneven spiral cords (9-12 on pen-

ultimate whorl); body whorl rounded with spiral lines
continued all over. Aperture pvriform; outer lip ortho-
chne, smooth inner surface, not thickened. Inner hp
thin, bordering narrow imibilicus. Shell color buff, uni-
form.

Habitat: On soft bottoms (muddy sand or mud) of
continental shelf, usually between 50 and 200 m.

Distribution: From Northern Noru'ay, the Faroes,
southwestern and southern Iceland, to Mediterranean
(Bouchet anil Waren, 1993), and in West Africa south to
southernmost Angola at least.

Remarks: The range of this species is suq^risingly
large, but there are no moi']:)hological or ecological
grounds on which the West African populations cotdd
be distinguished.

SubfamiK Rissoininae Stimpson, 1865

Genus Rissaina il'Orliignw 1840

Tvpe species: Rissoina inca d'Orbigny, 1840, by original des-
ignation.

Rixsoina piiiictostriata (Talavera, 1975)
(Figures 69-73)

Zihinii jiuiirltKtriata Tala\'era, 1975: 3; pi, 1 fig, I; pi. 4 fig. 7.

Description: Shell elongate, conical, sohd, adults

S. Gotas, 1999

Paee 97

Figure 69. Rissoina punctostriata (Talavera, 1975), living specimen from Caotinha (Angola), shell length 7.45

5.2X2.1 to 8.9X,3.5 mm. Protoconch with 2.8-3.2 con-
vex whorls, smooth with spiral thread runninsi adapicallv
along suture of first and second whorl; protoconcli/teleo-
conch transition well-defined, with very deep adapical
notch bordered bv thickened rim (indicating position of
velar lobes during \'eliger lanal stage). Teleoconch with
6-7 whorls, first 2 with strong shoulder, next one becom-
ing gradually slightl)' convex; body whorl rounded abapi-
cally. Surface with oblique axial ribs (ca. 30 on penulti-
mate whorl, becoming more numerous but less distinct
on bodv whorl), and composite spiral niicrosculpture ot
uneven spiral threads superimposed bv spiral micro-
striae.

Aperture pyriform, deeply channelled at insertion of
outer lip on pre\'ious whorl. Outer lip opistliocline,
thickened lint not demarcated hom body whorl; nmer
surface smooth, meeting external sculptured surface
along sharp hne. Inner lip slightlv thickened, appressed.

Head-foot colorless, with faint brownish hue at tip of
snout. Tentacles with parallel sides, slender; eves at base
oi tentacles. Snout elongate, markedl)' with two lobes,
with pink buccal mass visible by transparency. Metapo-
dial tentacle hardlv visible. Sole of foot colorless, with
pedal gland inconspicuous. One small, slender pallial
tentacle on right side of aperttire; 2 contiguous tentacles
on left side, of which innermost is smaller and stouter.

Type material (not seen): Holotyjie, shell from SAH-
MAS 1 cruise, sta. EO 8, now in Museo de Ciencias
Naturales, Tenerife, TFMC 317 (MO) (F.G. Talavera,
pers. comm. 1997).

Type locality: Off Mauritania (18°2()'N, 16°1()'W),
70 m.

Material examined: Senegal: Baie de Goree, 20-.30

m, 4 shells (8.3X3.4 to 8.9X3.5 mm), coll. Pin. Cote
d'lvoire: 22 shells (4.25x1.65 to 4.65X1.9 mm). Sao
Tome: Esprainha, 0-4 m, 2 shells (4.85X2.05 to 6.5X2.6
mm). Gabon: Cap Santa Clara 1-2 m, 3 specimens + 1
shell (7.75X3.2 mm). Cameroun: sta. CC40, (4°04,5 N,
8°42'E), 61 m, 1 .shell (5.75x2.45 mm) coll. Monteillet.
Angola (coll. Gofas): Off Ambrizete 80 m, 4 juvenile
shells; Corimba, rocks with sand 0-1 m, 3 shells
(7.4X3.05 to 8.5X3.35); Corimba on sand bar 10-20 m,
10 .shells (6.00X2.45 to 6.75X2.8 mm). Palmeirinhas
(Buraco), 0-2 m, 1 shell (5.25X2.15 mm); Palmeirinhas
30 m, rocks, 1 shell (5.4x2.3 mm); Off Mussulo 90-100
m, 4 juvenile shells; Caotinha, 1 specimen (7.45X3.05
mm; living animal obseived and drawn); Santa Maria, 2
shells (7.25X3.1 to 8.65X3.6 mm); Lucira (Cesar), 10
m, 1 shell (7.25X3.15 mm); Chapeu Armado, 1 speci-
men (5.55X3.6 mm, broken apex).

Habitat: Under stones among coarse sand and rubble
on rock-v bottoms, subtidal.

Distribution: West African coast from Western Sa-
hara to Angola; Sao Tome; Cape Verde Islands.

Remarks: This species was described from a West Af-
rican specimen as Zcbina punctostriata by Talavera
(1975), but was later (Garcia-Talavera, 1982) svnony-
mized with the West Indian species Ri.s.soina dvcussata
(Montagu, 1803).

The multispiral lanal shell, with deep notches for ve-
lar lobes, indicates planktotrophic development for both
species. However, it can be questioned whether the
small laivae o{ Rissoina can routinely cross the Atlantic.
R. dccussata has a more cylindrical protoconch than R.
punctostriata from Angola or Cape Verde Islands, and
lacks the shoulder on the very early teleoconch whorls

Page 98

THE NAUTILUS, Vol. 113. No. 3

Figures 70-73. Rissoina pimctostriata (Tala\era, 197.5). 70-71. Specimen from Caotinha (Angola), .same as fig. 69. shell length
7.4.5 mm. 72. Protoconch of a juvenile specimen from Caotinha. Scale bar = 100 (jim. 73. Detail ot the microsculpture on a
teleoconch whorl, same specimen as figs 70-71. Scale bar = 50 (j.m.

Figures 74-77. Sclncartziella africana (Dautzenberg. 1913). 74. Specimen from Dakar (Senegal), shell length 3.2 mm. 75.
Protoconch of another specimen from the same locality. Scale bar = 100 jjim. 76-77. Details of microsculpture of the teleoconch
whorls on unaltered (76) and on partially eroded (77) surfaces, same specimen as fig. 74. Scale bars = 10 fjini.

(A.A. Luque, pers. comm. 1999): these moiphological
differences support holding them as separate species.

The distribution of R. piinctostriata is spott\" and the
species is usuall\ rare, a pattern of occurrence which is
usual in species featuring long-ranging planktotrophic
larvae. The population e.xamined from Cote d'lvoire
consists of old, worn shells, possibly subfossil, and differs
markedly in having strong axial sculpture, with few
strong ribs, thus superficially resembling SchwciiizicIIa
africana. The protoconch is similar to that illustrated in
fie. 72.

Genus Schwaiiziella Ne\ill. 1S81

T)pe species: Rissoina oricntalis Nevill. 1881 ( =
triticea Pease. 1861), by onginal designation.

■f* Rifi

Schwaiiziella africana (Dautzenberg, 1913)
(Figures 74-77)

Rissoina africana Dautzenberg, 1913: 48—49, pi. 2 fig. 5-6.
Rissoina africana \ar crassior Dautzenberg, 1913: 49, pi. 2 fig.

7-8.

Description: Shell \ery high conical, sohd, 2.5X1.1 to
5.25X2.15 mm, Protoconch with 1.1-1.3 convex whorls,
e\enly covered N\ith rounded smeared spots; proto-
conch/teleoconch transition well defined. Teleoconch
with 5-6.5 convex whorls; body whorl broadh' rounded
aliapicalK. Surface with strong, oblique, slightK sinuous
axial ribs (ca. 12 on bod\ whorl), and complex spiral
microsculpture of lines of pores and slits, or of irregular
striae when eroded (these latter representing underlying
stnicture). Aperture pNiiform, not channelled at inser-

S. Gotas, 1999

Page 99

Figures 78-80. Zchina robnsiior new species, from Senegal. 78-79. Sliell (|]iili)t\])e) from oil Oakai, sliell iengtii 5,1 mm, 80.

Protoconcli ot another specimen trom off SVV Goree. Scale bar = 100 jjim

Figures 81-82. Zebinn broivnuinn (d'Orbigny, 1842) from Santiago de Cuba, Eiisenaila ile Nispero, 81. Protoconcli, Scale bar

= 100 |jLm. 82. Detail of the microscuipture on a teleoconch whorl. Scale bar = 10 |j.m.

Figures 8.3-86. Zehiiui paivensis (Watson, 1873). 83-84. Specimen from Selvagens Islands, shell lengtli .3.9 nnii. 85. Protoconcli

o( a juvenile specimen from the same locality. Scale bar = 100 (J.m, 86. Detail ol the microstiilptiire on a teleoconch whorl Scale

bar = 10 (Jim, same specimen as figs 8.3-84.

tion of outer Hp on pre\ioiis whorl. Outer lip slightK'
opisthocline, thickened but not difterentiuted from body
whorl; inner surface smooth, meeting external sculp-
tured surface along shaq") line. Inner lip slightiv thick-
ened, appressed.

Type material: Figured synfvpe ol R africdna (shell,
.3.1 XI. .3.5 mm), here designated as lectotype (MNHN);
figured .syntype of R. ajricana var. crassior (3.0.5X1.4
mm), here designated as lectotype (MNHN).

Type locality: Pointe de Bel-Air (l)aie dc Ilaim), Da-
kar (Senegal), intertidal.

Material examined: Senegal: (Jtl Salonni .50 m, sta.
.55.3.9, 2 shells (4.9X2 to 5.6x1.9 mm); Dakar .5.563D,
3 shells (3.25X1.2 mm), coll. Marche Marcliad; Dakar,

10 shells (3.25X 1.2 to 3.45X 1.45 mm), Staadt collection,
MNHN; Joal, 10 shells (3.5X1.45 mm), coll. Mauny;
Yoff, intertidal under stones, 5 specimens (2. .55X1.15 to
2.85X1.25 mm), coll. Bouchet; Baie de Goree .50 m, 1
shell (5.25X2.15 mm), coll. Marche Marchad; Baie de
Goree .50 m, 1 shell (5.25X2.1 mm), coll. Marche Mar-
chad; Bel Air, towards "Tacoma" shipwreck, 2 shells
(3,15X1.3 to 3.45X1.4 mm); Dakar area, 32 specimens
widi nisU- crust (2.95X1.15 to 3.5X1,4 mm); Baie de
Goree 1.5 m, 15 specimens (3.15X1.4 to 3.4X1.45 mm),
coll. Marche Marchad; Dakar area (14°29'N, ]7°23'W),
21 m, 4 shells, ("ape Verde Islands: "Svlvana" sta. 136,
Maio island, 1 shell (3,45X1.45 mm).

Habitat: Intertidal or shallow subtidal, under riiiks.

Distribution: Known with certaintv onl\ Ironi a small

Page 100

THE NAUTILUS, Vol. 113, No. 3

stretch of coa,stline around Dakar, Senegal. The record
from Cape Verde Island.s i.s probaliK' hased on a misla-
beled specimen, as this species was not met with in a
large material of Rissoina from this archipelago exam-
ined by A. A. Luque and E. Rolan (pers. comm., 1999).

Genus Zcbina H. and A. Adams, 1854

Type species: Rissoina semiglabrata A.Adanis, 1854 ( = ? Ris-
soa tridentata Michaud, 18.30), bv subsequent designation,
Rehder, 1980.

Zcbina rohustior new species (Figures 78-SO)

Description: Shell elongate, conical, solid, glos.sy,
4.1X1.9 to 5.3X2.4 mm. p'rotoconch wth 1.2 whorls,
very globose, smooth. Teleoconch with 6 whorls, spire
whorls striiight, bodv whorl rounded abapicalK'. Surfoce
apparently smooth, udth scarce tiny punctures \asible
only under very high magnification. Aperture pyriform,
channelled at insertion of outer lip on previous whorl.
Outer hp veiy strongly opisthocline, thickened but not
differentiated from bodv whorl, with rounded edge. In-
ner surface of outer lip with 2 shghtK' swollen sections
separated by central depression. Inner lip strongK thick-
ened, appressed.

Type material: Holotvpe (5.1X2.4 mm) from t\pe lo-
calibv-, coll. Marche Marchad, sta. 55-6-3D (MNHN).

Type locality: Off Dakar, Senegal, 14°19'N, 17°23'W,
78 m.

Material examined: Morocco: "Vanneau" sta. 110
(30°23'N, 09°54'W), 110 m, 6 old shells. Mauritania:
"Ndiago" sta. 116, 210 m, 1 shell (4.85X2.15 mm), coll.
Richer de Forges 1981. Senegal (coll. Marche-Marchad):
SW Goree 150-200 m, 7 (4 juvenile) shells (4.1X1.9 to
4.15X2.0 mm); Off Goree 50 m, 2 shells; Off Goree 50
m, sta. 55-7-5A (14°32'N, 17°25,5'VV), 1 shell (5.3X2.4
mm); SW Cap Manuel, 2 juvenile shells.

Habitat: Continental shelf

Distribution: From Southern Morocco to Senegal.

Etymology: The specific epithet alludes to the sturdy
robust shell.

Remarks: This species resembles Zcbina paivcnsis
(Watson, 1873), a species commonly found on the shores
of the Canarv' Islands and Selvagens, and which authors
(Odhner, 1931; Nordsieck, 1972; Garcia-Talavera, 1982)
have identified as the West Indian Zcbina browniana
(d'Orbigny 1842) or Zcbina vitrea (C.B. Adams. 1850).
The Macaronesian populations resemble the Caribbean
ones, but differ in having paucispiral instead of multispi-
ral protoconchs, and in having different surface micros-
culpture. I illustrate here (Figs 81-82) these details from
a specimen of Z. browniana originating from the same
lot as the shell illustrated by Desjardin (1949: pi. 10 fig.
4).

Zcbina paivcnsis has been originally introduced by
Watson (1873: 364-365; pi. 36 fig. 29)' as Eulima paiv-
cnsis. and was overlooked since then. Its type locality is
the coast of the Selvagens, where it is extremely com-
mon. I figure (figures 83-86) some topotvpes for com-
parison.

Zcbina robustior differs from Z. paivcnsis in being
larger, having a strighter profile of spire and whorls and
a more globose apex. It is an outer shelf species, in con-
trast with Z. paivcnsis. which inhabits in beach gravel on
the lower intertidal zone of rocky shores.

DISCUSSION

There is apparentlv verv little in common between the
West African rissoids group and those from Western Eu-
rope and Morocco. The 2 shared species {PusiUina in-
conspictta and ObttiscUa intcrsccta) belong to genera in
which species-level systematics is still confused and
where further research may reveal species complexes
where 1 single taxon is now recognized.

The Rissoa and Sctia radiations, important in the
Mediterranean and still quite well represented in Mo-
rocco and in the Canaries, are \irtuall}^ absent in West
Africa. The only representation is the citation of Rissoa
similis Scacchi, 1836 in Port-Etienne (now Nouadhibou,
at the extreme north of the West African province) by
Dautzenberg (1913), which is correct but marginal to
the West African province. Rissoa and Sctia encompass
species that live on algae or marine phanerogams on
shallow rock"v bottoms. Some lineages in the genus Al-
vania. including species such as the Mediterranean A.
scabra Phihppi, 1836 and A. discors (Allan, 1818), which
usuall}' live among the photophilous algae, are also ap-
parently absent in West Africa. Most species described
herein live either in cavities of rock-v substrates, or on
outer shelf soft bottoms, and their European relatives,
where known, have also the same kind of habitat. This
trend mav be related to the scarcity of rockv shores suit-
able for the development of the communitv of photophi-
lous algae, in most of tropical West Africa. However, it
is not expkiined for Southern Angola where shallow ma-
rine environments resemble very much temperate Eu-
rope, but where Barleeidae clearlv outnumber Rissoidae
on the rock-v shore (Gofas, 1995).

There are 4 pairs of species with planktotrophic de-
velopment, and with respectively West African and West
European nonoverlapping ranges. These are:

Alvania aJhcana/A. bcani
Alvania manoi/A. canccllata
Ahania gofasi/A. zctlanclica
Crisilla transitoria/C. semistriata

Each one of these species p;iirs occupies similar hab-
itats and shares similar conchological characters, which
suggest that thev are closelv related. This is in contrast
to the usual concept of local radiations where loss of
planktotrophv has accompanied, or induced, local dif-

S. Gofas, 1999

Page 101

ferentiation. In this case, it is siiqirisiiii^ that species with
good tlispersal aliilit\ have not persisted as single pan-
mictic units. A possible scenario is that a single wide-
spread species has been chsrupted when increasing
Nortli/South climatic contrast has rendered impossible
the existence of an extensive latitudinal range, leading
the species to either be extinct on part ot its range or
to split up. This scenario (Eniller, 1977) implies that se-
lective pressure induced bv changing environmental
conditions mav lead to discontinuous character distri-
bution and, eventually, speciation along ioruierlv large
geographic ranges. This model has been invoked to ex-
plain .speciation events in planktic species (Pierrot-Bults
and van der Spoel, 1979). The model fits with the dis-
tributional data of the 4 species pairs mentioned abo\e.
Two other species (Pit.siUino iiiconspiciia and ObtiiseUa
intersecta) with planktotrophic lan'ae have apparently
maintained their large, all-inclusive tropical and temper-
ate range.

ACKNOWLEDGMENTS

This work would not ha\'e been possible without the
sujjport of friends and colleagues in Angola, particularly
the late Francisco Femandes who has been a companion
for most of the field work, Mario Albano dos Santos
"Carqueja" in Namibe, Joaquin Pinto Afonso in Ben-
guela, and Professor Eduardo Morais (UAN, Luanda).
The author is particurlarly grateful to Dr. Pierre Lozouet
(MNHN, Paris) for his insight into relevant Neogene
species and useful discussions on the manuscript, to Dr.
Angel A. Luque (Universidad Autonoma, Madrid) for
relevant information from his manuscript (with Dr. Eni-
ilio Rolan) on the Rissoina of the Cape Verde Islands,
and to Dr. Anders Waren (SMNH, Stockholm) for a con-
stnictive re\iew. The scanning electron micrographs
were taken by Juan-Jose Cuenca and Gregorio Caballero
at University of Malaga, or by Mrs. D. Guillaumin at
Centre Intenuiiversitaire de Microscopic Electronique,
Paris.

LITERATURE CITED

Bouchet, P. and A. Waren. 199.3. Re\ision of the Northeast

Atlantic bath\'al and ab\ssai Meso«;astropoda. Bollettino

Malacologico, Supplement .3:.579-84().
Dautzeiiberg, P. 1889. Contribution a la faune malacologique

des lies Azores. Resultats des Campagnes Scientifiques du

Prince de Monaco 1:1-112, 4 pis.
Dantzenherg, P. ("1912") 1913. Mission Gnivel sur la cote oc-

cidentale d'Afrique. Mollusques niarins. Annales de

rinstitut Oceanographi([ue .5(3):1-111, 3 pis.
Desjardin, M. 1949. Les Rissoina de lile de Cuba. Journal de

Conchyliologie 89:19.3-208, pis. 9-10.
Endler, J. A. 1977. Geographic variation, speciation, and dines.

Princeton University Press, Princeton.
Fretter, V. and A. Graham. 1978. The prosobranch molluscs of

Britain and Denmark. Part 4, Marine Rissoacca. fonnial

ot Molluscan Studies, Supplement 6:1.5.3-241,

Garci'a-Talaxeru, F. 1982. Los Moluscos Gasteropodos amfi-
atlanticos. Doctoral thesis, Coleccion Monografias 10,
Uni\ersidad ilc La Lasjuna, Tenerife, 3.52 pp., 7 pis.

Gofas, S. 1990. The littoral Rissoidae and Analiatliridae of Sao
Miguel, Azores. A^oreana, Supplement:97-1.34.

Gofas, S. 1995. A remarkable species richness of the Barleeidae
(Gastropoda: Rissoacea) in the eastern Adantic. The Nau-
tilus 109:14-37.

Gotas S. and A. Waren 1982. Ta.\onomie de quelrjues especes
du genre Alvania (Mollusca, Gastropoda) des cotes ibe-
riques et marocaines. Bollettino Malacologico 18:1-16.

Laniy, E. 1923. Canipagne du Si/lvaiui (fe\rier-juin 1913). Mis-
sion du Conite Jean de Polignac et de M. Louis Gain.
Mollusques testaces. Comptes Rendus du Congres des So-
cietes Sa\antes en 1922, Sciences:22-37.

Lozouet, P. 1990. Bcnthonellania nouveau genre de Rissoidae
(Gastropoda, Prosobranchia) du bathval atlantique. Bul-
letin du Museum national d'Histoire naturelle (4)12 (sec-
tion A, 2):31,3-328.

Moolenbeek, R. G. and M. J. Faber 1987. The Macaronesian
species of the genus Manzonia. De Kreukel 23(1):1-16,
pi. 1; 23(2-3): 23-31; 23(10): 166-179, pi. 2-3.

Moolenbeek, R. G. and H. J. Hoenselaar 1989. The genus Al-
vania on the Canary Islands and Madeira (Mollusca, Gas-
tropoda) part 1. Bulletin, Zoologisch Museum, Uni\ersi-
teit van Amsterdam 11:215-228.

Moolenbeek, R. G. and T. Piersma. 1990. A new Sctia species
from Mauritania (Gastropoda, Rissoidae). Gloria Maris 29:
31-.33.

Nickles, M. 1950. Mollusques testaces marins de la cote occi-
dentale d'Afrique. Manuels ouest-africains, 2. Leche\alier,
Paris, x + 269 pp.

Nordsieck, F. 1972. Die europaischen Meeresschnecken (Op-
isthobranchia mit Pyramidellidae; Rissoacea). Gustax-
Fischer, Stuttgart, .\ii -I- .327 pp.

Odhner, N. H. 1931. Beitriige zur Malakozoologie der Kanar-
ischen Inseln. Lamellibranchien. Cephalopoden, Gastro-
poden. Arkix' for Zoologi, 23A(14):1-116, pis. 1-2.

Pallary, P. 1920, Exploration scientifi(jue du Maroc. Malacolo-
gie. Institut Scientifique Cherifien, Rabat and Larose, Par-
is, 108 pp., 1 pi., 1 map.

Pierrot-Bults, A. C. and S, \an der Spoel. 1979. Speciation in
macrozooplankton. //!. S. \an der Spoel and AC. Pierrot-
Bults (eds.). Zoogeography and di\ersit\' of plankton. Ed-
ward Arnold, London, and Bunge, Utrecht., pp. 144-167.

Ponder, W. 1985. A review of the genera of the Rissoidae (Mol-
lusca: Mesogastropoda: Rissoacea). Records of the Austra-
lian Museum, Supplement 4:1-221.

Rolan, E. and F. Femandes. 1990. Tres nuexas especies del
genero Manzonia (Mollusca, Gastropoda) para la costa oc-
cidental de Africa. Publica^oes Ocasionais da Sociedade
Portuguesa de Malacologia 15:6.3-68.

Talavera, F. G[arcia-]. 1975. Moluscos de sedimentos de la pla-
taforma continental de Maiuitania. Boletin del Instituto
Espaiiol de Oceanografia 192:.3-18.

Waren, A. 1996. Ecology' and svstematics of the North Euro-
pean species of /S/.s.soc; and PnsiUina (Prosobranchia: Ris-
soidae). Journal of the Marine Biological Association of
the United King(k)m 76:101.3-10.59.

Watson, R. B. 1873. On the Marine Mollusca from Madeira,
including a new genus of the Muricidae, a new Eulima,
and the whole of the Rissoae of the group of islands. Pro-
ceedings of the Zoological Societv of London 1873:361-
.391, pis. .34-36.

THE NAUTILUS 113(3):102, 1999

Page 102

Book Review

Catalogue and Bibliography of the Marine
Shell-hearing Molltisca of Japan

hij Shun'ichi Higo, Paul Calloinon, and Yoshiliiro Goto.
1999. Elle Scientific Publications, Osaka, Japan. 749 pp.
$130.00

After compiling notes on Japanese Mollusca for some
30 years, Mr. Shun'ichi Higo pubBshed, in 1973, a list
of Japanese Mollusca, followed by a greatlv enlarged and
expanded version bv Higo & Goto in 1993. These two
important catalogues have not been widely utiUzed as
they were written in Japanese. This new catalogue, cov-
ering onlv marine shell-bearing mollusks, is entirely in
English and is a verv important tool for marine mala-
cologists, hsting 6,683 nominal species. This is the first
of a two-volume set, the second of which will consist of
new color plates of as many type specimens as possible,
including all Japanese tvpe specimens in major museums
such as the Smithsonian's National Museum of Natural
History (USNM), Japan's National Science Museum
(NSMT), the Natural Historv' Museum in London
(BM(NH)), and in important private collections in Ja-
pan, supplemented by photos of non-type specimens
from Japan.

The authors' intent is to list every species ever cited
from Japan and/or originallv described from Japan with
original combination, date, and publication reference
given for each. Given the nature of the malacological
literature there will almost certainly be a few errors and
omissions as in all such compilations. However, given the
large number of citations and references cited, the num-
ber of omissions, if any, must be very few. All ti«a are
attributed up through the family level and type species
are given for genera. The type locality and distribution
are given for each species. Habitat, when known, is also
included.

There are no manuscript names fisted and no fossils
unless the species also is found in the Recent fauna. The
authors stress that, because this work is compiled from
the existing fiterature, the number of species recortled
will almost certainly exceed the number of species ac-
tually existing in the area. This compilation provides a
platform from which systematists can work.

In addition to a complete citation for each species,
the Japanese vernacular name is given. Instead of these

names being in kana as usually rendered, thev have been
transcribed into the Roman alphabet with h\phens add-
ed to assist in pronunciation. The introduction to the
Catalogue gives an overview of the status and impor-
tance of these vernacular names, an explanation which
has long been needed in English as manv non-Japanese
wrongly consider the Japanese nomenclature to be a
competing system. An index of the Japanese vernacular
names is given in the Roman alphabet for the first time.

The systems used to transcribe Japanese kanji into the
Roman alphabet are explained. The two most widely
used systems have both been utifized in malacology,
sometimes with odd results. One example of the prob-
lems caused by ha\ing two systems is that we thus have
hvo verv different names, Mr. Ranzi Tiba and Mr Ranji
Chiba, being used for the same person. Some species
names based on variant transcriptions are "frozen" into
the binominal system, such as chlhanum and tibanum,
both proposed for species from Chiba Prefecture.

The short but very important and interesting intro-
duction to the Catalogue explains the use of new distri-
bution maps to indicate ranges for each species and in-
cludes information and insights about the Japanese mol-
luscan faima and its literature not to be found elsewhere.
Information about such classic locafities as "Ku" and
'Tosa", which show up frequently on specimens, but
which no longer exist under those names, is also includ-
ed.

The Catalogue's bibfiography is of great importance as
it contains not only all original references for marine
mollusks described from Japan, but it also fists manv
works originalK- written in Japanese with titles translated
into Engfish for the first time. This is the largest bibfi-
ograph\ of Japanese molluscan works ever published. A
fisting of Japanese serial publications, inclucfing address-
es of present publishers if still being printed, is of es-
pecial importance as manv of these journals are seldom
seen outside of Japan.

The book is a fine example of book production. It is
A4 in size, printed on cream stock paper and bound in
Damascene cloth with a gold stamped leatherette spine.
And, as is the case with most Japanese books, it comes
in a sfipcase.

Richard E. Petit

P.O. Box .30

North Myrtle Beach, SC 29582 USA

repetit@conipusei"ve.coni

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THE NAUTILUS

DEC 2 7 1999

Voliiinc 113, Niiiiil)cr4
December 16, 1999
ISSN 0028-1344

A quaiierh/ delated
to malacology.

EDITOR-IN-CHIEF

Dr. Jose H. Leal

The Bailey-Matthews Shell Museum
3075 Sanibel-Captiva Road
Sanibel, FL 33957

EDITOR EMERITUS

Dr. M. G. Harasewych
Department of Invertebrate Zoology
National Museum of
Natural Histon'
Smithsonian Institution
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ADMINISTRATIVE ASSISTANT

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3075 Sanibel-Captiva Road
Sanibel, FL 33957

CONSULTING EDITORS

Dr. Riidiger Bieler
Department of Invertebrates
Field Museum of
Natural History
Chicago, IL 60605

Dr. Arthur E. Bogan

North Carolina State Museum of

Natural Sciences

Raleigh, NC 27626

Dr. Philippe Bouchet
Laboratoire de Biologic des
Invertebres Marins et Malacologie
Museum National d'Histoire Naturelle
55, rue Buffon
Paris, 75005 France

Dr. Robert T.Dillon, Jr.

Department of Biology
College of Charleston
Charleston, SC 29424

Dr. William K. Emerson

Department of Living Invertebrates

The -American Museum of Natural

IlistorN'

New York, NY 10024

Dr. Eileen H. Joldnen
Institute of Water Resources
Universitv of Connecticut
Storrs,cf 06269-4018

Mr Richard I. Johnson
Department of Mollusks
Museum of Comparative Zoology
Hanard University
Cambridge, MA 02138

Dr. Douglas S. Jones
Florida Museum of Natural Histoiy
University of Florida
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Dr. James H. McLean
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Los Angeles County Museum of
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Dr. Arthur S. Merrill
% Department of Mollusks
Museum of Comparative Zoology
Harvard University
Cambridge, MA 02138

Dr. Paula M. Mikkelsen

Department of Living Invertebrates

The American Museum of Natural

Histor)'

NewYork, NY 10024

Dr. Gustav Paulay
Marine Laboratory-
University of Guam
Mangilao, Guam 96923

Mr. Richard E. Petit

RO.Box30

North Myrtle Beach, SC 29582

Dr Edward J. Petuch
Department of Geolog)'
Florida Atlantic University'
Boca Raton, FL 33431

Dr Gary Rosenberg
Department of Mollusks
The Academy of Natural Sciences
1900 Benjamin Franklin Parkwa\'
Philadelphia, PA 19103

Dr. Ruth D. Turner
Departnient of Mollusks
Museum of Comparative Zoology
Harvard University
Cambridge, MA 02 138

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Dr. John B. Wise

Houston Museum of Natural Science

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THEt^NAUTILUS

Vohimc 113. Number 4
DEC 2 7 1999 December 16, 1999

ISSN 0028-1344

CONTENTS

Eugene V. Coan Tht- eastern Pacific species of S))licitia (Bi\al\ia: Mvidae) 103

Roland Houart Two new species of the genus Chicorciis (Siratu.s)

(Gastropoda: Muricidae) from the western Atlantic 121

Michael J. Sweeney Harald A. Rehder (1907-1996): biographical sketch and

M. G. Harasewyeh malacological contributions 127

Notices 151

STATEMENT OF OWNERSHIP. MANAGEMENT AND CIRCULATION

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Puhlication Title, THE NAUTILUS

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THE NAUTILUS 113(4): 103-120, 1999

Page 103

The eastern Pacific species of Sphenia (BivaMa: Myidae)

Eugene V. Coan'

Department of Invertebrate Zoology
California Academy of Sciences
Golden Gate Park'
San Francisco, CA 9411S-4599 USA

ABSTRACT

There are four eastern Pacific Ocean species of the genus
Sphenia: (1) Sphenia fragilis (H. Adams and A. Adams, 1854),
occurs in a \'ariety of nestling situations from the inteitidal zone
to shallow water, from Santa Barbara County, California, to
Guayas Province, Ecuador, and has as synonyms S. fragilis Car-
penter, 1857; S. pacificensis de Folin, 1867; and S. tnincubis
Dall, 1916; as well as the western Atlantic Corbida iheringiana
Pilsbrv-, 1897; and S, antiUen.si.s Dall and Simpson, 1901. (2)
Sphenia gulfensis. a new species, is restricted to soft bottoms
in the Gulf of Cahfomia. (3) Sphenia hafcheri Pilsbr), 1899,
occurs, probably in relati\eK soft substrata, from Rocha De-
partment, Uruguay, through the Estreclio de Magallanes, as far
north as Isla Chiloe, Chile; S. subequalis Dall, 1908, is a syn-
onym. (4) Sphenia hiticola (\'alenciennes, 1846), occurs off-
shore in rock caxities from Jefferson Counts, Washington, to
San Diego Count\'. California, and has as sviionvms S. pholad-
iclea Dall, 1916; Cuspidaiia nana Oldrovd. 1918; and S. ajobnla
Dall, 1919. Sphenia bihrata Gabb, 1861. may have been based
on Recent specimens of Hiatella arctica (Linnaeus, 1758).
Sphenia ovoidea Carpenter 1864, is based on a juvenile Mija.
most probably .^/ arenaria Linnaeus, 1758

INTRODUCTION

Bernard (1983: 58) pro\icled the most recent list with
svnonvniies of the eastern Pacific species of the myoi-
dean genus Sphcnio. He recognized three: S. hatchcri
Pilsbry, 1899, in southern Chile; S. Ittticola (Valenci-
ennes, 1846), with a number of synonyms, from north-
em Cahfornia to northern Peru; and S. ovoidea Caq^en-
ter, 1864, from southeastern Alaska to Puget Sound,
Washington. At the same time he erected a new sul)fam-
ilv, the Spheniinae, based on two morphological features
(Bernard, 1983: 58, 70).

Mv attention was drawm to some material that proved
to belong to Sphenia from the Colfo de Cahfornia and
that had been misidentified as a new species of Ba.ste-
rotia, a genus of the Sportellidae, which 1 was then
studying. When that review was completed (Coan, 1997;
1999), I returned to this material, which proved instead

' Mailing address: 891 San Jude Avenue, Palo Alto, CA 94306-
2640 USA; 2ene.coan@sierraciub.org

to be a new species oi Spltcnia Examination of the avail-
able eastern Pacific material of Sphenia, including the
ty|De material of the described species, has resulted in
conclusions that differ significantly from those of Ber-
nard (1983),

Pre\ious taxonomic lists and studies of Sphenia in-
clude tho.se of A. Adams (1851), Tryon (1869: 67), E. A.
Smith (1893), Lamv (1927: 176-184), Lewis (1968), and
Hanks and Packer (1985). Bahik and Radwariski (1979)
discussed the moiphological variabihtv of the Miocene
Sphenia anatina (Basterot, 1825: 92) caused by its nes-
thng site.

Forbes and Hanley (1848: 189-193) were the first to
describe the anatomv of a species oi' Sphenia. S. hingha-
mi Turtf)n, 1822, from notes provided b\ WilUam Clark.
Yonge (1951) reviewed the functional moq^hology of S.
bint^Iunni, and Narchi and Domaneschi (1993) that of S.
antillensis Dall and Simpson, 1901

Some species, such as S. hinghanii, nestle in rock
crevices and similar situations, whereas others, such as
S. antillcnsis, are somewhat more widespread nestlers,
occurring among colonial ascidians, polychaete tubes,
and the byssal threads of mviilid clams. Sphenia sincera
Hanks and Packer, 1985, occurs in soft bottoms, but
there is not yet an account of how its functional mor-
pholog\- differs from the two taxa studied thus far.

FORMAT

In the following treatment, each \alid taxon is charac-
terized bv its svnonvmv, description, information on tvpe
specimens and tvpe locafities, notes on distribution and
habitat, and an additional discussion.

The synonymies include all major taxonomic accounts
about the species, but not most minor mentions in the
literature. The entries are arranged in chronological or-
der under each species name, with changes in generic
allocation from the previous entry, if any, and other notes
given in parentheses.

The distributional information is based on Recent
specimens I have examined, except as noted. Fossil oc-
currences are taken from the hterature.

The following abbreviations are used in the text:
AMNH, American Museimi of Natural Historv, New

Page 104

THE NAUTILUS, Vol. 113, No. 4

York, New York, USA; BM(NH), The Natural History
Museum, London, England; CAS, California Academv
of Sciences, San Francisco, CaUfomia, USA; LACM,
Natural History Museum of Los Angeles County, Cali-
fornia, USA; MACN, Museo Argentino de Ciencias Na-
turales, Buenos Aires, Argentina; MNHN, Museum Na-
tional d'Histoire Naturelle, Paris, France; SBMNH, San-
ta Barbara Museum of Natural Histor\', Santa Barbara,
California, USA; UCMP Universit)' of California Mu-
seum of Paleontology, Berkeley, Caliiornia, USA;
USNM, National Museum of Natural Historw Smithson-
ian Institution, Washington, DC, USA; UMML, Marine
Invertebrate Museum, Rosenstiel School of Marine and
Atmospheric Science, Miami, Florida, USA; Skoglund
Collection, collection of Carol C. Skoglund, Phoenix, Ar-
izona, USA.

MORPHOLOGICAL CHARACTERS

Due to the nesthng habitat of most species, overall shell
shape is not an easil)' used character in identification ot
species. Variabiht)' in shape is illustrated in the material
figured herein. Some species of Spltcnia, such as S. Iti-
ticola and S. anotino, are highlv \'ariable in shape, in-
habiting pre-established cavities in relatively hard sub-
strata. Splienio fragilis, nesthng amongst such materials
as brvozoan colonies, is onl\ slightK' less variable. Those
species occurring in softer substrata, such as S. sinccra,
and presmnabK' S. ffilfensis and S. hatchcii, retain a
more defined shape.

While the shape of the ends, especially the posterior
end, is also variable, the degree of elongation and infla-
tion of the anterior end is a useful character that helps,
for example, to chstinguish S. fragilis from S. gulfcnsis.

The periostracum differs among species. The extent
to which the periostracum and, in one case, the outer
shell layer extend the posterior end is particularK im-
portant in species distinction.

The chondrophore in the left valve, while exliibiting
xariabilitv in size and shape, pro\ddes at least three use-
ful characters. In small specimens of all species, it is
approximately horizontal. As some species reach their
maximum size, it may become more vertical, such as in
S. liiticola. Also useful is the degree of elongation of die
posterior attachment (the "posterior buttress") and the
length of the posterior ridge (figures 1, 18-22).

The placement ot the anterior muscle scar — located
either close to the ventral margin or more toward the
middle of the anterior margin — proves to be a usefid
specific character. The extent to which the posterior ad-
ductor muscle scar and the pallial sinus are recessed
from the posterior end is also a useful character These
features are illustrated with Hne drawings (figures 18-
22).

Finally, the size of the discontinuous scars defining the
palhal sinus differs among the four eastern Pacific spe-
cies.

The most important specific characters are gix'en in
Table 1.

1

Figure 1. Contrasting features ot the chondrophores of two
species of Sphenia. (a) S. gi/Z/c/i-sw and (b) S. luticola. The first
has a long posterior bvittress (arrow) in contrast to the latter
The second has a long, projecting posterior ridge (asterisk) in
contrast to the first.

SYSTEMATICS

Family Mvidae Lamarck, 1809; 319, as "Myaires"
Subfamily Crsptomvinae Habe, 1977: 279

This subfamily is thus far distinguished from the M\'inae
only on the basis its short siphons and shallow pallial
sinus.

Bernard (1983: 58, 70) proposed the Spheniinae
based on ^^vo characters: (1) the "resilifer" in the left
\'alve was said to be "only superficiall)' similar" to the
"chondrophore" oi Mya, and (2) "the lateral tubercles of
the right xaKe ha\'e no counteqiart in Mya"

I behe\e that this rationale is indefensible. In the first
instance, a chontlrophore is merely a hollowed-out, pro-
jecting resihfer, and the left \'alves of both Sphenia and
Mya have similar structures. Mya attains a much larger
size than Sphenia, and its chondrophore ultimateK' be-
comes more prominent. However at an equivalent size,
the chontlrophores and their features in the two genera
are \irtuallv identical. Indeed, in describing a new spe-
cies of Splicnia, Hanks and Packer (1985) use terminol-
ogy for chondrophore features first proposed for those
oi Mya by MacNeil (1965) and earlier authors. Bernard's
reference to "lateral tubercles" in Sphenia is equally per-
plexing. The eastern Pacific species of Sphenia have a
small, slightly projecting anterior carchnal tooth in the
right \'al\e, varying in prominence among species; there
is no posterior tooth. There is no projecting anterior car-
chnal tooth in Ml/a of equivalent size, but the posterior
end of its anterodorsal margin ma\' be expanded and
turned ventralK' just anterior to the resilifer The pres-
ence or absence of a small tooth ma\' differentiate mem-
bers of the two genera, but it does not seem to be a
fundamental difference.

Bernard's differentiation of the Spheniinae from Mtfa

E. V. Coan, 1999

Page 105

Tabic 1. Ke^• differentiatiiii; characters, size, and frequencv of eastern Pacific Spliciiiti. Total 204 lots.

Shell thickness/
shape

Anterior end

Chondropliore

orientation/
posterior ridge

Anterior

adductor

muscle scar

Posterior

aikluctor

nniscle scar

and pallial

Number of
eastern Pa-
M;l\. cific lots
size, nnn studied

Sphenia fragilis Thin/\ariable —
o\'ate, ovate-
elongate, sub
quadrate

Sphenia giilfensis Thin/elongate

Sphcniii liatcluii

Moderate/elon-
gate

Proportionately
nitiated

More inflated,

but tapering

gradualK
.More inflated,

but tapering

gradualK

Sphenia liiticola Thick/\ ariable — More inflated,
ovate, elongate but tapering
gradualK

Horizontal, but a
little \-entrally/
ridge not pro-
jecting

Horizontal/ridge
not projecting

Equal horizontal-
\\ and \entral-
K/ridge slightK
projecting

More xentral/
ridge project-
ing

Near ventral
margin

Central

Near \entral
margin

Near x'entral
margin

Clo.se to pos-
terior end

Recessed
from poste-
rior end

Reces.sed
from poste-
rior end

Close to pos-
terior end

12.'

1.3.1

.30.1

20.6

1.38

31

34

i.s still more puzzling, because Sphenia had until then
been placed in the Cryptomyinae Habe, 1977, which he
also recognized. It was originally erected for in\ids with
short siphons, and into it Habe placed Cn/ptdiin/a Con-
rad, 1S49: 121; Distiigonia Iredale, 19.36: 2S.3; Paranuja
Conrad, 1860: 232 [noincn novum pro Myalina Conrad,
1845: 65, non Konnick, 1842: 125]-; Sphenia- and Ve-
nafomi/a Iredale. 19.30: 403.

Oxerall relationships among the genera oi Myidae
could be inferred from cladistic analysis of an array of
anatomical and shell moi-phological characters. The siib-
familv Crvptom\inae currentK' stands on minimal
grounds.

Genus Sphenia Turton, LS22

Sphenia Turton, 1822: x\qi, x.xiii, .36-38, 261 (ex Leacli ms, ac-
cording to Gray, 1847). T\pe species: Sphenia binghanu
Turton, 1822: .vdii, .36-37' 2.57, 277, pi. 3, figs. 4, .5; bv
the subsequent designation of Gra\ (1847: 190). A lecto-
t\pe for this species was designated and figured bv Hanks
and Packer (198.5: figs. 1, 2).'

Sliaenia, Spaena. Sphaena, Sphaenin. Sphaeria. Sphena. Spen-
ia, Sphenica, Sphoenia. Sijphonia. misspellings of authors.

Tijleria H. Adams and A. Adams, 18.54: 418. T)pe species: Tij-
leria fraaihs H. Adams and A. Adams, 1854: 418; by
rnonotspx'.

- Placement of the genus Paranu/a remains a iiixsterv. The re-
silifers in the two valves are equallv recessed and U-shaped,
very- different from those of other mvids. There is no pallial
sinus, and the external shell smface is pustulose, the latter sug-
gesting to Campbell (1993:.33) that it the species might be a
member of the Sportellidae, Moreover, like the sportellid ge-
nus Anisodonta Deshayes, 1858:542, Paranuja has a thick in-
ternal radial rib just posterior to the anterior adductor How-
ever, sportellids ha\e not been reportetl to have large internal
hgaments. Onlv an examination of the anatomy of this genus
will resoKe its taxonomic position.

Tilerin. misspelling ot authors.

Description: Shell mecUum-sized to small, irregular in
shape in nestling species. Posterior end truncate. Chon-
drophore narrow to somewhat projecting. Small anterior
cardinal present in right valve. Pallial sinus shallow. Pal-
lial hue a series of discontinuous scars. Siphons short and
united, with tips surrounded bv single, common row of
short tentacles. Incurrent siphon circled by additional
row of tentacles, excurrent siphon with tubular NaKiilar
membrane, Ctenidia large and unplicated, inner demi-
branch larger, with supra-axial extension. Palps small and
differing in size between species. Foot small, narrow,
serving in S. hinghanii and ,S. antiUcnsis to attach the
bvssus to hard substrata.

Splienia fragiUs (H. Adams and A. Adams, 1854)
(Figures" 2-6, 18, 24)

Ti/Ieria fragilis H. Adams and A. Adams, 1854: 418; 1856: 368,
pi 97. figs. 3, .3a; Carpenter, lS57a; 245, .300; 1857b: 25
(as a possible svnonvm of Sphenia fraojUs Carpenter),
547-548 (as something unique): Keen. 1968: 400—401 (as
a senior .sviiouNiii of S. frao'ilis Carpenter); Bernard, 1983:
58 (as a s\iionvm of S. liiticola)

Sphenia fra0lis Caipenter, 1857a: 244, .300, nomen nuchon.
1857b: 24-25, 530; 1864: 543 (as a possible svnion\m of
P luticola). 553, 619 (1872: 29, .39, 105); de Folin,1867:
53 [ = 15], pi. 2, figs. 7-9; Trvon, 1869; 67; E. A. Smidi,
1893; 279; Oldroyd, 1924: 62^3; 1925: 200; Lamy, 1927:
179-180 (as a .smiouvto of P luticola. but under die head-
ing of S. fragility. Palmer, 1951: 13, 1958: 116; Keen.
19.58: 207, fig. .521: Fnmc, I960: 2120-2121, fig. E; Ols-
son, 1961: 424-425, 550, pi. 77, fig. 9, 9a, b; Brann, 1966;
pi, 6, fig. 35 (tablet 80); Keen, 1968: .394, 400-401, fig,
22a-<'; 1971: 262, 26.3-264, fig. 673; Bernard, 1983: 58 (as
a synonym of S. luticola]

Sphenia pacificensis de Folin, 1867: 5,3-54 [ = 15-16]; de Fol-
in and Perier, 1867: 8 (as pacificiensis): Crosse, 1868: 218
(as paciftca. unjustified emendation); \on Martens, 1868:

Page 106

THE NAUTILUS, Vol. 113, No. 4

E. V. Coan, 1999

Page 107

586 (as pacifica); E. A. Smith, 1893; 280, pi. 15A, fig. 7

(as S. pacifica): Keen, 1958: 207 (as a varieh' of S. frap,ilis);

Kisch, 1960: 149, 162; Lamy, 1927: 181; Keen, 1971: 263

(as a svmonvii) of S. fragilis); Benianl, 1983: 58 (as a svii-

on\iii of S. Iiiticola)
Corbula ihcrini!jann Pilsbn', 1897a: 8, nomen nudum; 1897b:

295, pi. 7, figs. 2-)-26;' Figneiras and Sicaiili, 197()a: 411,

pi. 5, fig. 78'
Splu'iiia aiitillcnsii Dali and Simpson, 1901: 474, pi. .55, fig.

14; Lamv, 1927: 181: VVarmke and Abbott, 1961: 20.5-206,

pi. 43, fig. e; Abbott, 1974: 537; \'oi<es and Voi<es, 1983;

44, 65, 164, pi. 48, fig. 17; Narchi and Domaneschi, 1993:

19.5-210; Rios, 1994: 291. pi. 99, fig. 1419
Sphenia tninculiis Daii, 1916a: 41, noincn nudum: 1916b: 415;

Oidroyd, 1925: 201; Lamy, 1927; 181; Bernard, 1983: 58

(as a sviionviii of S. luticola)
Sphenia luticola Valenciennes, auctt.. non Valenciennes, 1846.

Gemmeii et al.. 1987; .58, fig. 71

Description: Shell thin, ovate to ovate-elongate to
.subqiiadrate; shape variable, depending on habitat; right
valve slightlv more inflated, often overlapping left valve
at postero-ventral margin in less distorted specimens;
anterior end generalK- short, broadK rounded, particii-
larlv inflated; posterior end from short to produced,
truncate posteriorly, sometimes twisted to left or right,
often e.xtended bv periostracum, slightlv gaping. Exter-
nal surface with fine commarginal ribs; periostracum
thin, dark to light tan, adherent. Left valve with medi-
um-sized to large, triangular chondrophore, projecting
towards right valve, but also somewhat ventraJlv; poste-
rior buttress short; posterior ridge not much extended
beN'ond ventral edge of chondrophore. Right valve with
resilifer under beaks, more deeply recessed than that of
S. luticola, and with small anterior cardinal tooth. An-
terior adductor muscle scar positioned ventrally. Poste-
rior adductor scar and pallial sinus close to posterior end.
Pallial sinus a series of 2-4 narrow sears; pallial Une a
discontinuous series of small, rounded scars (Figure 18).
Length to 12.7 mm (USNM 733245; Fort Amador, Ca-
nal Zone, Panama); in the western Atlantic, to 14.6 mm
(Narchi and Domaneschi, 1993: 196, misprinted as "146
mm").

Type material and localities:

T /rag(/K— BM(NH) 1878.1.28.64, lioiotype (figure 2a, b):
lengtii. 6.3 mm; height, 5.0 mm; tliickness cannot be mea-
sured because ot breakage. Mazatlan, Sinaloa, Me.xico
(23.2°N); nestling in a sand-filled cavity in a Spondi/lus
valve; R. W. Tyler. The hoiotype is now badiv broken.
Copies of the original figures as well as photographs of
tlie present state of the specimen are provided here.

S. fragdis — BM(NH) Carpenter Mazatlan Collection
1857.6.4.82/1 [species .35, tablet 82], iectotvpe herein des-
ignated, the largest set of paired valves, length, approxi-

mately 8 mm; height, approximately 4.9 mm; tliickness
cannot be measured because .specimen glued on slide
(Figure 3); this specimen was figured bv Keen (1968: figs.
22a, b). Paralectotypes; BM(NH) 1857.6.4.82/2, one left
valve glued on same slide; 1857.6.4.81/1-6, 5 smaller
pairs, .some of which are broken, on slide, one loose valve;
1857.6.4.80/1-14, 10 still smaller pairs and four valves on
slide; USNM 71.5649, 40 pairs, valves, and fragments. Ma-
zatlan, Sinaloa, Mexico (23.2°N); inhabiting the burrows
of worms and molhisks in shells of chamids and Spondtj-
lus: also in dead barnacles on Stromhus: F. Reigen.

S. pacificensis — BM(NH) 196460/1, Iectotvpe herein designat-
ed, the only intact pair; length, 5.8 nnn; height, 3.6 mm;
thickness. .3.4 mm (Figure 4). BM(NH) 196460/2-3, par-
alectotypes, 1 pair with right valve broken anteriorly,
length, 5.2 mm; 1 pair with both valves broken, length,
appro.xiniately 6.2 mm. Archipielago de las Perlas, Panama
(approximately 8.4°N).

C. iheringiana — ANSP 70541, lectotype herein designated,
closed pair; length, 9.0 mm; height. 5.6 mm; thickniess,
3.8 mm (Figure 24); ANSP 4013.3.3, paralectotypes, 1 right
valve, length 7.4 mm: 1 open pair, length, 6.5 mm; 1
closed pair, length, 3.9 mm. Bahia Maldonado, Maldonado
Department, Uruguay (34.9°S); ;3-6 fms. [6-11 m].

S antillensis — USNM 160495, holohpe, pair; length, 4.0 mm;
height, 2.5 mm; thickness, 4.5 mm (Figure 5). Plava del
Ponce, Puerto Rico (1S.0°N, 66.6°W). The right valve of
this pair is now split but not broken.

S. tnincidus — USNM 160116, lectotype herein designated,
closed pair; length, 6.9 mm; height, 5.0 mm; thickness.
3.8 nun (Figure 6): USNM 880250, paralectotvpe, smaller
closed pair; length, 6.6 mm. San Diego, California
(.32.7°N); among barnacles on wharf piles.

Distribution and habitat: Carpinteria, Santa Barbara
County, Cahfoniia (.34.6°N) (LACM 77-30.22, SBMNH
31575), throughout the Golfo de California, to its head
at Puerto Pefiasco, Sonora, Mexico (31.3°N), south to
Salinas, Guayas Province, Ecuador (2.2°S) (CAS
110971); intertidal zone to 55 m " (mean, 4 m; n = 50),
nestling in pre-existing holes in rock, wood, and other
mollusks, and among brv'ozoan colonies and in similar
situations. A record from Zorritos, Tumbes Province,
Peni (3.7°S) (Olsson, 1961; 425), could not be verified
and was perhaps based on a specimen of Alii^cna
(UMML 30.10456): in the western Adantic from Hunt-
ing Island State Park, South Carolina (32..3°N) (L. D.
Campbell, in correspondence, 19 Julv 1997), Saint Lucie

'The specimen forming the basis of Durham's (1942:121) re-
cord oi "Sphenia sp." from 393 m west of Isla Tiburon, Sonora,
Mexico, cannot now be located in the UCMP, and thus a deep
record of either this or the next species cannot be confirmed.
The specimen might have proven to belong to some other ge-
nus and has now been filed elsewhere.

Figures 2-4. Sphenia fragdis (H. Adams and A. Adams). 2. Holohpe of Ti/leiia fragdis H. .^dams and A. Adams, BM(NH)
1878.1.28.64, length = 6.3 mm. (a) Figures of H. Adams and A. Adams (1856); (b) photographs of current state of specimen. 3.
Lectotype of Sphenia fragdis. BM(NH) Carjienter Mazatlan Collection 1857.6.4.82/1, length = 8 mm (external view through glass
shde). 4. Lectotype of Sphenia pacificetisis, BM(NH) 196460/1, length = 5.8 mm.

Page 108

THE NAUTILUS, Vol. 113, No. 4

E. V. Coan, 1999

Page 109

(USNM 861964) and Sanihel Island (SBMNH 144631),
Florida, and South Padre Island, Texas (Abbott, 1974),
to Santa Catarina State, Brazil (approx. 28°S) (Rios,
1994), and south to Bahia Mandonado, Maldonado De-
partment, Uniguav (34.9°S) (t\pe locidit)' of Corbula
ihcring^iana- SBMNH 145228), I have seen 138 eastern
Pacific lots and about 30 western Atlantic lots.

Di.scussion: Ti/lcria fra>^ilis was proposed as a new ge-
nus and species In' H. .A.danis and A. Adams in 1854 for
a "curious little shell" found nestling in a sand-filled cav-
it\- in a Spoiuli/ltis vAw. Figured by these authors two
N'ears later, it was characterized as having a chondro-
phore and a short pallial sinus like a Sphcnia, but it was
also said to ha\e a partlv external ligament and the an-
terior end was said to have a ridge below the dorsal
margin supported by a series of calcareous septa dorsal
to it (figure 2a).

When Carpenter (1857b: 24) first discussed Ti/Iciia
frazil is (in a portion of the Mazatlan Catalogue typeset
in 1855), and having seen only a drawing of it, he said,
"I have occasionalK' noticed valves of Sphcnia with a
tendencv towards the same crenation at the posterior
end, apparentK' through the irritation of sand. ' (He ev-
idently confused or misstated which end had the "ore-
nation. ") Later, in a portion of his book typeset in 1857
(p. 547), he said that he had seen the actual specimen
and concluded that it was unique, not a Sphcnia; in the
same work he described Sphcnia fraf^ihs as a new spe-
cies, thus selecting the same species name.

When Mvra Keen examined the holotvpe of Tijleiia
fra^iUs in The Natural Histors' Museum in London she
concluded that it realK' was a pathological specimen of
Sphenia in which the shell material was "either not de-
posited or was resorbed" along the dorsal margin, leav-
ing "a few islands of solid calcium carbonate embedded
in v\hat is mostK' periostracum" (Keen, 1968). (Perios-
tracuni bridging the two valves probably accounts for the
Adams' observation of an external ligament.)

The holotvpe of Ti/lciia fragilis remains a puzzle. I
have been able to examine onK photographs of the spec-
imen, which is now too fragile to travel. The specimen
was badh' eroded from the start and has since been
much broken; the outer periostracal layer is now mostly
gone. The "calcareous septa" seen bv the Adams' and
the "islands of solid calcium carbonate " observed by
Keen now appear with loss of the periostracum to re-
semble short radial ribs (figure 2b). No other specimens
of Sphenia have been seen like this, and it may have
been caused h\ the animal's nestling site. On the other

hand, the shallow pallial sinus defined by a series of dis-
continuous scars comports with Sphcnia. The chondro-
phore and resilifer also fit this genus: while both seem
distorted, such distortion is not unusual. A greater com-
fort level with the identity- of the specimen could be
achieved by close examination of the chondrophore, be-
cause that of nivids has unitjue features. While I ha%e
not seen other specimens of Sphcnia with its singular
morphology, no other known nesthng clam better ex-
plains the specimen, and it is best to leave the svnonviny
of the two species named fragiUs as it is.

The t\pe specimen oi' Spltcnia antillcnsis is small, flat,
and now chipped (figure 5). All other western Atlantic
material I have examined, including the 23 lots available
in the USNM, specimens in musouns on the west coast,
and a set of the material studied In Narchi and Doma-
nesehi (1993) [SBMNH 145246],' is indistinguishable
from eastern Pacific specimens of Sphcnia fragiUs in all
features here found to be useful here in distinguishing
species Sphcnia — shell shape and thickness, maximum
size, chondrophore orientation and moq:)holog)', pallial
sinus details, and positions of the adductor muscle scars,
as well as in reported habitat.

The nomenclatural situation is further complicated by
the fact that Sphcnia antiUcnms is not even the earliest
name available in the western Atlantic. The first name
proposed for a Recent western Atlantic Sphenia is Cor-
bula iheiingiana based on material from Urugua%', with
a lectohpe figured here with a photograph for the first
time (figure 24).

Because of the fact that western Atlantic material can-
not be distinguished from eastern Pacific material bv any
criteria found useful at the species level in this genus, it
is inescapable to use the first name proposed until such
time as significant differences are discovered.

Given its shallow habitat and presence in fouling com-
munities, it is not impossible that Sphcnia fragiJis has
been transported h\ ship traffic during the last 300 years
either to the east or the west. As evidence that such
transport can occur in this genus is the recent discover\'
of a few .specimens of Sphcnia in Pearl Harbor, Oaliu,
Hawaii (SBMNH 144633). This material has been ten-
tatively identified as the western Pacific Sphcnia corca-
nica Habe, 1951:76, based on its heavier comniarginal
sculpture and pallial sinus positioned further from the
posterior end. (Habe's original material was small and
oval, and had not grown the elongate posterior end t\p-
ical of larger specimens.) However, the relationships of
this material and of western Pacific Sphcnia corcanica

Figures 5-7. Sphenia fragiUs (H. Adams and A. Adams). 5. Holotvpe of Sphcnia antiUciisis; USNM 160495; leiigtii, 4.0 mm. 6.
Lectotype o{ Sphenia tntncuhis. USNM 160116, length = 6.9 mm. 7. SBMNH 1446.30, Puerto Peiiasco, Sonora, Mexico, lengtli
= 9.1 mm.

Figures 8-10. Sphcnia giilfcnsis new species. 8. Holotvpe, left valve, SBMNH 144627, length = 11.7 mm. 9. Paratvpe, right
valve, SBMNH 144628, length, = 0.6 mm. 10. Lot sliovving variability in shape, SBMNH 144345, Puertecitos, Baja Califoniia
(Norte), 18 m.

Page 110

THE NAUTILUS. Vol. 113. No. 4

to die eastern Pacific S'. fra^ilis merits additional stud)-,
ideally augmented with information from biochemical
genetics.

Sphenia gulfensis new species
(Figures la, 8-10, 19, 20)

Description: Shell thin, elongate; anterior end pro-
duced, broadlv roimded, tapering in thickness gradualh'
from anterior to posterior end; right valve more inflated,
generally extending well beyond posteroventral margin
of left valve; posterior end often very elongate, tapered
verticallv, broadlv truncate, sometimes flared, narrower
in left valve, not extended b\- periostracum, slightK gap-
ing. External surface with fine commarginal ribs. Peri-
ostracum thin, verv' light tan. adherent. Left valve with
narrow chondrophore. projecting horizontalK- toward
right valve, even in large specimens; posterior buttress
elongate in large specimens; posterior ridge not extend-
ing beyond ventral margin of chondrophore. Right valve
with resilifer more deeplv recessed than that of S. fra-
gilis, and with more conspicuous anterior cardinal tooth.
Anterior adductor scar at middle of anterior end. Pos-
terior adductor scar and pallial sinus well recessed from
posterior margin. Pallial sinus a series of 3^ narrow-
scars; pallial line a discontinuous series of small, rounded
scars (figures 19, 20). An additional lot is illustrated to
show the variabihty in shape of this species (Figure 10).
Length to 13.1 mm (SBMNH 144345; Puertecitos, Baja
California [Norte], Golfo de California).

Type material: SBMNH 144627, holotype, left valve;
length, 11.7 mm; height, 6.6 mm; thickness, 2.2 mm
(Figure 7, 19); SBMNH 144628, figured parat\pe, right
valve; length, 10.6 mm; height 5.9 mm; thickness. 1.9
mm (figures 8, 20); SBMNH 144629, four additional
paratypes, of lengths 12.8 mm (right valve), 12.0 mm
(left valve), 11.1 mm (right valve), 11.0 mm (left valve),
all from type locality, Peter and Sally Bennett, March
1976.

T)pe locality: San Felipe, Baja California (Norte),
Gulf of California, Mexico (31.0°N, 114.8°W), 15-22 m,
mud bottom.

Distribution: Puerto Escondido, Baja California Sur
(25.8°N) (CAS 112929). northward to the head of the
Gulf of California at Bahi'a ChoUa, Puerto Peiiasco, So-
nora (31.4°N) (SBMNH 115752, UCMP E8398, Sko-
glund Coll.), south to Mazatlan, Sinaloa (23.2°N)
(UCMP E9704), Mexico, from the intertidal zone to 61
m (mean = 15 m; n = 13), on soft bottoms. I have
examined 31 lots.

Discussion: This species has sometimes been separat-
ed in coUections as differing from S. fragilis. Stillman

Berrs' recognized it in material he had collected at
Puerto Peiiasco, Sonora, in 1949 and 1952 (SBMNH
115752), and I isolated it aside in material I had col-
lected at San Felipe, Baja California (Norte), in 1963
(UCMP E9705).

This new species differs from S. fragilis as follows: (1)
the anterior end of S. gulfensis is longer and propor-
tionately less inflated; (2) overall, it is more elongate; (3)
while varsing somewhat in shape, not as much as S. fra-
gilis and never as irregular, suggesting a non-nestling
habitat, (4) the right valve overlaps the left valve more
extensivelv postero-ventrallv; (5) the chondrophore in
the left valve, while varying somewhat in shape, is nor-
malK' smaller and projects more horizontally, and has a
longer posterior buttress; the resilifer in the right valve
is consequently more deepK* recessed; (6) the cardinal
tooth in the right viilve is more con.spicuous; (7) the pos-
terior end is sometimes flared and is not extended by
periostracum; and (8) the posterior adductor and the
pallial sinus are positioned further from die posterior
end.

In the Golfo de California, where their distributions
overlap, specimens of S. fragilis and S. gulfensis smaller
than 2 mm are difficult to differentiate. In such material,
S. fragilis is most easilv distinguished by its more round-
ed outline, more inflated posterior end. its more poste-
riorly positioned pallial sinus, and its proportionately
larger chondrophore.

This new species differs from S. sinccra Hanks and
Packer, 1985, which inhabits soft bottoms in 30 to 63 m
in northern New England, in being longer and more
tapered posteriorK-, having a narrower less projecting
chondrophore, having a thinner almost transparent peri-
ostracum, and in attaining a larger size.

Etymology: The name gulfensis refers to the fact diat
the known distribuHon of this new species is chiefly re-
stricted to the Gulf of California.

Sphenia hatcheri Pilsbry, 1899
(Figures 11-13, 21)

Sphenia hatcheri Pilsbrv, 1899: 128, pi. 1, fig. 5. 6: Lamv, 1927:
181: Carcelles, 1950: 82, pi. 5, fig. 94; Carcelles and Wil-
liamson, 1951: 347; Castellanos. 1965: 173-175. pi 1. figs.
1-8; Fi^ueiras and Sicardi. 1970b: 22, pi. 7. fi^. 103: Cas-
tellanos", 1970: 278-279, 348, pi. 25, figs. 9-ir[349]: Ber-
nard, 1983: 58.

Sphenia subequalis Dall, 1908: 422-423; Lamy, 1927: 181-182;
Carcelles, 1950: 82; Castellanos, 1965: 174 [as a svnonvTn
of S. hatcheri]; Bernard, 1983: 58 [as a synonym of S.
hatcheri].

Description: Shell of moderate thickniess, elongate;
subequivalve; anterior end produced, rounded, width ta-
pering gradually towards posterior end; posterior end

Figures 11-13. Sphenia hatcheri. 11. Holotvpe; ANSP 18083, length = 17.2 mm. 12. Holotvpe of Sphenia sabeipialis, left valve,
USNM 110719. length = 6.9 nnn 13. Specimen from Isla Chiloe, Chile, SBMNH 144635, length = 30.1 mm.

E. V. Coan, 1999

Page 111

Page 112

THE NAUTILUS. Vol. 113. No. 4

elongate, tapering, truncate posteriorly, extended by
periostracum, scarceK- gaping. E.xtemal surface with fine
to moderate commarginal ribs. Periostraciun of moder-
ate thickness, thicker on posterior slope, light tan. erod-
ed in large specimens. Left \alve with a large chondro-
phore. projecting as much ventrallv as horizontalK'; an-
terior buttress wide; posterior buttress short; posterior
ridge extending a little beyond ventral margin of
chondrophore. Right valve with resOifer not verv deepK-
recessed, with a conspicuous anterior cardinal tooth. An-
terior adductor placed antero-ventralK'. Posterior adduc-
tor and palhal sinus recessed from posterior margin. Pal-
hal sinus of one broad and several smaller scars; pallial
line a discontinuous series of small scars (Figure 21).
Length to 30.1 mm (SBMNH 144635; Isla Chiloe.
Chile).

Type material and localities:

S. hatcheri — ANSP 1808.3, holot\'pe, pair; length. 17.2 mm;

height. 9.2 mm; thickness. 7.3 mm (Figure 11). "Cape

Fairweather" (Cabo Buen Tiempo). Santa Cruz Province.

Argentina (.51..5°S).
S. sitbequalis — USNM 110719. holotvpe. left valve; lengtli, 6.9

mm; height. 4.3 mm; thickness. 1.3 mm (Figure 12). Se-

conda Angostura, eastern end of Estrecho de Magallanes.

Magallanes Province, Chile (.52.6°S, 70.2°W); Albatross

stn. 2779; 20 fms. [37 m]. on gravel.

Distribution: La Paloma. Rocha Department. Uru-
guav (34.6°S) (Figueiras and Sicardi, 1970b); Mar del
Plata. Buenos Aires Province (38.0°S) (Castellanos. 1970:
279; MACN 10757, not seen); Puerto Madmi. Golfo
Nuevo, Chubut Province (42.8°S) (ANSP 170466). and
Cabo Domingo. Tierra del Fuego Province (53.7°S)
(ANSP 236074). Argentina, in die Estrecho de Magal-
lanes, at 52.6°S (USNM 110719). and nordi in the east-
ern Pacific to Isla Chiloe. Chiloe Province. Chile
(approx. 43°S) (Figure 13) (SBMNH 144635). The only
depth record available is that with the hpe specimen of
S. snbequalis, 37 m. The material that has been illus-
trated and that I have studied is of similar shape, sug-
gesting that the species occurs in relativelv soft substrata.
Records from north of the Golfo Nuevo — the Uniguav-
an and Mar del Plata specimens cited above — probablv
represent sporadic larval settlement. Only a single east-
em Pacific lot has thus far come to my attention, and 1
have been unable to discover what material Bernard
(1983) relied upon for his eastern Pacific record of this
species.

Discussion: The largest species of the genus. S. hat-
cheri is most similar to S. luticola. The former differs in
having a thinner, more regular shell with a broader pos-
terior end. The chondrophore in the left valve projects
more toward the right valve, with the resilifer in the
right valve more deeply recessed. The posterior ridge
does not extend as far beyond the v entral edge of the
chondrophore.

Sphenia luticola (Valenciennes. 1846)
(Figures lb, 14-17, 22)

Corhula luticola Valenciennes, 1846; pi. 24. fig. 6. 6a; Carpen-
ter 1864: .543 (as Sphenia, and as a possible senior svno-
nv-m of S, fragilis): E. A. Smith. 1893: 279; Lamy, 1927:
179-180 (as Sphenia and as senior sviionvin of S. fragiUs.
but under the latter heading); Hertlein and Grant, 1972:
321-322, pi. 56, figs. 21, 22 (as "of. luticola")- Bernard,
1983: .58.

Sphenia pholadidea Dall, 1916a: 41. nomen nudum: 1916b:
415; Oldroyd. 1925: 201; Lamy. 1927: 181; Bernard. 1983:
58 (as a sviionvin of S. luticola)

Cuspidaria (Tropidomija) nana Oldrovd. 1918: 28; Oldrovd.
1924: 99. pi. 13. figs. 8, 9; J. T Smith, 1978: 3.50; Bernard,
1983: 58 (as a sviionvm of S, luticola); Coan and Kellogg,
1990: 178.

Sphenia glohula Dall, 1919: 370-371; Oldrovd, 1925: 201;
Lamy 1927: 181; Schenck and Keen, 1940:' ,391, pi. 2, figs.
4-7;'Schenck, 1945: 517, 519, pi. 67, figs. 5-8; Woodring
and Bramlette, 1951: 66, 91; J. T. Smith. 1978: .340: Ber-
nard, 1983: 58 (as a svnonvm of S. luticola).

Description: Shell relativelv thick, highly variable in
shape, from oval to elongate; right valve slightlv more
inflated than left; anterior end short, broadlv rounded;
tapering in thick-ness evenlv from anterior to posterior
ends; posterior end from short to somewhat produced,
and from truncate to somewhat pointed, sometimes
hvisted in various directions depending on crevice or
hole inhabited, generall)- extended bv" outer shell layer
and periostracum, scarcely gaping. External surface with
fine commarginal ribs; periostracum hght to dark tan,
trequentlv eroded in large specimens. Left valve with
large, heaw chondrophore, projecting more ventrallv
than toward right valve; posterior and anterior buttresses
of chondrophore short, thick; posterior ridge often pro-
jecting well bevond ventral margin of chondrophore.
Right valve with large, heaw, rounded resilifer under
beaks, not deeply recessed, and sometimes with slightly
projecting anterior cardinal tooth. Anterior adductor scar
large, positioned ventrallv. Posterior adductor scar and
palhal sinus near posterior end. Palhal sinus of hvo rel-
ativelv broad, elongate scars, and generallv also with one
or two smaller scars; palhal line a series of small, dis-
continuous scars (figure 22). Length to 20.6 mm (para-
tvpe of C. nana).

Type material and localities:

C. luticola — MNHN, lectohpe herein designated, the larger of
two pairs, likel)' that figured bv Valenciennes (1846: pi.
24, fig. 6a) as an opened specimen; length, 10.7 mm;
height, 6.3 mm; thickness, 6.0 mm (figure 14). Paralec-
totvpe, pair, perhaps that figured by Valenciennes (1846;
pi. 24, fig. 6) as a closed pair in situ in a rock crevice;
length, 9.3 nun. LociJitv not given in original account. The
label says "Califoniie." to which someone has added "Ma-
zatlan." The latter is undoubtedlv incorrect, and the lo-
calitv is here restricted to Monterev Bay. Monterev Coun-
ts'. California (36.7°N). from where other of Valenciennes'
material of bivalves boring or nestling in dredged rocks
seems to have come.

S. pholadidea — USNM 2581, holotype, pair; lengdi, 13.1 mm:
height, 6.0 mm; thickness, 5.1 mm (Figure 15). Santa Bar-
bara, Santa Barbara Count)-, California (34.4°N); WiHiani
Rich.

E. V. Coan, 1999

Page 113

Figures 14-17. SplwnUi luticola. 14. LectoUix- of Corhiila luticola MNHN lendth - in? ■„„, iix w I », c c i, ■

Page 114

THE NAUTILUS, Vol. 113, No. 4

18

19

20

21

o

22

Figures 18-22. Comparative pallial lines, muscle scars and hinges. 18. Sphcniafra^iUs. SBMNH 14463() Pnerto PeMsc.o, Sonora,
Mexico length = 9.1 mm. 19, 20. Sphcnia ffdfcms new species. 19. Holot>pe, SBMNH 14462., lett valve length- ll>""m . 20-
Paratvpe SBMNH 144628, nght valve, len|ih = 10.6 mm. 21. Sphrnia hntchcri. SBMNH 1446,3.5. Isla Chiloe, Chile, length -
,30.1 mm. 22. Sphenia luticola, LACM 60-22, Monterey, California, 18 m, length = 14.1 mm.

E. V. Coan, 1999

Page 115

C nana— CAS 060981, holohpe, pair: length, 19.4 mm;
height, 11.1 mm; thickne.ss, 10.1 mm (Figure 16). AMNH
58.306, paratvpe, 20.6 mm. Monterey Bay, Monterey
Counbv-, Cahfoniia (36.7°N).

S. glubnla — USNM 218977, lectotvpe herein designated, open
pair; length, 12.8 mm; height, 8.7 mm; thickness. 8.4 mm,
the only specimen cited in the original acconnt (Figure
17). CAS 064102, open pair, paralectotype; length, 14. .5
mm. CAS 08.5834, closed pair, paralectohqie; length, 9.7
mm. Bolinas Bay, Marin Connb,-, California (37.9°N);
Henn- Hemphill. The last specimen was figured by
Schenck and Keen (1940) and Schenck (1945) and may
also be the Bolinas specimen of Cuspidnria nana referred
tob\ OldroN-d (1918).

Distribution: West of Destruction Island, Jefferson
County, Washington (47.7°N) (SBMNH 140087), south
to San Diego, San Diego Countv, California (.32.8°N)
(USNM .322064, .32206.5'), from L5 to 155 m (mean, 50
m; n = 15), nestling in holes in rock and shell. I have
seen 34 Recent lots. This species has also been recorded
in the Pliocene Foxen Mudstone of Santa Barbara Coun-
t>' (Woodring and Bramlette, 1951: 66, 91, as S. globula)
and San Diego Formation of San Diego Count)' (Hert-
lein and Grant, 1972: 321-322, pi. .56, figs. 21. 22, as
"cf luticola"). California.

Discussion: The t)pe specimens of this species and its
SMiomins demonstrate the potential of species of Sphc-
nia to assume a varieh' of shapes. Nestling in pre-estab-
lished cavities in subtidal rocks, the shell conforms to
the shape of the available space in the same manner as
the European Miocene S. anatina discussed bv Baluk
and Radwariski (1979). The lectotvpe oi'Corbula luticola
and the holohpe of Cuspklaria nana are inflated and
acuminate. The holot\pe of Sphcnia pholadklca is flat-
tened and truncate posteriorly, and the lectotype of S.
globula is oval and inflated.

A small specimen (CAS 111000; Monterey, California:
length, 5.8 mm) is regular in outline and only somewhat
longer posteriorly. It is similar to juvenile Mya, but is
easily distinguished by its shallow pallial sinus composed
of broad pallial scars, as well as by features of its chon-
drophore, such as those outlined by Hanks and Packer
(1985: 324) for their new species of Sphcnia. In this
small Monterey specimen, the chondrophore projects
more horizontally and the resilifer in the left valve is
more deeply recessed than those of large specimens,
demonstrating the morphological change with growth.

Sphenia luticola and S. fragilis overlap in southern
California. The former can be distinguished by its thick-
er shell, more ventrallv directed chondrophore, with a
larger, often projecting posterior ridge, and a pallia) line
formed by larger scars. It also occurs offshore and attains
a larger size.

Of the eastern Pacific taxa, S. luticola is the most sim-
flar to the type species of the genus, .S. binghami. which
has a more regularly cjuadrate shape, with a broadly
truncate posterior end, and its chondrophore is thinner
more horizontal, and does not have a projecting poste-
rior ridge (USNM 171240 [lectotype], 171245. ,3.372,53).

Its pallial sinus is composed of narrower scars, and at-
tains only about 10 nun.

OTHER TAXA

Several fossil taxa of Sphcnia have been named in North
and Central America. While it is beyond tlie scope of
the present paper to anal)'ze their relationships, I list
them here to facilitate access to this literature.

Sphcnia scntcifciti Gardner (19.36: 45, pi. 8, figs. 2-
4). described from the Miocene Oak Grove Formation
of the Alum Bluff Group of Florida, has an obfique out-
line, a broadk- flared posterior end, and a relatively
smafl, horizontal chondrophore [USNM 372905], Sphe-
nia (hibia (Lea. 1843: 2) [originally as Mija clubia] from
the Pliocene Yorktowm Formation of Virginia is relatively
thick shelled, variable in shape, but generally short and
oval, and it has an especially large chondrophore in the
left valve and resilifer in the right; Mi/a rcflcxa Lea
(1843: 2) is regarded as a .s\Tionym (Campbell' 1993: 47,
213. fig. 187. 187a) [see also Lea. 1845: 236-237, pi. .34,
figs. 9. 10] [ANSP 1588, USNM 40.3.361, SBMNH
144632]. Sphcnia attcnuota Dall (1898: 860, pi. 35, fig.
9) [USNM 107806], from the Plio-Pleistocene Caloosa-
hatchee Formation of Florida, is ver\' similar to S. dubia
and is perhaps a svnonyTn of it.

Sphcnia wallacci Dall (1912: 3), described from the
late Pleistocene of Panama, seems to have been over-
looked since it was proposed. The tvpe material [USNM
214354] consists of a large right valve (length, 10.3 mm),
a medium-sized left valve (length. 9.1 mm), and a smafl
left valve (length, 3.7 mm). The right valve, which would
be the logical candidate for an eventual lectotspe des-
ignation, is oval and inflated and seems to be same thing
as die later-named S. tumida Lewis (1968: 26-30. pi. 1,
figs. 1-4) [USNM 64.566.3-645667] from the Pleistocene
of Florida. The larger left valve in the type lot of S.
wallacci is elongate antl mav be S. fragilis; the other left
viJve is too small to identify. Sphcnia tuinida was re-
ported from a Texas beach, although it was thought to
have possibly been a fossil (Ode and Speers. 1971: 71;
Ode. 1971: 74-75). However Sphcnia tumida has been
reported and figured as living on the coast of Yucatan,
Mexico (Yokes and Yokes, 1983: 44, pi. 48, fig. 16). Thus,
there may be a second living species of Sphcnia in the
western Atlantic, which should be called S. wallacci.

EXCLUDED TAXA

Sphcnia hilirata Gabb (1861: 369) was described from
Santa Barbara, California, presmnablv from the Tertiary,
as is implied by the paper's title ( "Descriptions of new
species of American Tertiary Fossils and a new Carbon-
iferous Cephalopod from Texas"). Sphcnia hilirata was
not figured, and the tvpe material has never been locat-
ed. There must have been more than one original spec-
imen, because the diameter was said to be "variable." As
has been noted by previous workers (Carpenter, 1864:
632 [1872: 118]; Lamy 1927: 177), the description close-

Page 116

THE NAUTILUS, Vol. 113, No. 4

Figure 23. Mi/a ?nrennria Linnaeus, 17.58. Holotype of Sphenia ovoidea, USNM 4.5.52, lengtl

1, I .o mm.

ly matches small Hiatclla arctica (Linnaeus, 17.58). How-
ever, Hiotclla has not been recorded from the Santa Bar-
bara Formation (Arnold. 1907: .32; Dibblee, 1966: .58),
the outcrop of which at Bathhouse Beach is the most
likel)' loc;Jit\- for the rest of the Santa Barbara taxa de-
scribed in Gabbs paper^. Additional study of the Santa
Barbara Formation is needed, because its fauna remains
poorK' documented. It is more likelv that S. biliratci
came from the Recent fauna, because, while it is re-
corded by Gabb simply as "Santa Barbara Cal.," each of
the other Santa Barbara fossil t;ixa also adds "Miocene"
or "Miocene?" (The Santa Barbara Formation is now
regarded as being of earK Pleistocene age.)

Sphenia californica Conrad (1837: 2.34, pi. 17, fig. 11)
is the original combination for Cn/ptomi/a californica
(Conrad. 18.37) (Keen, 1966: 171). '

Sphenia? mc^anosensi.s Clark and Woodford (1927:
106, 134. pi. 18, figs. 15, 16), from die lower middle
Eocene Meganos Formation of central California, is
based on two closed specimens [UCMP 31326, 31327]

' Another Santa Barbara bivalve described in Gabbs paper.
Cardita inonilico.sta (p. .371). has been snbse(juentlv recorded
Ironi this formation. A third Santa Barbara bivalve, Venus rhij-
somia (pp. 369-370), which has been regarded as a svmonviii
of Ntitricola tantiHa (Gould. l.S.53:406) (Caqienter 1864:632:
S. Gray, 1982:108-109, as Transennelln- generic allocation:
Coan and Scott, 1997:14, 25), has not been recorded from the
Santa Barbara Formation, but N. barbarensis (Arnold, 1907:
440-441, pi. 58, fig. 3, as Psephidia) is so recorded, and I dius
suspect that V. rhtjsomia is a senior synonvm of Arnold's ta.\on
instead. The two gastropods described from the same station
have subsequently been recorded there, both now regarded as
junior ,svTion\ins of other taxa.

said to resemble material of Sphenia from the Eocene
of the Paris Basin. No additional specimens of this spe-
cies have been recorded, nor has the genus been re-
ported in anv other strata earlier than the Pliocene in
western North America. Until more material becomes
available and a hinge e.>qDosed, one cannot be certain that
this material represents Sphenia.

Sphenia ovoidea Carpenter (1864: 602, 637 [1872: 88,
123], 1865: .54) was described from a single specimen,
paired valves obtained in Puget Sound from a fish stom-
ach in the 1860s. (Sphenia ovalis Carpenter, 1864: 682
[1872: 168], is a typographical error for ovoidea in the
same work.) Although previous workers have listed it as
a Sphenia, examination of tlie holot\pe [USNM 4552;
length, 7.5 mm; height, 4.5 mm; thickTiess, 2.8 mm]
(Figure 23)^, which was well figured bv Palmer (1958:
116, .340, pi. 15, figs. 7-10), confirms C'aqienter's "man-
tle bend large" (i.e., a large pallial sinus), demonstrating
tliat it is a juvenile Mija, not a Sphenia. Juvenile Mt/a
are most easilv distinguished from Splienia bv' their larg-
er, continuous pallial sinuses. This specimen appears to
be Ml/a arcnaria Linnaeus (1758: 670); it presents a
quandars' because this species, having become extinct in
the Pacific Northwest by the Pleistocene (Carlton, 1979:
518-.543, 1992: 495), was first recorded in the 1880s,
when it became reestablished through introduction from
Atlantic popidations with imports of Crasso.strea virgi-
nica (Gmelin, 1791). The only Mija thought to have been
present in the 1860s is Mija tnincata Linnaeus (1758:

■ A broken valve in a separate capsule with the hoiohpe is an
unrekited species, most likely a Ivonsiid, mixed into the lot
between the time of Garpenter and that of Palmer

E. V. Coan, 1999

Page 11"

Figure 24. Sphenia fragilis. Lectot\pe ot Corbula iheiingi-
ana Pilsbry-, 1897; ANSP 70541, length, 9.0 mm.

670)'\ but small specimens of that species are very dif-
ferent from those of areuaria. It is possible that: (1) the
specimen might not actually have come from Fuget
Sound, although all the other material collected there bv
C. B. R. Kennerley of the Northwest Boundan- Survey
has proven to authentic; or that (2) Mija arenaria was
actually present in Puget Sound 20 years earlier than
previously thought as a result of introduction with oys-
ters or ship fouling (J. T. Carlton, e-mail, 19 November
1997).

ACKNOWLEDGMENTS

I appreciated the assistance of curators and others, who
have allowed me to visit collections, to borrow speci-
mens, and provided needed information. These include
Elana Benamv, Ned Gilmore, and Gar)- Rosenberg,
Academy of Natural Sciences, Philadelphia: Philippe
Bouchet and Philippe Maestrati, Museum National
d'Histoire Naturelle, Paris; Steven Coles, Bishop Mu-
seum, Honolulu, Hawaii; Jay Cordeiro, William K. Em-
erson, American Museum of Natural History, New York;
Amanda Diaz and Nancy Voss, University of Miami;
Warren C. Blow, Raye Germon, Alan R. Rabat, Guido
Pastorino, and Thomas R. Waller, National Museum of
Natural History, Washington, DC; Cristian F. Ituarte,
Museo de La Plata, Buenos Aires, Argentina: Lindsey
Groves and James H. McLean, Natural Histoiy Museum
of Los Angeles Count\'; Elizalieth Kools and Robert Van
Syoc, California Academy of Sciences, San Francisco;
Gustav Paulay, Universits' of Guam; Paul Valentich Scott,
Santa Barbara Museum of Natural Histon'; Kathie Way
and Joan Pickering, The Natural Histor)' Museum, Lon-
don; and Karen Wetmore, Museum of Paleontology,

'The second species of Sphenia described by Turton (1822)
along with the genus and S. biiighami. its type species, was S.
sivainsoni (pp. .37-38, pi. 2, fig. 3, pf 19, fig. 2). It was regarded
as having been based on a juvenile Mya tmncata by MacNeil
(196.5;.38).

Uni\ersit\' of California, Berkeley. Carol C. Skogiund
generousK made available material from her collection,
as did the late Elsie Marshall. David B. Packer of the
National Marine Fisheries Service, Highlands, New Jer-
sey, provided specimens of Sphenia sinccra, and Lyie
Campbell, of Spartanburg, South Carolina, provided
specimens of the Pliocene Sphenia diibia and the Recent
S. antillensis. Osmar Domaneschi of the Universidade
de Sao Paulo provided Brazilian specimens of S. antil-
h^nsis. I obtiuned advice from Roland Anderson, Seattle
Aquarium: [ames T Carlton, Williams College, Mvstic,
Connecticut: Nora Foster Universits* of Alaska: Akihiko
Matsukuma, Kvxishu University', Fukuoka, Japan; Fabri-
zio Scarabino, Museo Nacional de Historia Natural,
Montevideo, Uniguav; and Richard L. Squires, Califor-
nia State Universih, Northridge. Lvle Campbell, Osmar
Domaneschi, Lindsey T Groves, Paula M. Mikkelsen,
Walter Narchi, David B. Packer, Paul Valentich Scott,
and Carol C. Skogiund provided useful comments on the
manuscript. Sharon Williams helped to prepare the
plates.

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Adams, A., 18.51. Monograph of Sphnenin. a genus of lamelli-
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Adams, H. and A. Adams. 1854. Description of a new genus
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Adams, H. and A. Adams. 18.56 [185:3-1858]. The Genera of
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Baluk, W. and A. Radwaiiski. 1979. Shell adaptation and eco-
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Basterot, M. 1825. Memoire Geologique sur les Environs de
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Bernard, F. R. 1983. Catalogue of the living Bivalvia of the
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+ 102 pp.

Brann, D. C. 1966. Illustrations to "Catalogue of the Collection
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Campbell, L. D. 1993. Pliocene molluscs from the Yorktown

Page 118

THE NAUTILUS, Vol. 113, No. 4

and Chowan River formations of Virginia. Publication of
the Department of Mines, Minerals, and Energ); Division
of Mineral Resources, Commonwealth of Virginia 127, vii
+ 259 pp., 43 pis.

Carcelles, A. R. 1950. Catalogo de los nioluscos marines de la
Patagonia. Anales de la Museo Huapi 2:31-100, 6 pis., 1
map.

Carcelles, A. R. and S. I. Williamson. 1951. Catalogo de los
moluscos niarinos de la Provincia Magallanica. Rexista
(Ciencias Zoologicas), Institute Nactional de Investigacion
de las Ciencias Naturales 2(5):22.5-283.

Carlton, J. T. 1979. Histor)-, Biogeography, and Ecology of the
Introduced Marine and Estuarine Invertebrates of the Pa-
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THE NAUTILUS 113(4):121-126, 1999

Page 121

Two new species of the genus Chicoreiis (Stratus) (Gastropoda:
Muricidae) from the western Atlantic.

Roland Houart

Institut Royal des Sciences Natuielles

de Belgique
Rue Vautier, 29
1000 Bnixelles
BELGIUM

ABSTRACT

Two new species of the family Muricidae are described. Chi-
coreiis (Siratus) colellai, from off Veracruz, eastern Mexico, is
compared with C. (S.) articulatus (Reeve, 1845), and differs
from this latter species in manv aspects. C. (S. ) caudacurta.
from off Puerto Rico, is somewhat similar to C. (S.) bentiii
(Fischer & Bernardi, 1857), but differs in having a broad, pau-
cispiral protoconch, a higher spire, and a shorter siphonal ca-
nal.

Key words: Gastropoda, Muricidae, Chicoreiis (Siratus). west-
em Atlantic, Mexico, Puerto Rico, new .species.

INTRODUCTION

At least 19 Recent species of Chicorcus (Siratus) are
known, of which onl\' 2, C. (S.) alabaster (Reeve, 1845),
and C. (S.) pliciferoiclcs Kuroda, 1942, occur in the Indo-
West Pacific region (Houart, 1992), and 17 in the west-
em Atlantic. The western Atlantic species have been re-
viewed and illustrated in several works (Fair, 1976; Rad-
win & D'Attilio, 1976; Rios, 1985; Yokes, 196,3, 1980,
1990, 1991). Additional names have been proposed, but
are synonymized under the 17 taxa herein recognized.
The Recent western Atlantic species of Siratus are listed
in Table 1.

1

Figures 1, 2. Protoconchs. 1. Chicoreiis (Siratus) caudacur-
ta new species, off Veracruz, dredged in 400-500 m, paratype
R. Houart Collection. 2. C. (S) beauii (Fischer & Bernardi,
1857), Florida, off Egmont Key, 7.3 m, R. Houart Collection.
Scale bar = 0.5 mm.

Recently, Jose Coltro (Sao Paulo, Brazil) sent me sev-
eral lots of Chicoreiis (Siratus) dredged from deep water
in different regions of the Caribbean Sea. Within these
lots, 1 immecliately noticed the presence of unusual
forms. Some of these were identified as representing ex-
tremes of variation of known species, but two remained
unidentified. One, C. (S.) colellai new species, differs
from all Recent or fossil species. The other, C. (S.) cau-
dacutia new species resembles a small, spineless C.
beauii, but the protoconch whorls are distinct in the two
ta.\a. Abbreviations and text conventions are: dd, empty
shell; MORG, Museu Oceanografico "Prof. Eliezer de
C. Rios", Funda^ao Universidade do Rio Grande, Rio
Grande do Sul, Brazil.

SYSTEMATICS

Family Muricidae Rafinesque, 1815
Subfamily Muricinae Rafinesque, 1815
Genus Chicorcus Montfort, 1810

Type species: Murex ramosus Linnaeus, 1758; Re-
cent, Indo-West Pacific, by original designation.

Subgenus Siratus Jousseaume, 1880

Type species: Puqmra Sirat "Adanson" Jousseaume,

ISSO (= Murcx senegalcnsis Gmehn, 1791), Recent,
Brazil, bv original designation.

Chicorcus (Siratus) colellai new species
(Figures 3-5)

Description: Shell medium sized for die subgenus, up
to 47 mm in length at maturity (holotype), weakly spi-
nose, nodose, lightly built. Spire high, up to 6 broad,
convex, weaklv spinose teleoconch whorls. Suture im-
pressed. Protoconch imkiiown (broken).

Axial sculpture of teleoconch consisting of 3 low, nar-
row, rounded, weakly spinose varices from first to last
whorl. Other axi;il sculpture of low, narrow, rounded,

Page 122

THE NAUTILUS, Vol. 113, No. 4

Table 1. Geographic and batlmiietric range of western Atlantic species of Stratus
considered as valid):

ipecies (species names printed in boldface are

Name of species

Geographical
distribution

Bath\iiietric range
(live specimens)

Chicoreus (Sirattts) aguayoi (Clench & Perez Farfante,

194.5)
C. (S.) articulatus (Reeve. 1845)
Murex nodatus Ree\e, 184.5 (not Gnielin, 1791)
Murex gundlachi Dunker, 188.3
Murex finlayi Clench, 1955
C. (S.) beauii (Fischer & Bemardi, 1857)
Murex percoidcs Lobbecke, 1879
Murex bronchi Clench, 195.3
C. (S.) cailletti (PeHt. 18.56)
Murex siinili.s Sowerb\-, 1841 (not Schroter, 1805)
Murex kugleri Clench & Perez Farfante, 194.5 (new name

for M sitnilis Sowerbv, 1841, not Schroter)
C (S.) carolynae Yokes. 1990

C (S.) cibonetj (Clench & Perez Farfante, 1945)
Murex trilinentus Reeve, 1845 (not Sowerbv, 1813)
Chicoreus reevei Vokes, 1965 (new name for M. trilineatus

Reeve, 1845, not Sowerbv)
.\/((;y.v t/iimuriniis Sarasua & Espinosa, 1978
C. (S.)coUrortim Vokes, 1990
C. (S.) consuehi (Verrill. 19.50)
Murex pidcher A. Adams, 1853 (not Sowerb)', 1813)
Murex pulcher var consuela Verrill, 1950
C (S.) formosiis (Sowerbv, 1841)
C. (S.) motacilla (Gmelin, 1791)
Murex hriskasii Wrrill, 1953
C. (S.) perelegans Vokes, 1965

Murex elegans Beck in Sowerb\, 1841 (not Dono\'an, 1804)
Chicoreus perelegans Vokes, 1965 (new name for M. elegans

Beck in Sowerbv, 1841. not Wood, 1828)
C (S.) senegalensis (Gnielin, 1791)
Murex costatus Gmelin, 1791
Murex brasiliensis Sowerbv, 1834
Purpura sirat dOrbigny, 1841
C. (S.) springeri (Bull'is, 1964)
C (S.) tenuivaricosus ( Dautzenberg, 1927)
Murex calcar Kiener, 1843 (not Scacchi, 1835)
Murex tenuivaricosus Dautzenberg, 1927 (new name for M.

calcar Kiener, not Sowerbv)
Chicoreus carioca Vokes, 1968 (new name for calcar Kiener,

1842)
C. (S.) thompsoni (Bullis, 1964)

C (S.) colellai new species

C (S.) caitdacurta new species

Baliamas to Cuba

E Florida to Ceara, Brazil

Florida to Uniguay

Florida to Honduras, Colombia, and
Barbados

Biiliia, Brazil

Cuba to Lesser Antilles

338^521 m
16-^00 m

7:3-463 m
46-400 m

Unknown (probabK- 10—40
ni, R. Houart collection)
110-155 m

Northern Brazil

2-100 m

Gulf of Mexico to

Lesser .\ntilles

14-110 m

Jamaica and Cuba

20-100 m

Lesser Antilles

170 m

Haiti to Barbados

100-120 m

Surinam and Brazil (Espi'rito Santo
to Santa Catarina)

Northern South .\merica
Brazil (Espi'rito Santo to Santa Cata-
rina)

Surinam, Venezuela, Brazil (Amapa

to Baliia)
Off Puerto Rico

Mexico, off Veracniz

2-15 m

20
2-40 m

L8-40 m

Unknowni (914-1097 m for

eniph' shells)
Unknouii (400-500 m for

enipt\' shells)

inter\'arical ribs: 2 from first to third whorl; 2 or 3 on
fourth: 3 on fifth: 3, occasionally with low fourth one,
on last whorl. Spiral sculpture of low, narrow, smooth,
primary, secondary', and tertiary cords. Spire whorls with
2 or 3 primar)' cords, and one secondary' cord in each
interspace; penultimate with 4 primary cords, and one
secondarv cord in each interspace: last whorl with 5 pri-

mary cords that interconnect strongest varical spines,
with secondar}' and tertiary' cords in each interspace.
Cords more obvious on axial varices. Shoulder of last
whorl with 4 or 5 spiral cords. Aperture large, broad,
rounded. Columellar lip narrow, with 3 weak kmobs
abapically. Rim partialK' erect, adlierent at adapical ex-
tremitv. Ana! notch broad, deep. Outer lip weakl\' erect.

R. Houart. 1999

Page 123

Figures 3-7. New species oi Chicorens. 3-5. C. (Siratus) colellai, t\pe localitv' (off Puerto Rico, 914-1097 m). 3. 4. Holotype,
MORG 39519, 47 mm. 5. Paratope, R. Houart Collection, 42.9 mm. 6, 7. C. {S.j caudaauia Holohpe MORG .39520, t\pe lociility
(off Veracmz, 400-500 m), 45.8 mm.

Page ]24

THE NAUTILUS, Vol. 113, No. 4

R. Hoiiart, 1999

Page 125

smooth within. Siphonal canal short, hut partially hroken
in all specimens, \veakl\- annulate, open, with 1 or 2 short
spines adaperturalK', and 2 or 3 spiral cords. Shell white,
with presence of hrown iilotches on 2 or 3 prinian'
cords. Operculum and radula unknown.

Type material: HolotApe, MORG 39519, dredt;ed
(dd); paratopes (From tApe localitA), R. Houart collection
(Ezemaal, Belgium), 1 parat\pe; J. & M. Coltro collec-
tion (Sao Paulo, Brazil), 1 paratope.

T>pe locality-: Off Puerto Rico, 914-1097 m.

Etymology: Named in honor of Mr Julio ("olella. one
of the founding members of the two Brazilian shell-re-
lated organizations, "Sociedade Brasileira de Malacolo-
gia" and "ConquiUoIogistas do Brasil".

Discussion: C. colcllai new species differs from all
other species of Chicorciis fS'(raf«,s) in having rounded,
broad, last teleoconch whorl, rounded aperture, narrow
varices, and short siphonal canal. C. colcllai is most
closely related to C. (S.) articulatiis (Figs 8-10), a spe-
cies with \er\' variable shell, however. C. colcllai differs
in having more vvidelv spaced, narrower, a\ial ribs, nar-
rower varices, broader, more rounded aperture, and nar-
rower (relative to the shell width) siphonal canal.

Chicoreus (Stratus) caudacuiia new species
(Figures 1, 6, 7)

Description: Shell medium sized for subgenus, up to
50 mm in length at maturitv, lanceolate, weakly spinose,
lightly built. Spire verv high with 1.45 protoconch
whorls, and up to 7 conve.x, low, weakly spinose teleo-
conch whorls. Suture impressed. Protoconch large,
broad, irregularly shaped, smooth. Terminal varix vm-
known (eroded). Axial sculpture of teleoconch consisting
of low, narrow, weakly spinose varices. Other axial sculp-
ture of weakly nodose, intervarical ribs. Shoulder spine
longest. First whorl with 15 ribs, second with 14, third
to last whorl with 3 varices and 3 or 4 intervarical ribs.
Spiral sculpture of primary, and secondary', low cords
and threads. First to third whorl with 4 primaiy cords
and 1 or 2 secondar)' cords; fourth and fifth with 6-8
cords and threads; last whorl with 13-15 cords and
threads; shoulder with 3 or 4 cords. Aperture large,
rounded. Columellar lip narrow, with 2 or 3 weak, low
knobs abapicallv. Rim partially erect, adlierent at adap-
ical extremity. Anal notch narrow, deep. Outer lip weakly
erect, smooth within. Siphonal canal short in examined
specimens, narrow, open, weakly angulate, with 3 or 4
low, spiral cords, and one short spine adaperturally. Shell

ivorv'-white or creamy-white, with brown tinge on some
spiral cords, more apparent on two cords abapicallv of
shoulder, two on periphery, and two above siphonal ca-
nal. Aperture white. Operculum and radula unknown.

Type material: Holohpe MORC 39520. dretlged
(dd(; R. Houart collection, 1 paratvpe, from tspe locality.

Type locality: oil Veracruz, eastern Mexico, 400-500
m.

Etymology: Latin for "short tail", alluding tf) its short
siphonal canal.

Discussion: Chicoreus (Stratus) caudacuiia somewhat
resembles C. (S.) bcauit (Figures 2, 11, 12) from which
it differs in having apparently different larval develop-
ment. Chicoreus (S.) caudacurta has large, irregularly
shaped, broad protoconch of 1.45 whorls (Figure 1),
which suggest non-planktotrophic and possibly intracap-
sular development, while C. (S.) hcauti has conical pro-
toconch of 34- whorls and small embrvonic whorl (Fig-
ure 2), which suggests planctotrophic larval develop-
ment. Chicoreus (S.) caudacuiia also has a higher spire
relative to the height of the aperture, and the length of
the siphonal canal. Chicoreus (S.) caudacurta differs
from C. (S. ) cailleti (Petit de la Saussave. 1856) (Figure
13) by higher spire, shorter siphonal canal, broader ap-
erture, and protoconch twice as large. The new species
differs from the 'form" C. kugleii (Clench and Perez-
Farfante, 1945) in having shallower, more numerous,
narrower spiral threads and cords, higher spire relative
to the height of the aperture and the length of the si-
phonal canal, and a protoconch twice as large.

ACKNOWLEDGMENTS

I am very grateful to Jose and Marcus Coltro, Sao Paulo,
Brazil, who provided this very interesting material. 1 am
also thankful to Dr Emily H. Yokes, Tulane University
(retired), for her constructive comments on the manu-
script. Thanks are due also to two anonymous referees
for additional remarks.

LITERATURE CITED

Fair. R. H. 1976. The Mtirex Book, an illustrated catalogue of
Recent Muricidae (Muricinae, Muricopsinae, Ocenebri-
nae), Stiirgis Printing Co.. Honolulu. 138 pp.

Houart, R. 1992. The genus Chicoreus and related genera
(Gastropoda: Muricidae) in the Indo-West Pacific. Me-
nioires du Museum national d'Histoire naturelle (A) 154:
1-188.

Radwin G. and A. D'attilio, 1976. Miircx shells of the world.

Figures 8-13. Other species of Chicoreus. 8-10. C (Siratus) aiiiculatus (Reeve, 1845). 8. Off Haiti. R. Houart Collection. 52.1
mm. 9. Off Honduras, 200-300 m, R. Houart Collection, 50.4 nun. 10. Off Paramaribo, Surinam, .300-400 m. R. Houart Collection,
32.2 mm. 11, 12. C. (S.) hcmiii (Fischer & Bemardi. 1857). 11. Bahia de Campeche, otTVeracniz, eastern Mexico, 200-300 m,
R. Houart Collection, 108.2 mm. 12. Off Egmont Key, Florida, 73 m, R. Houart Collection. 56.5 mm. 1.3. Clticoreus (Siratus)
cailleti (Petit de la Saussaye, 1856). St. James, Barbados, 94 m, R. Houart Collection, 52.7 mm.

Page 126

THE NAUTILUS, Vol. 11.3, No. 4

An illustrated guide to the Muricidae. Stanford University
Press, Stanford, 284 pp.

Rios, E. de C. 1985. Seashells of Brazil, Funda^ao Universi-
dade do Rio Grande, Museu Oceanografico, Rio Grande,
Brazil, 329 pp.

Voices, E. H. 196.3. Cenozoic Muricidae of the Western Atlantic
region. Part 1 — Miirex sensu stricto. Tulane Studies in Ge-
ology and Paleontology 1:9.5-123.

Voices, E. H. 1980. What is my name? Or, will the real Murex
antillanim please stand up! Of Sea and Shore 11:91-92.

Vokes, E. H. 1990. Cenozoic Muricidae of the western Atlantic
region. Part VIII — Murex s.s., HaiistcUuin, Chicoreus,
Hcxaplcx; additions and corrections. Tulane Studies in Ge-
olog\' and Paleontologv" 23:1-96.

Vokes, E. H. 1991. The other species of Sirntus in the western
Atlantic. Siratus 8:7-13.

THE NAUTILUS 113(4): 127-150, 1999

Page 127

Harald A. Rehder (1907-1996): biographical sketch and
malacological contributions

Michael J. Sweeney
M. G. Harasewych

Department of Invertebrate Zoology
National Museum of Natural History
Smithsonian Institution
Washington, DC 20560-0118 U,SA

Dr. Harald Alfred Rehder, Zoologist Emeritus at the Na-
tional Museum of Natural Histors' of the Smithsonian
Institution, and a leading authority' on die s>stematies
and biogeographv of the niolluscan fauna of Polynesia,
died on November 10, 1996, of cardiac arrest. An emi-
nent scholar, he enjoyed a long and distinguished career,
and was highly respected by the international malaco-
logical community', as well as b\' the many amateur nat-
uralists interested in moUusks.

Harald Rehder was bom on June 5, 1905, in Boston,
Massachusetts, the son of a botanist and curator at Har-
vard University's Arnold Arboretum. He grew up in the
Boston suburb of Jamaica Plain, were he attended Rox-
bury Latin School. With the encouragement of his father
and of Charles W. Johnson of the Boston Societv of Nat-
ural History, his interest in natural history gradually be-
came focused on mollusks. After earning a Bachelor of
Arts degree in Chemistry from Bowdoin College in
1929, he entered Harvard University' as William J.
Clench's first graduate student in Malacology.

Prior to completing his degree at Harvard, he ob-
tained a position as a Senior Science Aide to Dr. Paul
Bartsch in the Division of Mollusks at the National Mu-
seum of Natural History in 1932. Harald Rehder re-
ceived his Master of Arts degree in Malacology from
Harvard University in 1933, and his Ph. D. in Malacol-
ogy from George Washingttjn Universit)- (Washington,
DC) in 1934. His dissertation, "A contribution to our
knowledge of the genus Succinea in North America " was
completed under the guidance of Dr Bartsch. After re-
ceiving his doctoral degree, he became an Assistant Cu-
rator in the Division of Mollusks in 1934, an Associate
Curator in 1942, and succeeded Paul Bartsch as Curator
in 1946. Dr. Rehder was named Senior Zoologist in
1965, and held this position until his retirement in June
1976. He continued, as Zoologist Emeritus, to publish
popular books and research papers on mollusks until his
death.

Dr Rehder devoted an extraordinary amount of time
and effort throughout his career to the care and curation
of the National Collection of Mollusks. During the Sec-
ond World War, Dr Rehder and Dr. J. P. E. Morri.son,
also of the Division of Mollusks, completed the daunting
task of identifying, documenting, and segregating thou-
sands of tvpe specimens of mollusks in preparation for

removal from the general collections to the caverns in
Luray, Virginia, for safekeeping. Upon their return to the
National Museum after the war, these collections re-
mained segregated as the type collection. Dr. Rehder
oversaw the move of the collections from the central
portion of the museum to the newly constructed re-
search wing in the 1960s, and continued to curate count-
less new acquisitions well into his retirement.

Dr. Rehder began his career working primarily on the
systematics of land and freshwater gastropods, but soon
expanded his work to include marine faunas. He took
part in expeditions to the Caribbean early in his career,
but his major research interest, the study of the niollus-
can faunas of the Central and South Pacific, developed
during his participation in the Smidisonian-Bredin Ex-
pedition to French Polynesia in 1957. He was a member
of the Pacific Science Board's expeditions to the Mar-
shall Islands in 1960, and led additional expeditions to
the South Pacific in 1963, 1964, 1967, 1969, 1970, and
1973. supported by grants from the National Geographic
Society, The National Science Foundation, and by pri-
vate donors. Dr. Rehder conducted field work at more
than fifty- islands and atolls during his career. The Na-
tional Collection was enriched by over thirteen thousand
lots and nearly one hundred thousand specimens that
were collected during these expeditions.

Dr. Rehder served as advisor to Helen L. Hayes, Jo-
seph C. Britton, and George Radwin, who completed
their doctoral degrees in malacology at George Wash-
ington University between 1968 and 1972.

Harald Rehder was the last surviving founding mem-
ber of the American Malacological Union (now the
American Malacological Society). He served as President
of this organization in 1940^1, was elected as Honorary
Life Member in 1978 and Honorarv' Life President in
1985.

Dr. Rehder was a Fellow of the American Association
for the Advancement of Science and the California Acad-
emy of Science, and a member of numerous professional
societies, among them the Washington Academy of Sci-
ences, the Biological Society of Washington, the Mala-
cological Society of London, the Paleontological Societ)',
the Societv for Systematic Zoolog); tlie American Littoral
Societv', the IntemaHoniil Societv for Reef Studies, and
the Senckenbergische Naturforschende Gesellschaft. He

Page 128

THE NAUTILUS, Vol. 113, No. 4

Figure 1. Dr. Harald Rehder. June 1971.

served as Editor of the Jounjal of the Wo.shiufSton Acad-
emy of Sciences from 1944—1946, as co-editor of Iiulo-
Pacific MoHusca from 1959-1973, and on the ecbtorial
board of numerous scholarK' publications.

He also belonged to man\ shell clubs, including Boston
Malacological Club, and the Hawaiian Malacological Soci-
ety. A Charter member of tlie National Capital SheO Club,
he was elected as its first president in November 1960.

In addition he was a member of the Cosmos Club,
The International Club of Washington, DC, The Ex-
plorers Club of New York City', and served on the Ad-
ministrative Board of the Foundrv United Methodist
Church and its Susanna Weslev House, and was \er\'
active in the Homeless Mission. He enjoved member-
ship in the "Fossils," an organization of distinguished
retired men.

Two genera and 24 species of Mollusca, 1 species of
Foraminifera and 1 species of Pvcnogonida have thusfar
been named in his honor.

His wife of 50 )'ears, Lois Corea Rehder, died in 1988.
He is survived by a daughter, Anne F. van Beek-Rehder,
of Amsterdam, The Netherlands; a son, Alfred L. Reh-
der of Germantown, Marvland, and three grandchildren.

n. MOLLUSCAN TAXA OF HARALD A. REHDER

During his career Harald Rehder introduced 397 mol-
luscan taxa, of which 58 were genus-level ta,xa (34 genera
and 24 subgenera, 1 subgenus as a replacement name),
and 339 were species-level taxa (320 as species, 19 as
subspecies, 8 species as replacement names). These are

Figure 2. Dr. Rehder at tlie .^imniai] Malacological Union
meeting in Fort Lauderdale. Florida, July, 1981.

listed alphabeticalK', by categorv. Data for genus-level
taxa include taxon, author(s), year of publication, journal
and page number, t>pe species, and famiK' to which cur-
rentk' assigned. Data for species-level taxa include taxon
(original combination), author(s), vear of publication,
journal, page number and illustrations, tvpe material,
tvpe repositorx' and catalog number, general t\pe locality,
and famiK'.

Genus-level Taxa

Aharhaiia Dall, Bartsch and Rehder, 1938, as a subgenus
of Barbatia Gra\', 1847. Bemice R Bishop Museum Bul-
letin 153:29. Type species: Barbatia (Abarbatia) oahtia
Dall, Bartsch and Rehder, 1938, bv original designation.
Arcidae.

Achasmca Dall, Bartsch and Rehder, 1938, as a subgenus
o{ Leiocliasmca Dall, Bartsch and Rehder, 1938. Bemice
R Bishop Museum Bulletin 153:147. T%pe species: So-
lecardia thaanumi (Pilsbry, 1921), by original designa-
tion. Galeommaddae.

Anfiuipccten Dall. Bartsch and Rehder, 1938. Bemice P.
Bishop Museum Bulletin 153:92. T\pe species: Angui-

M. J. Sweeney and M. G. Harasevwch, 1999

Page 129

Figure 3. Foniuil portrait ot Dr Rehder, circa 1994.

pccfen grcgpriji Dall, Bartsch and Rehder, 1938, h\ orig-
inal designation. Pectinidae.

Antcmctitia Rehder, 1943. Proceedings of tlie United
States National Mu.seuni 93(3 161): 199, T\pe species
Bitcciniiin inctiila Hinds, 1S44, by original designation.
Bnccinidae.

Barharca Dall, Bartsch and Rehder, 193S, as a siihgeniis
of Calloaiva Cray, 1857. Bernice P. Bishop Museum
Bulletin 153:23. Type species: CoHoarca {Barharca) luta
Dall, Bartsch and Rehder, 1938. In' original designation.
Arcidae.

Baiischia Rehder, 1943. Proceedings of the United
States National Museum 93(3161):199. Tvpe species:
Bartschia significrins Rehder, 1943, by original designa-
tion. Buccinidae.

Cadella Dall, Bartsch and Rehder, 1938. Bernice R
Bishop Museum Bulletin 153:196. T\pe species: Tclliiui
Icchiograiuma MeKille, 1893, by original designation.
Tellinidae.

CcrodriUia Bartsch and Rehder, 1939. Proceedings of
the United States National Museum 87(3070): 127. T\pe
species: Cerodrillia clappi Bartsch and Rehder, 1939. by
original designation. Turridae.

Coniittcrcdo Dall, Bartsch and Rehder, 1938, as a sub-
genus ot Teredo Linne, 1758. Bernice P. Bishop Muse-
um Bulletin 153:209. Tvpe species: Teredo iConiiitere-
do) inilleri Dall, Bartsch and Rehder, 1938, by original
designation. Terechnidae. [see comments under Teredo
milleri]

Cras.'-;i.'if)irella Bartsch and Rehder, 1939, as a subgenus
oi Crassispira Svvainson, 1840. Proceedings of the Unit-
ed States National Museum 87(3070): 135. Tvpe species:
Crassispira (CrassispireUa) nigitecta (Dall, 1918), by
original designation. Turridae.

Crifpfopeeten Dall, Bartsch and Rehder, 1938. Bernice
P. Bishop Museum Bulletin 153:93. Tyj)e species: Cn/p-
topeeten alii Dall, Bartsch and Rehder, 1938, bv original
designation. Pectinidae.

Dalliiiuirex Rehder, 1946, as subgenus of Paziella Jous-
seaume, 1880. The Nautilus 59(4):142. Tvpe species:
Miirex iiiiftingi Dall, 1896, by original designation. Mur-
icidae.

Desinaidirs Rehder, 1943, as a subgenus of Bieatillii.s
Swainsou, 1840. Proceedings of the Biological Societ^• of
Washington 56:45. Tvpe species: Calijptraea extincto-
rium Lamarck. 1822, bv original designation. Crepidu-
lidae.

Diploplax Bartsch and Rehder, 1945, as a subgenus of
Maiiesia Sowerby, 1S24, Smithsonian Miscellaneous
Collections 104(11): 10. Tvpe species: Maiiesia (Diplo-
plax) auiericaiia Bartsch and Rehder, 1945, In original
designation. Pholadidae.

Diploflu/ra Bartsch and Rehder, 1945, as a subgenus, of
Maiiesia Sowerby, 1824. Smithsonian Miscellaneous
Collections 104(11):8. Type species: Martesia (Diplotluj-
ra) sinifhii (Tnon, 1862), bv original designation. Pho-
latlidae.

Dorisca Dall, Bartsch and Rehder, 1938. Bernice R
Bishop Museum Bulletin 153:159. Tvpe species: Dorisca
cookei Dall, Bartscli and Rehder, 1938, by original des-
ignation. Veneridae.

Dysinea Dall, Bartsch and Rehder, 1938. Bemice R
Bishop Museum Bulletin 153:173. Tvpe species: Solen
occideiis Gmelin, 1791, bv original designation. Psain-
inobiidae.

Egentelaria Rehder, 1980. Smithsonian Contributions to
Zoolog)' 289:93. T)pe species: Terehra sti/lata Hinds,
1844, by originiil designation. Terebridae.

Gli/phepifhema Rehder, 1943. Proceedings of the Unit-
ed States National Museum 93(3161): 196. Tvpe species:
Natica idiopoma Pilsbi-v and Lowe, 1932, by original
designation. Naticidae.

Halopseplins Rehiler, 1943. Proceedings of the United
States National Museum 93(3161 ):191. Tvpe species:
Halopsepluis pideher Rehder, 1943, bv original desig-
nation, Turbinidae.

Page 130

THE NAUTILUS, Vol. 113, No. 4

Hautrwa Dall, Bartsch and Rehder, 1938. Bemice P.
Bishop Museum Bulletin 153:86. T\pe species: Haumea
juddi Dall, Bartsch and Rehder, 1938, By original des-
ignation. Pectinidae.

Hawaiarca Dall, Bartsch and Rehder, 1938. Bemice P.
Bishop Museum Bulletin 153:27. Type species: Ha-
waiarca rectangula Dall, Bartsch and Rehder, 1938, by
original designation. Arcidae.

Hitia Dall, Bartsch and Rehder, 19.38. Bemice P Bishop
Museum Bulletin 153:1.35. Type species: Hitia ovalis
Dall, Bartsch and Rehder, 1938, by original designation.
Sportellidae.

Kaneoha Dall, Bartsch and Rehder. 1938. Bernice P.
Bishop Museum Bulletin 153:142. Type species: Kaneo-
ha rosea Dall, Bartsch and Rehder, 1938, by monoUpy
Kelliidae.

Kona Dall, Bartsch and Rehder, 1938. Bemice P Bishop
Museum Bulletin 153:148. TyjDe species: Ki^na hucki
Dall, Bartsch and Rehder, 19.38, by original designation.
Montacutidae.

Kiirodina Rehder, 1969, as subgenus of Musa.shia Hay-
ashi, 1960. Venus, The Japanese Joumal of Malacology-
27(4):128. Type species: Musashia {Kiirodina) smithi
(Sowerby, 1901), by original designation. Volutidae.

Leiochasmea Dall, Bartsch and Rehder, 1938. Bemice P.
Bishop Museum Bulletin 1.53:146. T\pe species: Lei-
ochasmea chascax (Pilsbr\'. 1921), by original designa-
tion. Galeommatidae.

Lissodriiha Bartsch and Rehder, 1939, as a subgenus of
CerodiilUa Bartsch and Rehder, 1939. Proceedings of
tlie United States National Museum 87(3070): 129. Type
species: CerodriUia (Lissodriiha) schroedeti Bartsch and
Rehder, 1939, bv original designation. Turridae.

Lonoa Dall, Bartsch and Rehder, 19.38. Bemice P Bish-
op Museum Bulletin 153:178. T\pe species: Lonoa ha-
waiensis Dall, Bartsch and Rehder, 1938, by original
designation. Semelidae.

Loxoghjpta Dall, Bartsch and Rehder, 1938, as a sub-
genus oiJacteUina Iredale, 1929. Bemice P. Bishop Mu-
seum Bulletin 153:192. Type species: TeUina okhqiiih-
ncata Conrad, 1837, by original designation. Tellinidae.

Luetzenia Rehder, 1980. Smithsonian Contributions to
Zoolog}' 289:57. Type species: Luetzenia goodingi Reh-
der, 1980, by original designation. Eulimidae.

Microdochus Rehder, 1943. Proceedings of the United
States National Museum 93(3161):193. T\pe species:
Microdochus floridanus Rehder, 1943, by original des-
ignation. Elachisinidae.

Mirapecten Dall, Bartsch and Rehder, 1938. Bemice P.
Bishop Museum Bulletin 153:84. Type species: Mira-
pecten thaanumi Dall, Bartsch and Rehder, 1938, by
original designation. Pectinidae.

Monil'uipira Bartsch and Rehder, 1939. Proceedings of

the United States National Museum 87(3070): 137. Type
species: MoniUspira monihfera (Carpenter, 1857), by
original designation. Turridae.

Nanopitar Rehder, 1943, as a subgenus of Pitar Romer,
1857. Proceedings of the United States National Muse-
um 93(3161):188. Type species: Pitar (Nanopitar) pihda
Rehder, 1943, by original designation. Veneridae.

Nentsca Rehder, 1939, as a subgenus of Nenia H. & A.
Adams. 1855. Joumal of the Washington Academy of
Sciences 29(4):171. T\pe species: Nenia (Nenisca) bart-
schi Rehder, 1939, by original designation. Clausiliidae.

Ncsobomia Dall, Bartsch and Rehder, 19.38. Bernice P.
Bishop Museum Bulletin 1.53:136. T)pe species: Sole-
cardia ovata (Gould, 1850), by original designation. Kel-
hidae.

Nodochila Rehder, 1980. Smithsonian Contributions to
Zoology 289:79. Type species: Zafra pascua Hertlein,
1962, by original designation. Columbellidae.

Parancnia Rehder. 1939, as a subgenus of Nenia H. &
A. Adams, 18.55. Joumal of the Washington Academy of
Sciences 29(4): 173. Type species: Nenia perarata (von
Martens, 1873), bv original designation. Clausiliidae.

Particoma Bartsch and Rehder, 1945, as a subgenus of
Martcsia Sowerby, 1824. Smithsonian Miscellaneous
Collections 104(11):5. Type species: Martesia (Partico-
ma) cunciformis (Say, 1822), by original designation.
Pholadidae.

Paurodiscus Rehder, 19.35, as a subgenus of Pscudoma-
laxis Fischer, 1885. The Nautilus 48(4): 128. T>pe spe-
cies: Pseudornalaxis (Paurodiscus) lameUifera Rehder,
1935, by monotypy. Architectonicidae.

Pectinistcmma Rehder, 1940, as a subgenus of Propils-
bn/a Bartsch, 1906. Joumal of the Washington Academy
of Sciences .30(7):315. T\pe species: Propilsbn/a (Pectin-
istcmma) koestneri Rehder, 1940, by original designa-
tion. Urocoptidae.

Phh/cticoncha Bartsch and Rehder. 1940, as a subgenus
of Lijonsia Turton, 1822. The Nautilus 53(4): 137. Re-
placement name for Plih/ctidcnna Bartsch and Rehder,
1939, preoccupied bv Phhjctidenna D;ill, 1899. Lyonsi-
idae.

Phhjctidenna Bartsch and Rehder, 1939, as a subgenus
of Lijonsia Turton, 1822. Smithsonian Miscellaneous
Collections 9S(10):12. Tvpe species: Lijonsia (Phhjcti-
denna) htcasana Bartsch and Rehder, 1939, by original
designation. Lyonsiidae. [see above].

Phijcodrosus Rehder, 1943, as a subgenus of Rissoella
Gray-, 1847. Proceedings of the United States National
Museum 93(3161):194' T\pe species: Rissoella (Phijcod-
rosus) caribaea Rehder, 1943, by original designation.
Rissoelhdae.

Pitarenus Rehder and Abbott, 1951, as a subgenus of
Pitar Romer, 1857. Revista de la Sociedad Malacologica

M. J. Sweeney and M. G. Harasewych, 1999

Page 131

"Carlos de la Torre" 8(2):57. Type species: Pitaria cor-
clata Schwengel, 1951, by original designation. Veneri-
dae.

Policordia Dall, Bartsch and Rehder, 1938. Bemice P.
Bishop Museum Bulletin 153:217. Type species: Poli-
cordia diomedea Dall, Bartsch and Rehder, 1938, by
origin;il designation. Verticordiidae.

Privigna Dall, Bartsch and Rehder. 1938. Bemice P
Bishop Museum Bulletin 153:160. Type species: Privig-
na pikbnji Didl. Bartsch and Rehder, 1938, by original
designation. Veneridae.

Psarostola Rehder, 1943. Proceedings of the United
States National Museum 93(3161):198. Type species:
CohimbcUa tnonilifcra Sowerby, 1844, by original des-
ignation. ColumbeUidae.

Radobornia Dall, Bartsch and Rehder, 1938. Bemice P
Bishop Museum Bulletin 153:140. Type species: Rado-
bornia araia Dall, Bartsch and Rehder, 1938, by original
designation. Kelliidae.

RubcUatoma Bartsch and Rehder, 1939. Proceedings of
the United States National Museum 87(3070): 130. Type
species: RubcUatoma rubella (Kurtz and Stimpson,
1851), by original designation. Turridae.

Sigaluta Rehder, 1967. Pacific Science, 21(2):182. T\pe
species: Sigaluta pratasensis Rehder, 1967, by original
designation. Marginellidae. [junior synonym of Margi-
ncllona Martens, 1904, see Harasewych and Kantor,
1991]

Siphova.sum Rehder and Abbott, 1951, as a subgenus of
Vasuin Roding, 1798. Revista de la Sociedad Malacolo-
gica "Carlos de la Torre ' 8(2):61. Type species: Vasum
latirifonne Rehder and Abbott, 195i, by original desig-
nation. Turbinellidae.

Stcllatoma Bartsch and Rehder, 19.39. Proceedings of
the United States National Museum 87(3070): 132. Type
species: Stcllatoma stcUata (Stearns, 1872), by original
designation. Turridae.

Stenolena Dall, Bartsch and Rehder, 1938. Bemice P
Bishop Museum Bulletin 153:60. Type species: Steno-
lena hawaicn.sis Dall, Bartsch and Rehder, 1938, by orig-
inal designation. Mytilidae.

Tenia Dall, Bartsch and Rehder, 19.38. Bemice P Bishop
Museum Bulletin 153:57. Type species: Tenia pacifica
Dall, Bartsch and Rehder, 1938, by original designation.
Mytilidae.

Urotcuthis Rehder, 1945. Proceedings of the Biological
Society of Washington 58:21. Type species: Uroteuthis
bartschi Rehder, 1945, by original designation. Loligin-
idae.

Volutipysma Rehder, 1969, as subgenus of Fulgoraria
Schumacher, 1817. Venus, The Japanese Journal of Mal-
acology 27(4):130. T\pe species: Fulgoraria (Volutipijs-

mn) humerosa Rehder, 1969, by original designation.
Volutidae.

Species-level taxa

Holotyjoes are deposited in the following institutions:
ANSP, The Acadeni\' of Natural Sciences, Pliiladelphia,
Penn.sylvannia, USA; CAS, California Academy of Nat-
ural Sciences, San Francisco, Califomia, USA: DMNH,
Delaware Museum of Natural History, Wilmington, Del-
aware, USA; MCZ, Museum of Comparative Zoology at
Hanard University, Cambridge, Massachusetts, USA;
MNSH, Museo Nacional de Historia Natural, Seccion
Hidrobiologia, Santiago, Chile; NM, Natal Museum,
Pietermaritzburg, South Africa; NSMT, National Sci-
ence Museum, TokNO, Japan; SAM, South African Mu-
seum, Cape Town, South Africa; USNM, National Mu-
seum of Natural Historv, Smithsonian InstttuHon, Wash-
ington, DC, USA; and WAM, Westem Australian Mu-
seum, Perth, Australia.

abijssicola, Calliotropis (Solaricida) Rehder and Ladd,
1973. Science Reports of the Tohoku University (ser. 2,
Geology), Special volume 6:44, pi. 3, figs. 13-15. Holo-
t\pe USNM 703266, Hess Guyot, central Pacific Ocean.
Trochidae.

actinium. Ccrithidium Rehder, 1980. Smithsonian Con-
tributions to Zoologv 289:33, pi. 5, figs. 17-18. HoIot\pe
USNM 339456, off Waikild, Oaliu, Hawaii. Diastomati-
dae.

aequicostata, Pcronaeus Rehder, 1945. Revista Chilean
de Historia Natural, 48(1944):5. Replacement name for
Bulimus (Peronacus) scalarioides Philippi in Pfeiffer,
1867, preoccupied b\- Bulinus scalaroides Reeve, 1849.
Orthalicidae.

akuana, Planaxis (Hinea) Rehder, 1980. Smithsonian
Contributions to Zoolog\- 289:50, pi. 7, fig. 5. Holotype
USNM 756793, Hanga Omiti, Easter Island. Planaxidae.

alarconi, Eucliclus (Hcrpetopoma) Rehder, 1980. Smith-
sonian Contributions to Zoology 289:19, pi. 4, fig. 8. Ho-
lot\pe USNM 756195, Onetea, Hotuiti, Easter Lsland.
Trochidae.

albata, Leptarionta lenusta Rehder, 1942. Joumal of the
Washington Academ\- of Sciences 32(11):352, figs. 7-9.
Holotype USNM 5.36030, Chiriqui Province, Panama.
HelminthogKptidae.

albomaculatus, Nassarius (Telasco) Rehder, 1980. Smith-
sonian Contributions to Zoologv 289:82, pi. 10, figs. 4-
5. Holotype USNM 751615, fe Raa Raa, Easter Island.
Nassariidae.

alia, Hawaiarca Dall, Bartsch and Rehder, 1938. Bemice
P Bishop Museum Bulletin 153:28, pi. 3, figs. 7-10. Ho-
lotype USNM 173221, northeast coast of Hawaii. Arci-
dae.

alii. Cn/ptopcctcn Dall, Bartsch and Rehder, 1938.

Page 132

THE NAUTILUS, Vol. 113, No. 4

Bemice P. Bishop Museum Bulletin 153:93, pi. 23, figs.
1^, 7. Holot\-pe USNM 173194. off south coast of
Oahu, Hawaii. Pectinidae.

allisoni. Pletirotomclla Rehder and Ladd, 1973. Science
Reports of the Tohoku University (ser. 2, Geology), Spe-
ciiJ volume 6:46, pi. 3, figs. 7-8. Holot>pe USNM
703269, Agassiz Guyot, central Pacific Ocean. Turridae.

l\J'J^y.tiJ, /ItiCl^JlZ, \.JLiy\_/l, V.\„iILlCli A CIV.11X\„ V^\.»^«ll. xuiaivj

ambatensis, Naesiotus quitensis Rehder, 1940. The N
tilus 53(4):117, pi. 13, figs. 12, 14. Holot\pe US^
473973, Ambato, Prov. del Tunguragua, Ecuador Or-

53(4):117, pi. 13, figs. 12, 14

173, / ' ■ " ' ' '^-

thalicidae

amhJa, Ewilia {Spondervilia) Dall, Bartsch and Rehder,
1938. Bemice P Bishop Museum Bulletin 153:171, pi.
44, figs. 5-8. Holotype USNM 484195, Hilo, Hawaii.
M esodesmatidae .

americana, Martcsia (Diploplax) Bartsch and Rehder,
1945. Smithsonian Miscellaneous Collections 104(11):
13, pi. 2 figs. 1-2, pi. 3, figs. 3-4. Holot\iDe USNM
573550, Fort Dade. Florida. Pholadidae.

amydroghjptum. Caecum Rehder. 1980. Smithsonian
Contributions to Zoology 289:32, pi. 5, fig. 12. Holotype
USNM 757977, Onetea, Hotuiti, Easter Island. Caeci-
dae.

(lugiilata, Anisoclonta Dall, Bartsch and Rehder, 1938.
Bemice P. Bishop Museum Bulletin 153:125, pi. 34, figs.
5-6. Holot\pe USNM 427774, near Koko Head, Maun-
alua Bay, Oahu, Hawaii. SporteUidae.

antisana, Nocsiotus quitensis Rehder, 1942. The Nauti-
lus 55(3): 103. Holo^v-pe USNM 516940, Mt. Antisana,
Province of Pichincha, Ecuador. Orthalicidae.

aporema, Triphora Rehder, 1980. Smithsonian Contri-
butions to Zoology 289:43, pi. 6, fig. 13. Holot>pe ANSP
321077, Easter Island. Triphoridae.

aquilum, Cerithiopsis Rehder, 1980. Smithsonian Con-
tributions to Zoology 289:40, pi. 6, fig. 9. Holotvpe
MNSH 200388, Easter Island. Cerithiopsidae.

araia, Radohomia Dall. Bartsch and Rehder, 1938.
Bemice P. Bishop Museum Bulletin 153:140, te.\t-fig. 17,
pi. 37, figs. 5-8. Holotvpe USNM 484183, Keokea, Hilo,
Hawaii. Kelliidae.

arcella. Cirsotrema (Cirsotremopsis) Rehder, 1945. Pro-
ceedings of the Biological Societv' of Washington 58:128.
Holotvpe USNM 83725, soudiwest of Cape Hatteras,
North Carohna. Epitoniidae.

aresta, Pusioliiui Rehder, 1943. Proceedings of the Unit-
ed States National Museum 93(3161):201, pi. 20, fig. 1.
Holotvpe USNM 517056, Santa Rosa, Pinar del Rio,
Cuba. Costellariidae.

argoblijsis. Bulla (Leucophijsema) Rehder and Ladd,
1973. Science Reports of the Tohoku Universits' (ser. 2,
Geology), Special volume 6:47, pi. 3, figs. 5-6. Holotype
USNM' 703272, Horizon Guyot, Mid-Pacific Mountains,
central Pacific Ocean. Bullidae.

atlantica. Cooperella Rehder, 1943. Proceedings of the
United States National Museum 93(3161):187, pi. 19.
figs. 3-4. Holot)pe USNM 517058, off Peanut Island,
northem Lake Worth, Florida. Cooperellidae.

auaua, Lima Dall, Bartsch and Rehder, 1938. Bemice P.
Bishop Museum Bulletin 153:106, pi. 27, figs. 5-8. Ho-
lotype USNM 335607, Auau Channel, Hawaii. Liniidae.

aureofasciatus, Comts spuiius Rehder and Abbott, 1951.
Revista de la Sociedad Malacologica "Carlos de la Torre"
8(2):64, pi. 9, figs. 3-4. Holotype USNM 597521, off
Dry Tortugas, Florida. Conidae.

austini, Couus Rehder and Abbott, 1951. Journal of the
Washington Academv of Sciences 41(1):22, fig. 7. Ho-
lotype USNM 603017. southeast of Loggerhead Key,
Drv^ Tortugas, Florida. Conidae.

aviarius, Musculu.s Dall, Bartsch and Rehder, 1938.
Bemice P. Bishop Museum Bulletin 153:53, pi. 10, figs.
1-4. Holot>pe USNM 173231, near Mokii Manu, Ha-
w;ui. M\tilidae.

bahamensis, Maiicsia (Dioloplax) Bartsch and Rehder,
1945. Smithsonian Miscellaneous Collections 104(11):
11, pi. 3, figs. 15-16. Holotype USNM 573549, eastern
end of South Bight, Andros Island, Baliamas. Pholadi-
dae.

bahamondci, Hcmiliosfraca Rehder, 1980. Smithsonian
Contributions to Zoology 289:55, pi. 7, fig. 13. Holotvpe
USNM 766572, Easter Island. Euhmidae.

bahamondei, Zcidora Rehder, 1980. Smithsonian Con-
tributions to Zoology 289:17, pi. 4, figs. 4—5. Holot\pe
MNSH 200403, Vaihu, Easter Island. FissureHidae. '

bailci/i, Nitidella guttata Bartsch and Rehder, 1939.
Smithsonian Miscellaneous Collections 98(10):6. pi. 2,
fig, 6. Holotype USNM 472857, Elizabeth Bay, Alber-
marle Island, Galapagos Islands. ColunibeUidae.

baked, Botulina Dall, Bartsch and Rehder, 1938. Bem-
ice P Bishop Museum Bulletin 153:55, pi. 11, figs. 7-8.
Holot\pe USNM 484182 [incorrectly published as
484183]. Hilo, Hawaii. Mytilidae.

baldwini, Solccuiius Dall, Bartsch and Rehder, 1938.
Bemice P. Bishop Museum Bulletin 153:176, pi. 45. figs.
9-10. Holotype USNM 335618, off soudi coast of Mo-
lokai, Hawaii. Psammobiidae.

balesi, Asthenothaenis Rehder, 1943. Proceedings of the
United States National Museum 93(3161):189, pi. 19,
figs. 1.3-14. Holotype USNM 536052, Missouri Key,
Florida. Thracidae.

barken. Tegulo (Chlnrostoma) Bartsch and Rehder,
1939. Smithsonian Miscellaneous Collections 98(10):11,
pi. 2, figs. 10-12. Holotvpe USNM 472589, Elizabeth
Bay, Albemarle Island, Galapagos Islands. Trochidae.

baniardi, Fusivoluta Rehder, 1969. The Veliger 11(3):
207, pi. 40, fig. 9, pi. 43, figs. 40-43. Holotype DNMH
10751, off Nat;il coast. Volutidae.

M. J. Svveene)- and M. G. HarasevvAch. 1999

Page 1.33

bartschi, Nenia (Nenisca) Rehder, 1939. Journal of the
Washington Acadeni\' of Sciences 29(4): 171, figs. 1-5.
Holot^pe USNM 380074, Mt. Rochelle (or Rochelois),
south of Rochelois, Haiti. Clausiliidae.

bartschi, Ooconjs Rehder, 1943. Proceedings of the
United States National Museum 93(3161): 197, pi. 20,
fig. 16. Holot>pe USNM 535689, south of Tortugas,
Florida. Cassidae.

bai-fschi, Turbonilla (Carcliopsis) Aguayo and Redher,
1936. Memorias de la Sociedad Cuhana de Historia Nat-
ural 9(4):267, pi. 24, fig. 7. Holot\pe USNM 420975, La
Chorrera, Habana, Cuba. PvTamidellidae.

bartschi. Urotcuthis Rehder, 1945. Proceedings of the
Biological Socieh- of Washington 58:22, pi. 3, figs. 1-2.
Holotype USNM 573515, Jolo Harbor, Jolo, Philippine
Islands. Loliginidae.

baijeri, Natica (Ghjphcpithema) Rehder, 1986. The Nau-
tilus 100(1):38. Replacement name for Ghjphcpithema
foridiiiui Rehder, 1943, preoccupied by Natica (Cnjp-
tonatica) floridana Dall, 1892. Naticidae. [see teclbaijeii,
Natica Rehder, 1986]

bermudezi, Rissoina (Fohnia) Aguayo and Redher, 1936.
Memorias de la Sociedad Cubana de Historia Natural
9(4):265, text-fig. 1, pi. 24, fig. 5. HolotApe USNM
436307, Tortugas, Florida. Rissoidae.

blakci, VitrineUa Rehder, 1944. The Nautilus 57(3):97,
pi. 9, figs. 1-2. Holot>pe USNM 537834, Talbot for-
mation (Pleistocene) at Wailes Bluff (near Cornfield
Harbor), St. Marys Countv, Maryland. Vitrinellidae.

boswellae, Volutocorbis Rehder, 1969. The Veliger 11(3):
202, pi. 40, fig. 5, pi. 41, figs. 16-19. Holot\'pe USNM
683585, off Mosselbaai (MosselBay), South Africa. Vol-
utidae.

boitntyi. Fusinus galatheae Rehder and Wilson, 1975.
Smithsonian Contributions to Zoology 203:10, te.xt-figs.
6-8. HoloUpe USNM 707230, off NW coast of Pitcaim
Island. Fasciolariidae.

bourgeoisae, Dnjmaeus Rehder, 1943. The Nautilus
57(1):28, pi. 6, fig. 10. Holot>pe USNM 517550, near
Paraje Nuevo (near Cordoba), Vera Cruz, Mexico. Or-
thalicidae.

branhamae, Fasciolaria distans Rehder and Abbott,
1951. Revista de la Sociedad Malacologica "Carlos de la
Torre" 8(2):59, pi. 8, figs. 4-5. Holotype USNM 597513,
off Puerto Alvaro Obregon, Tabasco, Mexico. Fasciolar-
iidae.

hntnnescens, Fenimorea inoscri Rehder, 1943. Proceed-
ings of the United States National Museum 93(3161):
202, pi. 20, fig. 5. Holotvpe USNM 517055, off Fort
Walton, Okaloosa County', Florida. Turridae.

bucki. Kitna Dall, Bartsch and Rehder, 1938. Bemice P
Bishop Museum Bulletin 153:149, text-fig. 25, pi. 39,

figs. 13-16. Holotype USNM 333067, near Koko Head,
Oaliu, Hawaii. Montacutidae.

candcns, Actcon Rehder, 1939. The Nautilus 53(1 ):21,
pi. 6, fig. 7. Holotyi^e USNM 493407, off Fowey Light,
Florida. Acteonidae.

caribaca, RissocUa (Phi/codrosus) Rehder, 1943. Pro-
ceedings of the United States National Museum
93(316i):194, pi. 20, fig. 7. Holotype USNM 536046,
Bonefish Key, Florida. Rissoellidae.

caiibbaea, Tritoualia (Ociucbiina) Bartsch and Rehder,
1939. Smithsonian Miscellaneous Collections 98(10):7,
pi. 1, fig. 1. Hok)t>pe USNM 472617, Old Province Is-
land, Colombia [Caribbean], Muricidae.

canieopicta, Phenacovolva Rehder and Wilson, 1975.
Smidisonian Contributions to Zoology 203:6, frontis-
piece figs. 6, 9. 12. Holot>pe USNM 707232, Haava
Strait, Marquesas Islands. Ovulidae.

caijosensc. Caecum (Caecum) Rehder, 1943. Proceedings
of the United States National Museum 93(3161):190, pi.
20, fig. 9. Holot>pe USNM 536045, Bonefish Key Flor-
ida Keys. Caecidae.

cedrosensis, Aph/sia Bartsch and Rehder, 1939. Smith-
sonian Miscellaneous Collections 9S(10):2, pi. 4, figs. 8-
10, pi. 5. Holotxpe USNM 472859, east side of Cedros
Island, Lower California, Me.xico. Aplysiidae.

ccrnohorski/i, Ziba Rehder and Wilson, 1975. Smithson-
ian Contributions to ZoologN 203:13, te.xt-fig. 9, frontis-
piece fig. 13. Holotxpe USNM 707239, off NW comer
of Pitcaim Island. Mitridae.

chilcna, CaUiostoma Rehder, 1971. Proceedings of the
Biological Society of Washington 83(51):590, figs. 2, 5.
Holotxpe USNM 701669, NW of Valparaiso, Chile. Tro-
chidae.

clappi, CcrodiiUia (CcrodriUia) Bartsch and Rehder,
1939. Proceedings of the United States National Muse-
um 87(3070): 128, pi. 17, fig. 4. Holot>pe USNM 493408,
Hawk Channel, Florida. Turridae.

clarkci, Fusivohita Rehder. 1969. The Veliger 11(3):206,
pi. 40, fig. 8, pi. 43, figs. 37-39. Holotxpe DNMH 12833,
off Joao Belo, District Gaza, Mozambique. Volutidae.

clarki, Conus Rehder and Abbott, 1951. Journal of the
Washington Academy of Sciences 41(1 ):22, figs. 1-6.
Holohpe USNM 485740, southwest of Marsh Island,
Iberia Countv, Louisiana. Conidae.

ch'uchi. Pohjgyra Rehder, 1932. The Nautilus 45(4): 129,
pi. 10, figs.' 1-3. Holotyi^e MCZ 81347, Izard County
Arkansas. Polygyridae.

clippertom'Hsis, Ctena Bartsch and Rehder, 1939. Smith-
sonian Miscellaneous Collections 98(10): 13, pi. 3, figs.
1-5. Holotxpe USNM 472552, Clipperton Island, East
Pacific Ocean. Lucinidae.

codoceoae, Hehacus Rehder, 1980. Smithsonian Contri-

Page 134

THE NAUTILUS, Vol. 113, No. 4

butions to Zoology 289:32, pi. 5, figs. 13-15. Holotype
ANSP 321078, Easter Island. Architectonicidae.

cou.sohrinclla, Zafroiio Rehder, 19S0. Smithsonian Con-
tributions to Zoolog\- 289:76, pi. 9, figs. 12-13. Holot)'pe
USNM 756156, Haka Ea, Vaihu, Easter Island. Col-
umbellidae.

cookei, Chlami/s Dall, Bartsch and Rehder, 1938. Bern-
ice R Bishop Museum Bulletin 153:90, pi. 24, figs. 1^.
Halot\pe USNM 484166. off entrance to Honolulu Har-
bor, Oaliu, Hawaii. Pectinidae.

cookei, Dorisca Dall, Bartsch and Rehder, 1938. Bemice
P Bishop Museum Bulletin 153:160, pi. 42, figs. 5-8.
Holotype USNM 484190, Pearl Harbor, Oahu, Hawaii.
Veneridae.

costaricana, Si/strophia (Sijstrophiella) Rehder, 1942.
Journal of the Washington Academy of Sciences 32(11):
352, figs. 13-15. Holot>pe USNM 536023, Goto, on the
Golfo Dulce, Puntarenas Province, Costa Rica. Systro-
phiidae.

costariceusis, TJii/sanophora Rehder, 1942. Journal of
the Washington Academy of Sciences 32(11):352, figs.
1-3. Holot>pe USNM 536009, La Caja, near San Jose,
San Jose Province, Costa Rica. Thysanophoridae.

crassiila. Kcrmia Rehder, 1980. Smithsonian Contribu-
tions to Zoology 289:89, pi. 11, fig. 2. Holotype MNSH
200397, VaihuT Easter Island. Turridae.

crassula, Merelina (Mereliniopsis) Rehder, 1980. Smith-
sonian Contributions to Zoology 289:30, pi. 5, fig. 9. Ho-
lot\'pe USNM 767026, Onetea, Hotuiti, Easter Island.
Rissoidae.

cruda, Arcinella Dall, Bartsch and Rehder, 1938. Bem-
ice P Bishop Museum Bulletin 153:118, pi. 33, figs. 9-
10. Holotype USNM 335601, off south coast of Molokai,
Hawaii. Chamidae.

cn/pfa. Tenia Dall, Bartsch and Rehder, 1938. Bemice
P Bishop Museum Bulletin 153:58, pi. 11, figs. .5-6. Ho-
lotype USNM 333333, off Oalui, Hawaii. Mytilidae.

cubanum, Vexillum Aguayo and Redher, 1936. Memorias
de la Sociedad Cubana de Historia Natural 9(4):267, pi.
24, fig. 4. Holotype USNM 420978, t)pe locality not
published. [Holotype from La Chorrera, Habana, Cuba].
Costellariidae.

dalli, Cirsotrema (Cirsofremopsis) Rehder, 1945. Pro-
ceedings of the Biological Society of Washington 58:128.
Holot>pe USNM 515240, off Cape San Bias, Florida,
Gulf of Mexico. Epitoniidae.

dichroiis, Semelangulus Dall, Bartsch and Rehder, 1938.
Bemice P. Bishop Museum Bulletin 153:180, pi. 46, figs.
1^. Holot>pe USNM 362356, Midway Island. Tellini-
dae.

diodorus, Semelangulus Dall, Bartsch and Rehder, 1938.
Bemice P. Bishop Museum Bulletin 153:181, pi. 46, figs.

9-12. Holotype USNM 484201, off Launiupoko, Maui,
Hawaii. Tellinidae.

diomcdca, Gh/cymcris Dall, Bartsch and Rehder, 1938.
Bemice P. Bishop Museum Bulletin 153:38, pi. 6, figs.
3-4. Holot>pe USNM 172987, off south coast of Mo-
lokai, Hawaii. Glycymerididae.

diomcdca, Policordia Dall, Bartsch and Rehder, 1938.
Bemice P. Bishop Museum Bulletin 153:217, pi. 56, figs.
5-8. Holotype USNM 335685, Pailolo Channel, Haw^i.
Verticordiidae.

diomcdca, RuhcUatoma Bartsch and Rehder, 1939. Pro-
ceedings of the United States National Museum
87(3070):131, pi. 17, fig. 3. Holotype USNM 508645,
Sanibel Island, Florida. Turridae.

diomedcum, Propeamussium Dall, Bartsch and Rehder,
1938. Bemice P Bishop Museum BulleHn 153:83, pi. 20,
figs. 3-4, pi. 21. figs. 1-4. Holot>pe USNM 190442, Pail-
olo Channel, Hawaii. Propeamussiidae.

diomcdcus. Pcctcn Dall, Bartsch and Rehder. 1938.
Bemice P. Bishop Museum Bulletin 153:96, pi. 24, figs.
7-8. Holot>pe USNM 335570 [incorrectly published as
355570], off south coast of Molokai, Hawaii. Pectinidae.

dispai; Cuspidaria (Mijoncra) Dall, Bart.sch and Rehder,
1938. Bemice P Bishop Museum Bulletin 153:225, pi.
58, figs. 5-7. Holotype USNM 110752, off south coast
of Molokai, Hawaii. Cuspidariidae.

disparilis, Volutocorbis Rehder, 1969. The Veliger 11(3):
203, pi. 40, fig. 4, pi. 41, figs. 20-22. Holot>pe SAM
A3335, 125 miles off Saint Sebastian Bay '(36°40'S,
2r26'E), South Africa. Volutidae.

carh/i, Engiua Bartsch and Rehder, 1939. Smithsonian
Miscellaneous Collections 98(10):6, pi. 1, fig. 8. Holo-
type USNM 472566, Sulivan Bay, James Island, Gala-
pagos Islands. Buccinidae.

edmondsoni, Periglypta Dall, Bartsch and Rehder, 1938.
Bemice P. Bishop Museum Bulletin 153:162, pi. 43, figs.
1-4. Holot>pe USNM 484193, Hilo, Hawaii. Veneridae.

enajcrti, Rissoina (Rissolina) turricula Rehder, 1980.
Smithsonian Contributions to Zoolog) 289:26, pi. 5, fig.
6. Holot)pe USNM 755993, Hanga'Papara, Easter Is-
land. Rissoidae.

epiphanea, Pusia Rehder, 1943. Proceedings of the Unit-
ed States National Museum 93(3161 ):201,' pi. 20, fig. 14.
Holot>pe USNM 414278. off Tortugas, Florida. Costel-
lariidae.

espcranzac, Stomatclla Rehder. 1980. Smithsonian Con-
tributions to Zoolog)' 289:20, pi. 4, figs. 9-11. Holotype
USNM 756113, Haka Ea, Vaihu, Easter Island. Stoma-
tellidae.

cuchaiis, Triphora Rehiler, 1980. Smithsonian Contri-
butions to Zoologv^ 289:46, pi. 6, fig. 16. Holotype ANSP
339946, Easter Island. Triphoridae.

M. J. Sweeney and M. G. Harasewx'ch, 1999

Page 135

cufirantiiuitiis. Conns Bartsch and Rehder, 1943. Pro-
ceedings of the Biological Socieh' of Washington 56:85
Holotvpe USNM 173213, off north coast of Molokai Is-
land (near Mokapu Islet), Hawaii. Conidae.

exilirata, Licnardia Rehder, 1980. Smithsonian Contri-
butions to Zoolog\' 289:86, pi. 11, figs. 7-9. Holotype
USNM 758393, Haka Ea, Vaihu, Easter Island. Turridae.

exomilisca, Triphora Rehder, 1980. Smithsonian Contri-
butions to Zooiogv' 289:45, pi. 6, fig. 15. Holotyi:)e
USNM 756779, Onetea, Hotuiti, Easter Island. Triphor-
idae.

exquisita, Martcsia (Dioloplax) Bartsch and Rehder,
1945. Smithsonian Miscellaneous Collections 104(11):
10, pi. 3, figs. 17-18. Holotype USNM 573548, Stony
Cove, St. Mar\'s Parish, Jamaica. Pholadidae.

exquisita. Pinna Dall, Bartsch and Rehder, 1938. Bemice
R Bishop Museum Bulletin 153:75, pi. 17, figs. 1-2. Ho-
lotype USNM 335610, near Laysan Island, Hawaii. Pin-
nidae.

fasciola, Lithophaga Dall, Bartsch and Rehder, 1938.
Bemice P. Bishop Museum Bulletin 153:57, pi. 11, figs.
1-4. Holotype USNM 337458, off Launiupoko Camp,
Maui, Hawaii. Mvtilidae.

filosa, Pyrgoajthara Rehder, 1943. Proceedings of the
United States National Museum 93(3161):203, pi. 20,
fig. 2. Holotype USNM 27637, Marco, Florida. Arcidae.

fimbriata, Hipponix Bartsch and Rehder, 1939. Smith-
sonian Miscellaneous Collections 98(10): 10, pi. 2, figs.
7-9. Holotype USNM 472853, Clipperton Island, East
Pacific Ocean. Hipponicidae.

floridana, Anachis Rehder, 1939. The Nautilus 53(1):20,
pi. 6, fig. 6. Holotype USNM 473202, near Cape Ca-
naveral, Brevard County, Florida. Columbellidae.

floridana, Ghjphepithenm Rehder, 1943. Proceedings of
the United States NaHonal Museum 93(3161): 196, pi.

19, figs. 19-21. HoIot\pe USNM 517060, Peanut Island,
Lake Worth, Florida. Naticidae. [see bayeri, Natica Reh-
der, 1986]

floridana, Tiicla Rehder, 1939. The Nautilus 53(1):18,
pi. 6, figs. 2-3. HoIot\pe USNM 473118, Jensen Beach
(near Miami), Florida. Veneridae.

floridanus, Microdochiis Rehder, 1943. Proceedings ot
the United States National Mu.seum 93(3161):193, pi.

20, fig. 6. HoIot\pe USNM 536048, Bonefish Key Flor-
ida. Tvpe localit)' corrected (The Nautilus 57(1 j:32) to
Missouri Key. Elachisinidae.

frctcnsis. Gemma Rehder, 1939. The Nautilus 53(1):18.
pi. 6, figs. 8-9. Holotyi^e USNM 508650, Long Island
Sound, New York. Veneridae.

flinisicola, Martesia (Diploplax) Bartsch and Rehder,
1945. Smithsonian Miscellaneous Collections 104(11):
14, pi. 3, figs. 1-2, 13-14. Holotype USNM 573551,
Lake Worth, Florida. Pholadidae.

fuscobasis, Clathurella Rehder, 1980. Smithsonian Con-
tributions to Zoolog\- 289:87, text-fig. 11, pi. 11, figs. 3-
4. Holotjpe USNM' 756265, Onetea, Hotuiti, Easter Is-
land. Turridae.

gayi, Penmaeus Rehder, 1945. Revista Chilean de His-
toria Natural, 48(1944):4. Holotype USNM 5.37831, An-
tofagasta, Chile. Orthalicidae.

gloriosiis, Spondyliis Dall, Bartsch and Rehder, 1938.
Bemice P. Bishop Museum Bulletin 153:102, pi. 26, figs.
8-11. HoIot)pe USNM 1904.35, off Penguin Bank, south
coast of Oahu, Hawaii. Spondyhdae.

goodingi, Luetzenia Rehder. 1980. Smithsonian Contri-
butions to ZooIog^■ 289:58, pi. 7, fig. 16. Svntspes (79)
USNM, ZMC, AMS, MNSH, Easter Island. Eulimidae.

goodwini. Haiya Rehder, 1993. The Nautilus 106(4):
127, figs. 1-2. Holot>pe USNM 860312, French Frigate
Shoals, Hawaii. Haipidae.

gouhli, Halirardia Dall, Bartsch and Rehder, 1938.
Bemice P. Bishop Museum Bulletin 153:218, pi. 56, figs.
1-4. Holot>pe USNM 335682, off Hawaii. Verticordi-
idae.

graeilior, Benthovaluta Rehder, 1967. Pacific Science,
21(2):1S5, figs. 5-6. HoIot\pe USNM 6.37252. off Ca-
gayan Islands, northern Sulu Sea, PhiUppines. Turbinel-
hdae.

gregon/i, Anguipecfcn Dall, Bartsch and Rehder, 1938.
Bemice P. Bishop Museum Bulletin 153:93, pi. 23, figs.
5-6, 8. Holotype USNM 173227, near Kau;ii, Hawaii.
Pectinidae.

grcgon/i. Teredo (Tercdora) Dall, Bartsch and Rehder,
1938. Bemice P Bishop Museum Bulletin 153:212, pi.
55, figs. 1-5. Holotvpe USNM 337316, Keaukiilia. Hiio,
Hawaii. Teredinidae. [junior .sviionym of Tercdora prin-
cesae (Sivickis, 1928), seeTumer, 1966:103]

halitropiis, Conns Bartsch and Rehder, 1943. Proceed-
ings of the Biological Socieh' of Washington 56:88. Ho-
lotype USNM .338579, at dredger dump at Hololulu,
Oahu, Hawaii. Conidae.

hammafus, Contts Bartsch and Rehder, 1943. Proceed-
ings of the Biological Societ}' of Washington 56:86. Ho-
lotype USNM 173225, near Kauai Island, Hawaii. Con-
idae.

Iiataii. Calliotropis {Solaricida) Rehder and Ladd, 1973.
Science Reports of the Tohoku University (ser. 2, Ge-
olog)'). Special volume 6:43, pi. 3, figs. 16-18. HoIot>pe
USNM 703263, Hess Guyot, central Pacific Ocean, tro-
chidae.

hausfrelhim. Succinca Rehder, 1942. Joumal of the
Washington Academy of Sciences 32(11):350, fig. 19.
Holotype USNM 536013, Pedemal, Guanacaste Prov-
ince, Costa Rica. Succineidae.

Iiauaia, Barhatia (Abarbatia) Dall, Bartsch and Rehder,

Paee 136

THE NAUTILUS, Vol. 113, No. 4

1938. Bemice P. Bishop Museum BulleHn 153:32, pi. 4,
figs. 1-4. Holotype USNM 484154 [incorrectly pub-
lished as 485154], Keaukaha, Hilo, Hawaii. Arcidae.

hawaia, Lithophaga Dall, Bartsch and Rehder, 1938.
Bemice P. Bishop Museum Bulletin 153:56, pi. 11, figs.
9-10. Holotype USNM 337459, entrance of Hololulu
Harbor, Oaliu, Hawaii. Mvtilidae.

hawaiana, Lima Dall, Bartsch and Rehder, 1938. Bem-
ice P Bishop Museum Bulletin 153:105, pi. 28, figs. .5-
6. Holotype USNM 173202, off south coast of Molokai,
Hawaii. Limidae.

hawaiana, Meiocardia Dall, Bartsch and Rehder, 1938.
Bemice P. Bishop Museum Bulletin 153:121, pi. 34, figs.
17-18. Holotype USNM 173048, near Kauai, Hawii.
Cardiidae.

hawaicnsis, Acar Dall, Bartsch and Rehder, 1938. Bem-
ice P. Bishop Museum Bulletin 153:14, text-fig. 3, pi. 1,
figs. 5-8. Holot>pe USNM 427766. Keaukaha, Hilo, Ha-
waii. Arcidae.

hawaicnsis, Anguhis Dall, Bartsch and Rehder, 1938.
Bemice P. Bishop Museum Bulletin 153:198, pi. 50, figs.
9-12. Holotype USNM 484200, entrance to Honolulu
Harbor, Oahu, Hawaii. Tellinidae.

hawaicnsis, Arcinella Dall, Bartsch and Rehder, 1938.
Bemice P. Bishop Museum Bulletin 153:118, pi. 33, figs.
11-14. HoIot>pe USNM 3.35602, off northeast coast of
Hawaii. Chamidae.

hawaicnsis. Bcntharca Dall, Bartsch and Rehder, 1938.
Bemice P. Bishop Museum Bulletin 153:20, pi. 3 figs.
15-18. Holot>pe USNM 427759, off south coast "^of
Oahu, Hawaii. Arcidae.

hawaiensis, Boftila Dall. Bartsch and Rehder, 1938.
Bemice P. Bishop Museum Bulletin 153:59, pi. 12, figs.
1-4. Holot)pe USNM 484180, off south coast of Mo-
lokai, Hawaii. Mxtilidae.

hawaiensis. CorditcUa Dall, Bartsch and Rehder, 1938.
Bemice P. Bishop Museum Bulletin 153:120, pi. 33. figs.
5-8. Holot>pe USNM 484175, off Waikiki, Oaliu, Ha-
waii. Carditidae.

hawaicnsis, Chlanu/s Dall, Bartsch and Rehder, 1938.
Bemice P. Bishop Museum Bulletin 153:88, pi. 22, figs.
5-8. Holotype USNM 484165, Honolulu side of en-
trance to channel to Pearl Harbor, Oahu, Hawaii. Pec-
tinidae.

hawaiensis, Cuspidaria Dall. Bartsch and Rehder, 1938.
Bemice P. Bishop Museum Bulletin 153:226, pi. 58, figs.
1-4. Holot>pe USNM 173012, Pailolo Channel, Haw^i.
Cuspidariidae.

hawaiensis, Krllia Dall. Bartsch and Rehder. 1938. Bem-
ice P. Bishop Museum Bulletin 153:143, text-fig. 20, pi.
38, figs. 1.5-18. Holot^pe USNM 361563, Fort Arm-
strong, Oahu, Hawaii. Kelliidae.

hawaiensis, Lasaea Diill, Bartsch and Rehder. 1938.

Bemice P. Bishop Museum Bulletin 1.53:141. text-fig. 18,
pi. 38, figs. 1-4. Holot>pe USNM 484184, Keaau, Ha-
waii. Lasaeidae.

hawaiensis, Lonoa Dall, Bartsch and Rehder, 1938.
Bemice P. Bishop Museum Bulletin 153:179, pi. 46, figs.
.5-8. Holot>pe USNM 337471. Keokea. Hilo. Hawaii.
Semehdae.

hawaiensis, Loripiniis Dall, Bartsch and Rehder, 1938.
Bemice P. Bishop Museum Bulletin 1.53:127, pi. 34, figs.
11-12. Holot>pe USNM 337404. entrance to Honolulu
Harbor. Oahu. Hawaii. Lucinidae.

hawaicnsis, Maiicsia Dall. Bartsch and Rehder. 1938.
Bemice P. Bishop Museum Bulletin 153:205. pi. 52. figs.
1-7. Holotype USNM 484213, Pearl Harbor, Oahu, Ha-
waii. Pholadidae.

hawaicnsis. Pinna Dall. Bartsch and Rehder. 1938.
Bemice P. Bishop Museum Bulletin 1.53:73. pi. 17, figs.
8-11. HolotApe USNM 337508, off Kaanapali, Maui,
Hawaii. Pinnidae.

hawaiensis, Rocellaria Dall. Bartsch and Rehder. 1938.
Bemice P Bishop Museum Bulletin 153:202, pi. 51, figs.
1-5. Holotype USNM 484206, Ford Island, Oahu, Ha-
waii. Gastrochaenidae.

hawaiensis, Saxicava Dall, Bartsch and Rehder, 1938.
Bemice P Bishop Museum BulleHn 153:200. pi. .50. figs.
1.3-14. Holotype USNM 484204, Fort Armstrong, Oaliu,
Hawaii. Hiatellidae.

hawaicnsis. Spondi/his Dall, Bartsch and Rehder, 1938.
Bemice P. Bishop Museum Bulletin 153:100, pi. 25, figs.
1^. Holotxpe USNM 337515, off Honolii reef, Hilo,
Hawaii. Spondyfidae.

hawaicnsis. Stcnolcna Dall, Bartsch and Rehder, 1938.
Bemice P. Bishop Museum Bulletin 153:60, text-fig. 14,
pi. 12, figs. 6-7. Holot>pe USNM 110427. off south
coast of Molokai, Hawaii. Mvtilidae.

hawaiensis, Teredo (Teredops) Dall, Bartsch and Rehder,
19.38. Bemice P Bishop Museum Bulletin 153:213, pi.
55, figs. 6-8. Holot\pe USNM 3.35077. off south coast
of Oahu. Hawaii. Teredinidae. [junior s\Tionvm of Li/-
rodiis pcdiccUatus (Quatrefages. 1849), see Turner,
1966:104]

hawaiensis, Trachi/cardinni Dall, Bartsch and Rehder,
19.38. Bemice P Bishop Museum Bulletin 1.53:1.55, pi.
41, figs. 1-4. Holotype USNM 337390. Keaukaha. Hilo.
Hawaii. Cardiidae.

hawaiensis, Venus Dall. Bartsch and Rehder. 19.38.
Bemice P. Bishop Museum Bulletin 153:164. pi. 42. figs.
1.3-15. Hok)t\pe USNM 3.35581 [incorrectly pubfished
as 3.3.5881], near Kauai, Hawaii. Veneridae.

hcniphilli. Pi/rgoct/tliara Bartsch and Rehder. 19.39. Pro-
ceedings of the United States National Museum
87(3076):132. pi. 17. fig. 2. Holotype USNM 86898a,
Sarasota Bav, Florida. Turridae.

M. J. Sweeney and M. G. Harasewych, 1999

Page 137

luiulcr.soiii. Bdrhdtui UAhtirhafia) Dall, Bartsch and Reh-
der, 1938. Beniice P. Bishop Museum Bulletin 153:33,
text-fig. 12. pi. 5, figs. 5-6. Holot>pe USNM 427768,
outcrop in the road that leads from the main route into
the Pearl Harbor grounds, 0;Uiu, Hawaii. Arcidae.

hendersoni, Chcima Dall, Bartsch and Rehder, 1938.
Bemice F Bishop Museum Bulletin 153:150. pi. 40, figs.
.5-8. Holot>pe USNM 484174, Pearl Harbor, Oahu, Ha-
waii. Chamidae.

hendersoni, Cymatoica oricntalis Rehder, 1939. The
Nautilus 53(1):19, pi. 6, figs. 10-11. Holot^pe USNM
493384, Fowey Light, near Miami, Florida. TeUinidae.

hendersoni, Piisia Rehder, 1943. Proceedings of die
United States National Museum 93(3161):200, pi. 20.
fig. 12. Holotype USNM 414359, off Bears Cut. Miami,
Florida. Costellariidae.

hiloa, Barbatia (Abarbatia) Dall, Bartsch and Rehder,
1938. Bemice P Bishop Museum Bulletin 153:34, pi. 4,
figs. 13-16. Holotype USNM 427769, Keaukalia, Hilo,
Hawaii. Arcidae.

hihm. Scintilla Dall, Bartsch and Rehder, 1938. Bemice
P Bishop Museum Bulletin 153:146, text-fig. 22, pi. 38,
figs. 11-14. Holotype USNM 484187, Keokea, Hilo, Ha-
waii. Galeommatidae.

houstonius, Brachidontcs midtifonnis Bartsch and Reh-
der, 1939. Smithsonian Miscellaneous Collections
98(10): 14. pi. 4, figs. 4^1. Holotype USNM 472858, Su-
livan Ba\', James Island, Galapagos Islands. M\-tilidae.

hua, Calloarca (Barbarca) Dall, Bartsch and Rehder,
1938. Bemice P Bishop Museum Bulletin 153:24, te.xt-
fig. 8, pi. 4, figs. 5-8. Holotype USNM 427760, entrance
to Pearl Harbor, Oaliu, Hawaii. Arcidae.

humboldti, Bathybembix Rehder, 1971. Proceedings of

the Biological Societv of Washington 83(51):587, fig. 4.
Holot\pe'uSNM 70i665, NW of Valparaiso, Chile. Tro-
chidae.

huincrosa. Fulffirarin (Vohitipt/snia) Rehder, 1969. Ve-
nus, The Japanese Journal of Malacology 27(4): 130, pi.
7, figs. .5-7. Holotype USNM 237732, SSW of Tung-sha-
Tao (Pratas Island), South China Sea. Volutidae.

jacksoni, Naesiotus quitcnsis Rehder, 1940. The Nautilus
53(4):116, pi. 13, figs. 1, 5. Holotype USNM 47.3969,
Guaillabamba (northeast of Quito), Ecuador Orthalici-
dae.

jaumci, Diadora [sic] Aguayo and Rehder, 19.36. Me-
morias de la Sociedad Cubana de Historia Natural 9(4):
263, pi. 24, fig. 6. Holotype USNM 420974, Varadero,
Matanzas, Cuba. Fissurellidae.

jtiddi, Haumea Dall, Bartsch and Rehder, 1938. Bemice
P Bishop Museum Bulletin 153:86, pi. 22, figs. 1-4. Ho-
lotyi^e USNM 337.538, off Kaanapali, Maui, Hawaii. Pec-
dnidae.

kajiijainai. Haqui Rehder. 1973. Indo-Pacific Mollusca

3(16):244, pi. 188. figs. .3^. Holotype NSMT 414.50,
southern Philippines. Haqiidae. [junior s\non\in and ju-
nior honionxni ol Harjxi kujiijiinuii Habe in Habe anil
Kosuge, 1972, see Petit & Bieler, 1996:44]

kanaka. Roccllaria Dall, Bartsch and Rehder, 19.38.
Bemice P. Bishop Museum Bulletin 153:203, pi. 51, figs.
11-15. Holotype USNM .337379, Mokuoloe Island, Ka-
neohe Bay, Oahu, Hawaii. Gastrochaenidae.

katiaia. Clticijtncris Dall, Bartsch and Rehder, 1938.
Bemice P. Bishop Museum Bulletin 153:40, pi. 7. fig.s.
2-3. Holotype USNM 173043. near Kauai, Hawaii. Gly-
CAinerididae.

katiaia, Navicula Dall. Bartsch and Rehder, 19.38. Bem-
ice P Bi.shop Museum Bulletin 1.53:11, pi. 1, figs. 9-12.
Holotype USNM .3.35696, off Kauai, Hawaii. Arcidae.

katiaia, Ostrca Dall, Bartsch and Rehder, 19.38. Bemice
P Bishop Museum Bulletin 153:112, pi. 30, figs. 8-9.
Holotype USNM 3.35585, near Kauai, Hawaii. Ostrei-
dae.

kaiiaicnsis, Bentharca Dall, Bartsch and Rehder, 1938.
Bemice P. Bishop Museum Bulletin 1.53:22, te.xt-fig. 7,
pi. 3, figs. .5-6. Holotype USNM 173232, near Kauai,
Hawaii. Arcidae.

katiaicnsis, Clilaint/s Dall, Bartsch and Rehder, 1938.
Bemice P. Bishop Museum Bulletin 153:92, pi. 22, figs.
9-10. Holotype USNM 3.3.5674, near Kauai, Hawaii.
Pectinidae.

katiaicnsis, Spondt/Iiis Dall, Bartsch and Rehder, 1938.
Bemice P. Bishop Museum Bulletin 153:103. pi. 26. figs.
12-13. Holotype USNM .3356.38, near Kauai, Hawaii.
Spondylidae.

kauaiensis, Teredo {Teredops) Dall, Bartsch antl Rehder,
1938. Bemice P Bishop Museum Bulletin 153:214.
These authors affixed the name T. kauaiensis to Millers
(1924) description of what he believed to be Teredo die-
gensis Bartsch, 1916, from Pearl Harbor, Honolulu Har-
bor, and Nawiliwili. They comment that thes ha\e not
had specimens of this species for examination. There can
be no t}pe material for Teredo kauaiensis, as neither
specimens nor figs, were associated with the name by
either Miller or Dall, Bartsch and Rehder Teredinidae.
[s)Tion\m oi Lt/rodus pedicellatus Quatrefages, 1849, see
Turner, 1966:106]

kauaium, Propcainu.ssium Dall, Bartsch and Rehder,
1938. Bemice P Bishop Museum Bulletin 1.53:80, pi. 19,
figs. 4-7, pi. 20 figs. 1-2. Holotype USNM 17.3218, near
Kauai, Hawaii. Propeamussiidae.

kcohca, Lima Dall, Bartsch and Rehder, 1938. Bemice
P Bishop Mu.seum Bulletin 1.53:107. pi. 27, figs. 1.3-16.
Holotype USNM 337519, Kaneohe Bay, Oahu, Hawaii.
Limidae.

kilburni, Volutocorbis Rehtler. 1974. The Nautilus 88(2):
.36, figs. .5-8. Holotype USNM 709.3.52, off Durban, Na-
tal. X'olutidae.

Page 1.38

THE NAUTILUS. Vol. 113, No. 4

kingae, Ci/pmca Rehder and Wilson. 197.5. Smithsonian
Contributions to Zoology* 203:2. text-figs. 1-3, frontis-
piece figs. 1, 4-.5. 7-8. Holotype USNM 7072.34, off NW
point of Pitcaim Island. Cypraeidae.

kona, Ghjcymeris Dall, Bartsch and Rehder, 1938. Bern-
ice R Bishop Museum Bulletin 1.53:41, pi. 7, figs. 8-9.
Holot\pe USNM 427765. off south coast of Oahu, Ha-
waii. Glvcymerididae.

kocitncii. Propilsbn/a {Pcctitrntcmma) Rehder. 1940.
Journal of the Washington Academ\' of Sciences .30(7):
316, figs. 1-3. Holotype USNM 535762, Cerro Potosi at
Galeana, Nuevo Leon, Mexico. Urocoptidae.

kupua, Ostrea Dall. Bartsch and Rehder, 1938. Bemice
R Bishop Museum Bulletin 1.53:111, pi. .30, figs. 1-4.
Holot>pe USNM 484156, Pearl Harbor, Oahu, Hawaii.
Ostreidae.

laciniata. Ptcria Dall, Bartsch and Rehder, 1938. Bemice
R Bishop Museum Bulletin 1.53:70. pi. 17, figs. 5-7. Ho-
lotype USNM 3.3.5666, off northeast coast of Hawmi.
Pteriidae.

lahaina. Lima Dall, Bartsch and Rehder, 1938. Bemice
P Bishop Mu.seum Bulletin 1.53:105, pi. 27. figs. 1-4.
Holotype USNM 484169, off Laliaina, Maui, Hawaii.
Limidae.

lamcUifcra, P.seudomalaxis {Puurodisctis) Rehder 1935.
The Nautilus 48(4): 128, pi. 7, figs. 8-10. Holotype
USNM 4262.35. Florida Straits. Architectonicidae.

langfordi, Nodipecten Dall, Bartsch and Rehder, 1938.
Bemice P. Bishop Museum Bulletin 153:85, pi. 21, figs.
9-12. Holotype USNM 337.541, off Waildld. Oaliu, Ha-
waii. Pectinidae.

latilirata. CoraUiophila Rehder, 1985. The Nautilus
99(4):97, figs. 1-3. Holotype USNM 731531, Oeno, Pit-
caim Islands. Coralliophilidae.

latiriforme, Vasitm (Siphovasuryi) Rehder and Abbott,
1951. Revista de la Sociedad Malacologica "Carlos de la
Torre" 8(2):61, pi. 9, figs. .5-6. Holotype USNM .597517.
NW of Cape Catoche, Yucatan, Mexico. Turbinellidae.

Jaijsana, Acar Dall, Bartsch and Rehder, 1938. Bemice
P. Bishop Museum Bulletin 1.53:16. text-fig. 4. pi. 1, figs.
13-16. Holotype USNM .335647, off Laysan Island, Ha-
waii. Arcidae.

laijsana, ArcineUa Dall, Bartsch and Rehder, 1938. Bem-
ice P Bishop Museum Bulletin 1.53:118, pi. .33. figs. IS-
IS. Holotype USNM 484178, Laysan Island, Hawaii.
Chamidae.

lai/sana, Botula Dall, Bartsch and Rehder, 1938. Bemice
P' Bishop Museum Bulletin 153:60. pi. 12, fig. 5. Holo-
type USNM 335614, near Laysan Island. Hawaii. Mvtil-
idae.

latjsana, Ostrea Dall, Bartsch and Rehder, 1938. Bemice
P Bishop Mu.seum Bulletin 1.53:111. pi. .32, figs. .5-8.

Holotype USNM 484157, off Laysan Island, Hawaii. Os-
treidae.

laijsanus, Muscidus Dall, Bartsch and Rehder, 1938.
Bemice P. Bishop Museum Bulletin 153:54, pi. 10, figs.
11-12. Holotype USNM 190444, vicinity of Laysan Is-
land, Hawaii. Mytilidae.

k'ptocharactum. Ccrithhim (Tlicricium) Rehder, 1980.
Smithsonian Contributions to Zoology 289:38, pi. 6, figs.
.5-7. Holotype USNM 7.563.35, Hanga Piko. Easter Is-
land. Cerithiidae.

leucathcina. Triphora Rehder, 1980. Smidisonian Con-
tributions to Zoology 289:42, pi. 6, fig. 12. Holotype
USNM 7.56003, Hanga Papara, Easter Island. Triphori-
dae.

leviuscula. Fiilgoniiia (Fulgoraria) Rehder, 1969. Venus,
The Japanese Journal of Malacolog\' 27(4): 128, pi. 7, figs.
1-4. Holotype USNM 231770. NW of Tung-sha-Tao
(Pratas Island). South China Sea. Volutidae.

Umiianfi, Inijoris Rehder. 1980. Smithsonian Contribu-
tions to Zoology 289:41. pi. 6. fig. 10. Holotype USNM
7.56215, Onetea, Hoituiti, Easter Island. Triphoridae.

lircllata, Nerita (Hcminerita) Rehder, 1980. Smithsonian
Contributions to Zoology 289:23, pi. 4, figs. 14-15. Ho-
lotype USNM 7.56795, Hanga Omiti. Easter Island. Ner-
itidae.

livida, Sm/ella Rehder 19.35. The Nautilus 48(4):129, pi.

7, fig. 7. Holotype USNM 12.5.556, Corpus Christi Bay,
Texas. Pyramidellidae.

loisac, Acncator (Ellicea) Rehder. 1971. Proceedings of
the Biological Societ\^ of Washington 83(51):.593, figs. 7-

8. Holotype USNM 701667, type locality not indicated
[NW of Valparaiso, Chile]. BuccinuHdae.

loisac, Austroliarpa Rehder, 1973. Indo-Pacific Mollusca
3(16):264, pi. 237. figs. 3, 6. Holot^pe WAM 1,56-72,
NW of Rottnest Island, Western Australia. Harpidae.

loisac. Euplica Rehder, 1980. Smithsonian Contributions
to Zoology 289:77, pi. 10, figs. 6-8. HoIot^pe USNM
7.53940. Haka Ea, Vaihu, Easter Island. Columbellidae.

loisac. Triphora Rehder 1980. Smithsonian Contribu-
tions to Zoology 289:44, pi. 6, fig. 14. Holot^pe USNM
756778, Hanga Papara, Easter Island. Triphoridae.

hmgincjua, Merelina (Merelina) Rehder, 1980. Smithson-
ian Contributions to Zoology 289:29, pi. 5. fig. 10. Ho-
lotype USNM 766829, Onetea, Hotuiti, Ea.ster Island.
Rissoidae.

hicasana. Lt/onsia (Phh/ctidcnna) Bartsch and Rehder,
1939. Smithsonian Miscellaneous C'ollections 98(10): 12,
pi. 4, figs. 1-3. Holotype USNM 472526, off Punta Gor-
da. Cape San Lucas. Lower California. Mexico. Lvonsi-
idae.

lutea, Anisodonta Dall. Bartsch and Rehder, 19.38. Bem-
ice P Bishop Museum BuIIeHn 1,53:124. pi. 34, figs. 7-

M. J. Sweeney and M. G. Harasewych, 1999

Page 1.39

10. Holotype USNM 337468, a Honolulu beach, Oahu,
Hawaii. Sportellidae.

mafflalcncnsis. Coiuis Bartsch and Rehder, 1939. Sniidi-
sonian Miscellaneous Collections 98(10):4, pi. 1, figs. 5,
9. Holotype USNM 472521, Magdalena Bay, Lower Cal-
ifornia, Mexico. Conidae.

mariclac. Conns Rehder and Wilson, 1975. Smithsonian
Contributions to Zoolog\- 203:14, text-fig. 10, frontis-
piece figs. 10-11. Holot\pe USNM 70.3255, off Baie
Motu-Hee, Nuku Hiva, Marquesas Islands. Conidae.

nmuia. Cadella Dall, Bartsch and Rehder, 1938. Bemice
P. Bishop Museum Bulletin 1.53:197, pi. 50, figs. 1-4.
Holotxpe USNM .337.347, off Launiupoko, Maui, Ha-
waii. Tellinidae.

inaitid. Gh/ci/meris Dall, Baitsch and Rehder, 1938.
Bemice P. Bishop Museum Bulletin 153:38, pi. 6, figs.
.5-6. Holot)pe USNM .337443, off Mala Bay, Maui, Ha-
waii. Glycymerididae.

mania, Navicula Dall, Bartsch and Rehder, 1938. Bem-
ice P. Bishop Museum Bulletin 1.53:12, text-fig. 2, pi. 1,
figs. 1-4. Holotype USNM 190446, Aleunihana Channel,
Hawaii. Arcidae.

uiaunahiana. CaUoarca (Barharca) Dall, Bartsch and
Rehder, 1938. Bemice P. Biship Museum Bulletin 153:
26, pi. 4, figs. 9-12. Holotype USNM 427757, near Koko
Head, Maunalua Bay, Oahu, Haw;iii. Arcidae.

maijaonim, Dnjmaeus Rehder, 1966. Proceedings of the
Biological Societ>' of Washington 79:287, figs. .3^. Ho-
lotype USNM 251656, Isla Mujeres, Quintana Roo,
Mexico. Orthahcidae.

iiicdiofxicificensis, DentaUitm Rehder and Ladd, 1973.
Science Reports of the Tohoku University (ser. 2, Ge-
ology), Special volume 6:48, pi. 3, figs. 1-2. Holoty|3e
USNM 70.3273, Agassiz Guyot, Mid-Pacific Mountains,
central Pacific Ocean. Dentaliidae.

mcsolciica, Crassispiro (Crassispirclla) Rehder, 1943.
Proceedings of the United States National Museum
93(3161 ):202, pi. 20, fig. 15. Holotype USNM 411906,
Looe Key Reef, Florida. Turridae.

mighehi, Margarites Rehder, 1937. Proceedings of the
Biological Society of Washington .50:115. Replacement
name for Margarites johnsoni Dall, 1921, preoccupied
hv Margarita johnsoni Arnold, 1909. Trochidae.

miUeri, Teredo (Cormiteredt)) Dall, Bartscli and Rehder,
19.38. Bemice P Bi.shop Museum Bulletin 1.53:210. pi.
54, figs. 1-2. New name for Teredo affinis Deshayes,
1863, as described and figured by Miller, 1924. Dall,
Bartsch and Rehder comment that they have not seen
this species, and applv their new name to Miller's de-
scription, which they believed to be tlifferent from Te-
redo affinis Deshaves, 1863. The holot)pe of Teredo niil-
leri (CAS 12384) is the specimen designated as the neo-
type of Teredo affinis Deshayes, 1863 bv Moll, 1941

[Turner, 1966:111). Teredo milleri Dall, Bartsch and
Rehder, 1938, is therefore a junior objective ,s\Tion\in of
Teredo affinis Deshayes, 1863, now placed in the genus
Li/rodus [Turner, 1966:86]. Teredinidae.

niiltoplax, Chiton Rehder, 1932. The Nautilus 45(4):pl.
10, figs. 4—5. Nomen nudum.

minuda, Dimi/a Dall, Bartsch and Rehder, 1938. Bernice
P Bishop Museum Bulletin 153:78, pi. 19, figs. 1-2. Ho-
lotvpe USNM 173002, off south coast of Molokai, Ha-
waii. Dimvidae.

niimus, Sp(nidt/Jus Dall, Bartsch and Rehder, 1938.
Bemice P. Bishop Museum Bulletin 153:102, pi. 26. figs.
6-7. Holotvpe USNM 337509, entrance to Honolulu
Harbor, Oaliu, Hawaii. SpondyUdae.

molokaia, Barbatia {Aharbatia) Dall, Bartsch and Reh-
der, 1938. Bemice P Bishop Museum Bulletin 1.53:.32,
te.xt-fig. 11, pi. 5, figs. 7-10. Holot)pe USNM 17.3203,
off south coast of Molokai, Hawaii. Arcidae.

molokaia, Diuii)a Dall. Bartsch and Rehder, 19.38. Bern-
ice P Bishop Museum Bulletin 1.53:79, pi. 19, fig. 3.
Holotype USNM 190431, off north coast of Molokai,
Hawaii. Dimvidae.

molokaia, Gh/ct/meris Dall. Bartsch and Rehder, 1938.
Bemice P. Bishop Museum Bulletin 153:.35, pi. 6, figs.
1-2. HolotApe USNM 427761, off south coast of Mo-
lokai, Hawaii. GKcvmerididae.

molokaiinn, Propeannissiiim D;ill. Bartsch and Rehder,
1938. Bemice P Bishop Museum Bulletin 153:82, pi. 20,
fig. 8, pi. 21, figs. .5-6. Holot>pe USNM 335677, off
south coast of Molokai, Hawaii. Propeamussiidae.

inolokains, Muscidus Dall, Bartsch and Rehder, 1938.
Bemice P. Bishop Museum Bulletin 153:53, pi. 10, figs.
.5-6. Holot>pe USNM 190445, off south coast of Mo-
lokai, Hawaii. Mvtilidae.

mondis. Moiulispira Bartsch and Rehder, 1939. Pro-
ceedings of the United States National Museum
87(3070): 137, pi. 17, fig. 10. Holotvpe USNM 508649,
Waveland, Dade County, Florida. Turridae.

morrisoni, Microdaphne Rehder, 1980. Smithsonian
Contributions to Zoology 289:88, text-fig. 12, pi. 11, figs.
,5-6. HoIot^pe USNM 758.390, Motu Mataira, Raroia,
Tuamotu Islands. Turridae.

mozamhieana, Voiutocorlns Rehder, 1972. The Veliger
15(1):12, figs. .3-5, 7-9. Holotype USNM 717909,^95
miles SE of Inhaca Id., Mozambique. Volutidae.

nniriei, Anabathron Bartsch and Rehder, 1939. The
Nautilus 52(4):110, pi. 8, figs. 2, 2a. Holotype USNM
5.35345, Oghuga Island, Aleutian Islands. Barleeidae.

nana, Voiutocorlns Rehder, 1974. The Nautilus 88(2):35,
figs. 1-4. Holotvpe USNM 709.351, off coast of southern
Zululand, Natal. Volutidae.

ncbiilo.

Faritihim Rehder, 1943. Proceedings of the

Page 140

THE NAUTILUS. Vol. 11.3, No. 4

United States National Museum 93(3161):190, pi. 20,
fig. S. Holot>pe USNM 536042, Bonefish Key, Florida.
Type locality' corrected (The Nautilu.s 57(1 ):32) to Mis-
souri Key, 14 or 15 miles west of Bonefish Key. Caeci-
dae.

nchnlosns. Semclonguhi.s Dall, Bartsch and Rehder,
1938. Bemice P. Bishop Museum Bulletin 153:180, pi.
47, figs. 5-8. Holot>pe USNM 484202, off Launiupoko,
Maui, Hawaii. Tellinidae.

nesiotum, Propeamussiiiiii Dall, Bartsch and Rehder
1938. Bemice P Bishop Museum Bulletin 153:81, pi. 19,
figs. 12-15, pi. 20 fig. 7. Holot^pe USNM 190441, off
south coast of Molokai, Hawaii. Propeaniussiidae.

ncwcombi. Vohella {AimjgdaJum) Dall, Bartsch and Reh-
der, 1938. Bemice P. Bishop Museum Bulletin 153:45,
pi. 8, figs. 9-10. Holot)pe USNM 173022, off south
coast of Oaliu, Hawaii. Mytilidae.

nigrescens, Alvania Bartsch and Rehder 1939. Smith-
sonian Miscellaneous Collections 98(10):8, pi. 2, fig. 5.
Holot>pe USNM 472621, Old Providence Island,"Co-
lombia [Caribbean]. Rissoidae.

nigrita, St/napfccochica Rehder, 1939. The Nautilus
53(1):20, pi. 6, fig. 1. Holot>pe USNM 473205, Logger-
head Key, Dry Tortugas Keys, Florida. Stomatellidae.

notabilis, Adrana Rehder 1939. The Nautilus 53(1): 16,
pi. 6, figs. 4, 12. Holot>pe USNM 499337, Paraguana
Peninsula, Venezuela. Nuculanidae.

micella. Pinguitellina Dall, Bartsch and Rehder, 1938.
Bemice P Bishop Museum Bulletin 153:195, text-fig. 28,
pi. 49. figs. 9-12. Holot>pe USNM 484199, off Lihau,
Maui, Hawaii. Telfinidae.

niittingi, CaUoorca (Barharca) Dall, Bartsch and Rehder,
1938. Bemice P Bishop Museum Bulletin 153:26, pi. 4,
figs. 17-20. Holotype USNM 335648, Laysan Island,
Hawaii. Arcidae.

nux, Gh/ct/mcri.s Dall, Bartsch and Rehder, 1938. Bem-
ice P. Bishop Museum Bulletin 153:36, pi. 6, figs. 9-12.
Holot>pe USNM 173003, Pailolo Channel, Hawaii. Gly-
c)'merididae.

oahua, Barbatia (Abarbatia) Dall, Bartsch and Rehder,
1938. Bemice P Bishop Museum Bulletin 153:30, te.xt-
fig. 10, pi. 5, figs. 1-4. Holotype USNM 427767, Ka-
neohe Bay, Oahu, Hawaii. Arcidae.

oahua, Ghjcynwris Dall, Bartsch and Rehder, 1938.
Bemice P. Bishop Museum Bulletin 153:39, pi. 7, figs.
10-11. Holotype USNM 338229, off soudi coast of
Oahu, Hawaii. Glycymerididae.

oahua. Pinna Dall, Bartsch and Rehder 1938. Bemice
P Bishop Museum Bulletin 153:75, pi. 18, figs. .5-6. Ho-
lotype USNM 337.506, t)pe localit)' not stated. Pinnidae.

oahuana, Cadella Dall, Bartsch and Rehder, 1938. Bem-
ice P Bishop Museum Bulletin 153:197, pi. .50, figs. .5-

8. Holot>pe USNM 33.3069, near Koko Head, Oaliu,
Hawaii. Tellinidae.

oahuana, RoccUaha Dall, Bartsch and Rehder 1938.
Bemice P. Bishop Museum Bulletin 153:201, pi. 51, figs.
6-10. Holotype USNM 337314, off Waikiki, Oahu, Ha-
waii. Gastrochaenidae.

oahuensis. Lt/onsia (AUogramma) Dall, Bartsch and
Rehder 1938. Bemice P. Bishop Museum Bulletin 153:
215, pi. .56, figs. 9-12. Holotvpe USNM 17.3017, Pailolo
Channel, Hawaii. Ljonsiidae.

oahuensis, Senielanguhis Dall, Bartsch and Rehder,
1938. Bemice P Bishop Museum Bulletin 153:181, pi.
47, figs. 9-12. Holotvpe USNM 337346. entrance to
Honolulu Harbor Oaliu, Hawaii. Tellinidae.

oahuus, Muscuhis Dall, Bartsch and Rehder, 1938. Bem-
ice P Bishop Museum Bulletin 1.53:54, pi. 10, figs. 1.3-
14. Holotype USNM 484181, Pearl Harbor, Oaliu, Ha-
waii. M\tilidae.

ochrostigmata. Natica Rehder 1980. Smithsonian Con-
tributions to Zoolog\- 289:64, pi. 8, figs. 13-15. Holotvpe
USNM 339159, off Launiupoko, Maui, Hawaii. Natici-
dae.

oiinus, Naesiotus quitensis Rehder 1940. The Nautilus
.53(4):116, pi. 13, figs. 6, 10. Holot>pe USNM 47.3971,
Chimborazo (near Riobamba), Ecuador Ortliahcidae.

oialis, Hitia Dall. Bartsch and Rehder 1938. Bemice R
Bishop Museum Bulletin 1.53:136, text-fig. 15, pi. 37,
figs. 1-4. Holot>pe USNM 21120, "SandWich Islands"
[Hawaii]. Sportellidae.

pacifica. Tenia Dall, Bartsch and Rehder 1938. Bemice
P Bishop Museum Bulletin 1.53:58, pi. 11, figs. 11-14.
Holot>pe USNM 17.3020, off south coast of Oahu, Ha-
waii. Mvtillidae.

paiU^loana, Cuspidaria (Mijoncra) Dall, Bartsch and
Rehder, 1938. Bemice P Bishop Museum Bulletin 1.53:
225, pi. .58, figs. S-11. Holotvpe USNM 17.3015, Pailolo
Channel, Hawaii. Cuspidariidae.

pailohmm. Propeamussium Dall, Bartsch and Rehder
1938. Bemice P Bishop Museum Bulletin 153:82, pi. 19,
figs. 8-11, pi. 20, figs. .5-6. Holotype USNM 17.3209,
Pailolo Channel, Hawaii. Propeamussiidae.

pailolous. Musculus Dall, Bartsch and Rehder 1938.
Bemice P. Bishop Museum Bulletin 153:.55, pi. 10, figs.
7-10. Holotype USNM 3.35310 [incorrectly pubUshed as
3.3.5301], Napih Harbor Maui, Hawaii. Mytihdae.

palmeri, Humboldtiana Clench and Rehder 1930. The
Nautilus 44(1): 12, pi. 2, figs. 1-4. Holotvpe MCZ 79779,
Jeff Davis Counh', Texas. Helminthoglyptidae.

parallcia, Lima Dall. Bartsch and Rehder 1938. Bemice
P Bishop Museum Bulletin 153:106, pi. 27, figs. 9-12.
Holotype USNM 33.3074, off Lahaina, Maui, Hawaii.
Limidae.

M. J. Sweeney and M. G. Harasewych, 1999

Page 141

parvispimis, Spondtjhis Dall, Bartsch and Rehder, 1938.
Bemice P. Bishop Mtiseum Bulletin 15.3:99, pi. 25, figs.
5-(S. Holotvpe USNM 484161, Honaunau, Hawaii.
Spond\lidae.

pascua, Siphonaria Rehder, 1980. Smithsonian Contri-
butions to Zoologx- 289:97, pi. 12, figs. 4-7. Holotvpe
CAS 58706, Easter Island. Siphonariidae.

pascuana. Gramda Rehder, 1980. Smithsonian Contri-
butions to ZooIogN- 289:85, pi. 9, fig. 20. Holot>pe
USNM 710721, dnetea, Hotuiti, Easter Island. Cystis-
cidae.

pascucnsis, Conus iniliaiis Rehder, 1980. Smithsonian
Contributions to Zoolog\- 289:91, pi. 9, figs. 21-22. Ho-
lotvpe USNM 766496, Hanga Piko, Easter Island. Con-
idae.

pauhsciilpta. Spiraxis (Rccfaxis) Rehder, 1942. Journal
of the Washington Academy of Sciences 32(11):350, fig.
18. Holotvpe "uSNM 536016, Santa Maria, San Jos'e
Province, Costa Rica. Spiraxidae.

perductonim, Dnjinaeits Rehder, 1943. The Nautilus
57(1):29, pi. 6, figs. 6-9. Holotype USNM 517552, near
Las Gnitas de Cacahuamilpa, Guerrero, Me>dco. Or-
thalicidae.

pcrcgrina. Pohigifra Rehder, 1932. The Nautilus 45(4):
130, te.xt-fig. l'. Holot)pe MCZ 81349, Izard Count)- Ar-
kansas. Polyg\Tidae.

pernjae, Cerodrillia (Cc rod till ia) Bartsch antl Relider,
1939. Proceedings of the United States National Muse-
um 87(.3070):128, pi. 17, fig. 1. Holot%pe USNM 508644,
Sanibel Island, Florida. Turridae.

penyac. Kuiiziclla Bartsch and Rehder. 1939. Proceed-
ings of the United States National Museum 87(.3()7()):
134, pi. 17, figs. 7, 9. Holotype USNM 508646, Sanibel
Island, Florida. Turridae.

philippii, Peronaeus Rehder, 1945. Revista Chilean de
Historia Natural, 48(1944):3. Holot>pe USNM .5378.30,
near Copiapo, Chile. Orthalicidae.

pilshnji, Dcntalhim (Antalis) Rehder, 1942. The Nauti-
lus 56(2):69. Replacement name kn Dentalutm (Aittalis)
pwudohcxagonum "Ihering" Henderson, 1920, preoc-
cupied by Dentaliiim pseudohexagoniiin Arnold, 1903.
Dentaliidae.

pihbnji, Megasplra Rehder, 1945. The Nautilus 59(2):
67. New name for Mcga.spira data (Gould) Pilsbry,
1904, not Mcga.spira data Gould, 1847. Three s\Titypes
are enumerated, ANSP 3057, ANSP 25041, USNM,
number not pubHshed. Brazil. Megaspiridae.

pilshrtji, Ptivigna Dall, Bartsch and Rehder, 1938. Bem-
ice P ' Bishop Mu.seum Bulletin 153:161, pi. 42, figs. 9-
12. Holot\pe USNM 484192, Pearl Harbor, Oahu, Ha-
waii. Veneridae.

ington Academy of Sciences .32(11):352, figs. 10-12. Ho-
lotvpe USNM 536018, Santa Maria, San Jose Province,
Costa Rica. C'haropidae.

pilula, Pitar (Nanopitar) Rehder, 1943. Proceedings of
the United States National Mu.seum 93(3161): 188, pi.
19, figs. 5-10. Holotype USNM 517057, Lake Worth,
Florida. Veneridae.

pi.siiiiia, Melanella Rehder, 1980. Smithsonian Contri-
butions to Zoologv 289:54, pi. 7, figs. 10-11. Holotype
USNM 756117, Ilaka Ea, Vaihu, Easter Island. Eul'im-
idae.

pisiim, Batlu/arca Dall, Bartsch and Rehder, 1938. Bem-
ice P. Bishop Mu.seum Bulletin 153:19, text-fig. 6, pi. 3,
figs. 1-4. Holotype USNM 335684, Pailolo Channel, Ha-
waii. Arcidae.

planospira, SpiratvJIa Rehder, 1942. United States De-
partment of the Interior, Geological Survey, Professional
Paper 196-D:108, pi. 23, figs. 5-7. Holotvpe USNM
535416, 49°36'N 28°54'W, middle of North Atlantic
Ocean (sample H-63, core 8). Liniacinidae.

polyuesica. Metaxia Rehder, 1980. Smithsonian Contri-
butions to Zoology 289:47, pi. 7, fig. 1. Holotvpe MNSH
200413, Easter Island. Triphoridae.

polynesica, Nucida Rehder, 1980. Smithsonian Contri-
butions to Zoologv' 289:106, pi. 13, figs. 1-2. Holohpe
USNM 756244, bnetea, Hotuiti, Easter Island. Nucu-
lidae.

polipicsica, Williamia Rehder, 1980. Smithsonian Con-
tributions to Zoology 289:98, pi. 12, figs. 10-11. Holo-
hpe USNM 757897,' off Waikiki, Oahu," Hawaii. Siphon-
ariidae.

powclli, Comitas Rehder and Ladd, 1973. Science Re-
ports of the Tohokii University (sen 2, Geology), Special
volume 6:45, pi. 3, figs. 11-12^ Holotvpe USNM 703267.
Agassiz Guvot, central Pacific Ocean. Turridae.

praccipua. Monda Rehder, 1980. Smithsonian Contri-
butions to Zoology 289:70, pi. 9. fig. 5. Holotvpe USNM
751592, Hanga Piko, Easter Island. Muridicae.

pratasemis. Sigaluta Rehder, 1967. Pacific Science,
21(2):182, figs. 1-4. Holohpe USNM 2.37618 [incor-
rectly published as 237018], west of Pratas Reef, South
China Sea (20°37'N, 115°43'E). Marginellidae. [junior
synonym oi' MargiiicIIona gigas (Martens, 1904) see Har-
asewych and Kantor, 1991]

pidchcn Hal(ipsci)liiis Rehder, 1943. Proceedings of the
United States National Museum 93(3161):I9f, pi. 20,
figs. 3, 10. Holotvpe USNM 500638, off Lazaretto, Bar-
bados. Turbinidae.

piimilio, Peronaeus Rehder, 1945. Revista Chilean de
Historia Natural, 48(1944):5. Replacement name for Bu-
limus uantis Reeve, 1849, preoccupied by Buliiuiis nan-
us Lamarck, 1804. Orthalicidae.

/;;/.s7;n/(. fif7fr7f/(.sr(/.s Rehder. 1942. Journal of the Wash- rapauiiiensis. Lene<>noj)sis Rehder, 1980. Smithsonian

Page 142

THE NAUTILUS, Vol. 113, No. 4

Contributions to Zoolog>' 289:101, pi. 12, fig. 3. Holo-
t>pe USNM 756790, Onetea, Hotuiti, Easter'island. El-
lobiidae.

recta, Atiina (Atriiw) Dall. Bartsch and Rehder, 193S.
Bemice P. Bishop Museum Bulletin 153:76, pi. 17, figs.
3^. HolotApe USNM 484153, off south coast of Mo-
lokai, Hawaii. Pinnidae.

rectangula, Hawaiarca Dall, Bart:sch and Rehder, 1938.
Bemice P. Bishop Museum Bulletin 153:27, text-fig. 9,
pi. 3, figs. 11-14. Holot>pe USNM 173222, Alenuihaha
Channel, Hawaii. Arcidae.

riisei, Arene Rehder, 1943. Proceedings of the United
States National Museum 93(3161):192, pi. 19, figs. 17-
18. Holot\pe USNM 42858, St. John, Virgin Islands.
Turbinidae.

rooscvciti, Coiuis Bartsch and Rehder, 1939. Smithson-
ian Miscellaneous Collections 98(10):3, pi. 1, figs. 4, 7.
Holot>pe USNM 472854, Clipperton Island, East Pacific
Ocean. Conidae.

rooscvciti, Marginella Bartsch and Rehder, 1939. Smith-
sonian Miscellaneous Collections 98(10):5, pi. 1, figs. 2-
3. Holot>pe USNM 472610, Old Providence Island, Co-
lombia [Caribbean]. Marginellidae.

rooscvciti, Pcasiella Bartsch and Rehder, 1939. Smith-
sonian Miscellaneous Collections 98(10):8, pi. 2, figs. 1-
3. Holot\pe USNM 472575, Sulivan Bav, James Island,
Galapagos Islands. Littorinidae.

rooscvciti, Tcrebra (Subula) Bartsch and Rehder, 1939.
Smithsonian Miscellaneous Collections 9S(10):1, pi. 1,
fig. 6. Holot\pe USNM 472534, Socorro Island, Mexico.
Terebridae.

rosavittoriae. Volutocorhis Rehder, 1981. The Nautilus
95(4):169, figs. 1-5. Holot>pe USNM 784653, Off Kisi-
mayu, southern Somalia. Volutidae.

rosea, Kancoha Dall, Bartsch and Rehder, 1938. Bemice
F Bishop Museum Bulletin 153:142, te.xt-fig. 19, pi. 39,
figs. 9-12. Holot>pe USNM 484185, Mokuoloe Island,
Kaneohe Bay, Oahu, Hawaii. Kelliidae.

rostrata, Poromya Rehder, 1943. Proceedings of the
United States National Museum 93(3161):189, pi. 19.
figs. 11-12. Holotype USNM 536152. off Delray Beach,
Palm Beach Countv, Florida. Poromvidae.

rostratula. Ervilia Rehder, 1943. Proceedings of the
United States National Museum 93(3161 ):189, pi. 19,
figs. 1-2. Holot>pe USNM 517059, Lake Worth, Florida.
Mesodesmatidae.

ruhropicta, Charrui Bartsch and Rehder. 1939. Smith-
sonian Miscellaneous Collections 98(10): 13, pi. 3, figs.
6-10. HoIot)pe USNM 472553, Clippertf)n Island, East
Pacific Ocean. Chamidae.

rudK, Septifcr Dall, Bartsch and Rehder, 1938. Bemice
P. Bi.shop Museum Bulletin 153:50, pi. 9, figs. .5-8. Ho-

lotvpe USNM 173223, off Alenuihaha Channel, Hawaii.
Mvtilidae.

niizona, Limop.sis (Felicia) Rehder, 1971. Proceedings of
die Biological Societv of Washington 83(51):586, fig. 1.
Holot>pe USNM 701671, NW of Valparaiso, Chile. Um-
opsidae.

sagcnaiia, Kcnnia Rehder, 1980. Smithsonian Contri-
butions to Zoolog\' 289:90, pi. 11, fig. 1. Holohpe
USNM 756115, Haka Ea, Vaihu, Easter Island. Turridae.

sagittatus. Modiolus {Aimjgdahtm) Rehder, 1935. The
Nautilus 48(4):127, pi. 7, figs. 11-12. Holotvpe USNM
93999, off Cape San Bias, Florida. Mytilidae.

sallasi, Murcx (Murcx) recuniirostris Rehder and Ab-
bott, 1951. Revista de la Sociedad Malacologica "Carlos
de la Torre" 8(2):58, pi. 9, figs, 7-8. Holot\pe USNM
597515, NW of Cape Catoche, Quintana Roo, Mexico.
Muricidae.

sanihelcnsis, Crassispira (Crassispirclla) Bartsch and
Rehder, 1939. Proceedings of the United States National
Museum 87(3070):135, "pi. 17, figs. 11-12. Holot\pe
USNM 508647, Sanibel Island, Florida. Turridae.

sanihelcnsis, Succinca Rehder, 1933. The Nautilus 47(1):
20. Holot\pe MCZ 59645, Sanibel Island, Florida. Suc-
cineidae.

scaphoides, Adrana Rehder, 1939. The Nautilus 53(1):
17, pi. 6, fig. 5. Holot>pe USNM .364223, near Carta-
gena, Colombia. Nuculanidae.

schcjferi. Liocijma Bartsch and Rehder, 1939. The Nau-
tilus 52(4):111, pi. S, figs. 1-lb. Holot>pe USNM
535344, off Chuginadak Island, Aleutian Islands, Alaska.
Veneridae.

schmitfi. Litfoiina Bartsch and Rehder, 1939. Smithson-
ian Mi.scellaneous Collections 98(10):9, pi. 2, fig. 4. Ho-
lot\pe USNM 472547, Clipperton Island, East Pacific
Ocean. Littorinidae.

schrocderi, Ccrodiillia (Lisodrillia) Bartsch and Rehder,
1939. Proceedings of the United States National Muse-
um 87(.3070):13d, pi. 17, fig. 8. Holotype USNM 530585,
off Charlotte Harbor, Florida. Turridae.

seminigata, Voliitocorbis Rehder and Weaver, 1974. The
Nautilus 88(2):31, figs. 1-8. Holohpe NM 99.39, SE of
Bluff, Durban, South Africa. Tvpe catalog number and
localitv- corrected [The Nautilus 89(3):79]" to NM Moll,
no. G769, off Ilha Bazaruto, Mozambique. Volutidae.

sennottoniui, Contis Rehder and Abbott, 1951. Revista
de la Sociedad Malacologica "Carlos de la Torre" 8(2):
63, pi. 9, figs. 1-2. Hoiot>pe USNM 597519, SW of
Campeche, Yucatan, Mexico. Conidae.

scrratissimus, Spondijlus D;ill, Bartsch and Rehder,
1938. Bemice P Bishop Mu.seum Bulletin 153:97, pi. 25.
figs. 9-12. Holot>pe USNM 337513, off Honolulu,
Oahu, Hawaii. Spondyhdae.

M. J. Sweeney and M. G. Harasewvch, 1999

Page 143

significans, Barischia Relider, 1943. Proceedings of the
United States National Museum 93(3161): 199, pi. 20,
fig. 17. Holot)pe USNM 516493, off Tortugas, Florida.
Buccinidae.

smima, Conus Bartsch and Relider, 1943. Proceedings
of the Biological Society' of Washington 56:87. Holotype
USNM 173226, near Kauai Island, Hawaii. Conidae.

sparsipunctata, Psarostola inonilifera Rehder, 1943. Pro-
ceedings of the United States National Museum
93(316i):198, pi. 20, fig. 11. Holot)pe USNM 450778,
southeast of Fowe\' Light, Florida. Columbellidae.

sparsispiuosiis, Sponchiltis Dall, Bartsch and Rehder,
1938. Bemice P Bishop Museum Bulletin 153:98, pi. 26,
figs. 1-5. Holotype USNM 190436, Pailolo Channel, Ha-
waii. Spondylidae.

spiceri, Comptopallium Rehder, 1944. The Nautilus
58(2):52, pi. 2, figs. 1-2. Holotype USNM 518010,
Christmas Island, Line Islands, Central Pacific. Pectini-
dae.

spiceri, Conus Bartsch and Rehder, 1943. Proceedings
of the Biological Societv of Washington 56:87. Holotvpe
USNM 537792, Sand Island, Midway, Central Pacific.
Conidae.

springeri, Anadara Rehder and Abbott, 1951. Revista de
la Sociedad Malacologica "Carlos de la Torre" 8(2):54,
pi. 8, figs. 8-9. HoIot>pe USNM 597375, off Dry Tor-
tugas, Florida. Arcidae.

steamsi, Chlamt/s Dall, Bartsch and Rehder, 1938. Bem-
ice P Bishop Museum Bulletin 153:89, pi. 22, figs. 11-
12. Holot\pe USNM 495030, Waianae Quarn, Oahu,
Hawaii. PecHnidae.

tampaensis, Crassispira (Crassispirclla) Bartsch and
Rehder, 1939. Proceedings of the United States National
Museum 87( .3070): 136, pi. 17, figs. 5, 13. Holotype
USNM 493409, Tampa Ba>-, Florida. Turridae.

tedbatjeri, Natica {Ghjphepitheriui) Rehder, 1986. The
Nautilus 10()(3):112. Replacement name for Natica
(Glyplicpithcma) batjeri Rehder, 1986, preoccupied by
Natica {Naticina) baijeri Koperberg, 1931. See baijeri,
Natica Rehder, 1986. Naticidae.

ternjae, Helicina Rehder, 1942. Journal of the Washing-
ton Academy of Sciences 32(11):350, fig. 16. Holot^pe
USNM 536026 [incorrectly published as 539026], Chi-
riqui Province, Republic of Panama. Helicinidae.

thaanumi, Arcinella Dall, Bartsch and Rehder, 1938.
Bemice P. Bi.shop Museum Bullehn 153:117, pi. 33, figs.
1-4. Holotype USNM 484176, near Koko Head, Oahu,
Hawaii. Chamidae.

thaanumi, Mactra Dall, Bartsch and Rehder, 1938.
Bemice P. Bishop Museum Bulletin 153:172, pi. 44, figs.
1^. Holot\pe USNM 337324, off Kaanapali, Maui, Ha-
waii. Mactridae.

thaanumi, Mirapecten Dall, Bartsch and Rehder, 1938.

Bemice P. Bishop Mu.seum Bulletin 153:84, pi. 21, figs.
7-8. Holotvpe USNM 173195, off south coast of Mo-
lokai, Hawaii. Pectinidae.

thaanumi, Ostrea Dall, Bartsch and Rehder, 1938. Bem-
ice P Bishop Museum Bulletin 153:114, pi. 32, figs. 1-
4. Holotype USNM 484158, Mokuoloe Island, Kaneohe
Bay, Oahu, Haw;iii. Ostreidae.

thur.stoni. Fraguiu Dall, Bartsch and Rehder, 19.38.
Bemice P. Bishop Museum Bulletin 153:154, pi. 41, figs.
9-10. Holotvpe USNM 484189, Keokea, Hilo, Hawaii.
Cardiidae.

tita, Senwk Dall, Bartsch and Rehder, 1938. Bemice P
Bishop Museum Bulletin 153:177, pi. 45, figs. 1-4. Ho-
lotype USNM 484197, Keaukaha, Hilo, Hawaii. Seme-
lidae.

torrci, Actcon Aguayo and Redher, 1936. Memorias de
la Sociedad Cubana de Historia Natural 9(4):268, pi. 24,
fig. 8. Holotyi^e USNM 420977, La Chorrera, Habana,
Cuba. Acteonidae.

transversa, Ctcna Dall, Bartsch and Rehder, 1938. Bem-
ice P Bishop Museum BulleHn 153:131, pi. 35 figs. 9-
12. Holotvpe USNM 337409, off Waikiki, Oahu, Hawaii.
Lucinidae.

transversa, Poronu/a Dall, Bartsch and Rehder, 1938.
Bemice P. Bishop Museum Bulletin 153:224, pi. 57, figs.
5-6. Holot)pe USNM 333017 [incorrectly pubfished as
330317], off south coast of Molokai, Havyaii. Poromyi-
dae.

transversum, Microcarditim Rehder and Abbott, 1951.
Revista de la Sociedad Malacologica "Carlos de la Torre '
8(2):56, pi. 9, figs. 9-10. Holotvpe USNM 4857.39 [in-
correctly published as 485734], SSW of Marsh Island,
Iberia County, Louisiana. Cardiidae.

valerioi, Streptostyla (Streptostijla) Rehder, 1942. Jour-
nal of the Washington Academy of Sciences 32(11):351,
fig. 17. Holotvpe USNM 536020, Cervantes, Cartago
Province. Costa Rica. Spiraxidae.

vanhi/ningi, Arene Rehder, 1943. Proceedings of the
United States National Museum 93(3161):192, pi. 19,
figs. 15-16. Holotvpe USNM 536054, Sand Key, 8 miles
south of Key West, Florida. Turbinidae.

vanhyningi, Conus verrucosus Rehder, 1944. The Nau-
tilus 57(3): 106. Holotype USNM 537863, on beach off
Pompano, Broward Countv, Florida. Conidae.

vanhtjningi, Cumingia telUnoidcs Rehder 1939. The
Nautilus 53(1): 19, pi. 6, figs. 13-14. Holot>pe USNM
473123, Lower Matecumbe Key, Florida. Semelidae.

vargasi. Triphora Rehder, 1980. Smidisonian Contribu-
tions to Zoology 289:43, pi. 6, fig. 11. Holotvpe MNSH
200384, Easter' Island. Triphoridae.

vaughani, Septifer Dall, Bartsch and Rehder, 1938.
Bemice P. Bishop Museum Bulletin 153:50, pi. 9, figs.

Page 144

THE NAUTILUS, Vol. 113, No. 4

9-10. Holot>pe USNM 495045, Wailupe Quarry, Oaliu,
Hawaii. Mytilidae.

velascoi, Emarginula Rehder, 1980. Smithsonian Con-
tributions to Zoology 289:16, pi. 4, figs. 2-3. Holot\pe
USNM 756019, Hanga Papara, Easter Island. Fissurel-
lidae.

vcnustiila. Arcne Aguavo and Redher, 1936. Memorias
de la Sociedad Cubana de Historia Natural 9(4):264, pi.
24, figs. 1-3. Holotype USNM 420976, La Chorrera,
Habana, Cuba. Turbinidae.

veimstitlus, Chicorcus {Chicomurcx) Rehder and Wilson,
1975. Smithsonian Contributions to Zoologv 203:7, te.xt-
figs. 4-5, frontispiece figs. 2-3. Holotype USNM
707241, off SW coast of Tahuata, Marquesas Islands.
Muricidae.

verae, Noticoiius Rehder, 1947. The Nautilus 61(1): 19,
pi. 1, top figure. Holot)pe USNM 485562, Marco Beach,
Marco Island, Collier County, Florida. Naticidae.

vermiculatus, Naesiotus quitensis Rehder, 1940. The
NauHlus 53(4):117, pi. 13, figs. 17, 19. Holot\pe MCZ
64957, Agovan (near Baiios Tunguragua), Ecuador. Or-
thalicidae.

vcstalis, CrcpitaccUa Rehder, 1943. Proceedings of the
United States National Museum 93(3161): 194, pi. 20,
fig. 13. Holotype USNM 411896, off Ajax Reef, Florida.
Rissoidae.

vefiila. Area Dall, Bartsch and Rehder, 1938. Bemice P.
Bishop Museum Bulletin 153:17, text-fig. 5, pi. 2, figs.
1-2, 5-6. Holotype USNM 495025, near Kaelepulu
Pond, Oahu, Hawaii. Arcidae.

vitrea. Di/smea Dall, Bartsch and Rehder, 1938. Bemice
P Bishop Museum Bulletin 153:174, pi. 44, figs. 13-14.
Holotype USNM 173001, off south coast o^Molokai,
Hawaii. Psammobiidae.

waikikia, Limopsis Dall, Bartsch and Rehder, 1938.
Bemice P. Bishop Museum Bulletin 153:42, text-fig. 13,
pi. 7, fig. 1. Holot>pe USNM 484219, off Waikild. Oaliu,
Hawaii. Limopsidae.

ivaikikius, Pccten Dall, Bartsch and Rehder, 1938. Bem-
ice P. Bishop Museum Bulletin 153:95, pi. 24, figs. 5-6.
Holotvpe USNM 337523. off Waikild, Oahu, Hawaii.
Pectinidae.

wareni, Vitreolina Rehder, 1980. Smithsonian Contri-
butions to Zoolog\- 289:56, pi. 7, figs. 14-15. Holotype
CAS 59676, Easter Island. Eulimidae.

iiillianisonim. Tcrainacliia JDlinsoni Rehder, 1972. The
Vehger 15(1 ):8. figs. 1-2, 4. Holot\pe USNM 707229,
30 miles south of Tung-Chiang, Taiwan. Volutidae.

wihoni, Aiistraharpa Rehder, 1973. Indo-Pacific Mollus-
ca 3(16):267, pi. 237, figs. 1-2. Holot>pe WAM 3670,
NW of Rottnest Island, off Perth, W'estem Australia.
Harpidae.

III. PUBLICATIONS OF HARALD A. REHDER

Harald A. Rehder authored 169 research papers and
books over a span of 70 years (1924—1993). His first and
last papers were in The Nautihis, a journal in which he
published 55 articles. Dr Rehder published in a broad
spectnmi of journals and moUuscan newsletters on a
wide variet)' of topics and taxa. While he came to be
recognized as a gastropod systematist with a special in-
terest in the families Harpidae (6 publications) and Vol-
utidae (11 papers), Rehder began his career studying
land and freshwater gastropods (20 publications). He
also published on Polvplacophora, Bivalvia, Scaphopoda,
and Cephiilopoda, describing new taxa in each of these
molluscan classes. Dr Rehder devoted 35 pubfications
to documenting the molluscan fauna of the tropical Pa-
cific, and 44 papers to nomenclatural and bibliographic
issues.

The following list of publications is arranged in chro-
nological order, with dates of publication provided in
square brackets [ ] when thev could be determined.
Dates of papers published in The Nautilus are from
Coan and HarasewTch (1993). Publications of the Unit-
ed States National Museum are dated according to
Anonymous (1971).

Rehder, H. A. 1924 [15 Januarv]. A list of moUusks from
Intervale, N.H. [New Hampshire]. The Nautilus 37(3):
96-97.

Clench. W. J. and H. A. Rehder. 1930 [17 July]. A new
Huinbokltiana from Texas. The Nautilus 44(1):10-13, pi.
2, figs. 1-4.

Rehder, H. A. 1932a [9 April]. Two new Polyg\Tas from
northern Arkansas. The Nautilus 45(4):128-i31, text-
figs. 1-2, pi. 10.

Rehder, H. A. 1932b [9 April]. Pohjgt/ra kiowaensis. The
Nautilus 45(4): 141.

Rehder, H. A. 1933 [16 June]. A new Succinea from
Florida. The Nautilus 47(1):20-21, pi. 3, fig. 4.

Rehder, H. A. 1935 [24 April]. New Caribbean marine
shells. The Nautilus 48(4): 127-130, pi. 7, figs. 7-12.

Rehder, H. A. 1936 [.30 Januar\]. The tvpe o( Poh/ff/ra
Say The Nautilus 49(3):102-1()3.

Aguayo, C. G. and H. A. Rehder. 1936 [13 Januarv].
New marine moUusks from Cuba. Memorias de la So-
ciedad Cubana de Historia Natural 9(4):26.3-26S, pi. 24,
figs. 1-8.

Rehder, H. A. 1937a [4 May]. The type ofAmicula Gray
The Nautilus 50(4): 142.

Rehder, H. A. 19371i [4 Mav]. Congerio leucophacata
(Con.) in the Hudson River the Nautilus 5()(4):143.

Rehder, H. A. 1937c [3 Julv]. Rissoa hcnmidezi Aguayo
and Rehder. The Nautilus 51(l):35-36.

Rehder, H. A. 1937d [7 August]. Notes on the nomen-

M. J. Sweeney and M. G. Harasewych, 1999

Page 145

clature of the Trochidae. Proceedings of the Biological
Societs' of Washington 50:115-116.

Relider, H. A. 1937e [23 September]. The genus Xij-
inonis Gebler. The Journal of Conchology (London)
20(12):370.

Dall, W. H., P Bartsch and H. A. Rehder. 1938. [25 July]
A manual of Recent and fossil marine pelecypod mol-
Kisks of the Hawaiian Islands. Bemice P. Bishop Mu-
seum Bulletin 153:1-233, pis. 1-58.

Rehder, H. A. 1939a [15 April], A new Ncuia from Haiti
and some notes on this genus. Journal of the Washington
Academy of Science 29(4):171-173, figs. 1-7.

Bartsch, P and H. A. Rehder. 1939a [26 April]. Two new
marine shells from the Aleutian Islands. The Nautilus
52(4):110-112, pi. 8. figs. 1-2.

Bartsch, P and H. A. Rehder. 1939b [13 June]. Mollusks
collected on the Presidential Cruise of 1938. Smithson-
ian Miscellaneous Collections 98(10); 1-18, pis. 1-5.

Rehder, H. A. 1939b [21 July]. New marine mollusks
from the West Atlantic. The Nautilus 53(1):16-21, pi. 6,
figs. 1-14.

Rehder, H. A. 1939c [21 July]. Micrelenchiis versus Gib-
hiuni. The Nautilus 53(l):33-34.

Rehder, H. A. 1939d [September]. Review of: Lichtig,
Ignaz.1938. Die Enstehung des Lebens durch stetige
Schopfung (The origin of life by constant creation),
Noord-Hollandsche Uitgeners Maatschappij, Amster-
dam. Journal of Paleontology 13(5):539-541.'

Bartsch, P and H. A. Rehder 1939c [15 September].
New turrid mollusks from Florida. Proceedings of the
United States National Museum 87(3070): 127-138, pi.
17, figs. 1-13.

Rehder, H. A. 1940a [29 April]. New mollusks of the
genus Nacsiotii.s from Ecuador. The Nautilus 53(4): 111-
118, pi. 13. figs. 1-20.

Bartsch, P and H. A. Rehder. 1940 [29 April]. Phhjcti-
concha. new name for a subgenus of Lijonsia. The Nau-
tilus 53(4):137.

Rehder, H. A. 1940b [28 June]. On the moUuscan genus
Trimusculus Schmidt 1818, with notes on some Medi-
terranean and West African Siphonarias. Proceedings of
the Biological Societ\- of Washington 53:67-69.

Rehder, H. A. 1940c [15 July]. A new urocoptid mollusk
from Mexico. Journal of the Washington Academy of
Science 30(7):315-316, figs. 1-3.

Rehder, H. A. 1940d [2 November]. Hijalotcuthis pdag-
icus (Rose) in the North Pacific. The Nautilus 54(2):71-
72.

Rehder, H. A. 1940e [2 November]. Note on Thericium
auricoma Schwengel and stantoni Dall. The Nautilus

54(2):72-73.

Rehder, H. A. 1941a [5 May]. The tvpe of Neptuuca
"Bolten" Rciding [in part]. The Nautilus 54(4): 126.

Rehder, H. A. 1941b [5 Mav]. Mtjrsus H. and A. Adams
1858, the valid name for Apolymetis Salisbuiy 1929. The
Nautilus 54(4):141-142.

Rehder, H. A. 1941c [24 October]. Correction re Mt/nsus
and Apolymetis. The Nautilus 55(2):64-65.

Rehder, H. A. 1942a [12 January]. A new subspecies of
Naesiotus quitensis from Ecuador. The Nautilus 55(3):
103.

Rehder. H. A. 1942b [May]. Geology and biolog\- of
North Atlantic deep-sea cores between Newfoundland
and Ireland. Part 5, MoUusca. United States Department
of the Interior, Geological Survey, Professional Paper
196-D: 107-109, 1 pi. 9" figures

Rehder, H. A. 1942c [14 October]. A note on the genus
Anaplocamm Dall. The Nautilus 56(2):49-50.

Rehder, H. A. 1942d [14 October]. The genot\'pe of
Neptunea. The Nautilus 56(2):69.

Rehder, H. A. 1942e [14 October]. Dentahiim (Antalis)
pihbnji, new name. The Nautilus 56(2):69.

Rehder, H. A. 1942f [15 November]. Some new land
shells from Costa Rica and Panama. Journal of the Wash-
ington Academy of Science 32(ll):350-352, figs. 1-19.

Rehder, H. A. 1943a [20 Januar\]. New marine mollusks
from the Antillean Region. Proceedings of the United
States National Museum 93(3161):187-203, pi. 19-20.

Rehder, H. A. 1943b [16 June]. The molluscan genus
Trochita Schumacher with a note on Bicatillus Swainson.
Proceedings of the Biological Society of Washington 56:
41^6, pl.'3.

Rehder, H. A. 1943c [23 July]. Two new species of Dnj-
macus from Mexico. The Nautilus 57(l):28-29, pi. 6,
figs. 6-10.

Rehder, H. A. 1943d [23 July]. CorrecHons and ecolog-
ical notes on some recently described Florida marine
shells. The NauHlus 57(l):32-33.

Bartsch, P and H. A. Rehder. 1943a [1 October]. New
cones from the Hawaiian Islands. Proceedings of the Bi-
ological Societv of Washington 56:85-88.

Bartsch, P and H. A. Rehder. 1943b [30 October]. Notes
on the names Potcria. PtijchontchUs, and Apcwsioma.
The Nautilus 57(2):62-64.'

Rehder, H. A. 1944a [9 February]. A new VitriucUa from
Man'land. The Nautilus 57(3):97, pi. 9, figs. 1-2.

Rehder, H. A. 1944b [9 Februan]. Conns icmicosiis
vanhi/uiiijs^i, new subspecies. The Nautilus 57(3);1()6.

Rehder, H. A. 1944c [4 May]. A note on the names Zozia
and Azorinus. Proceedings of the Malacological Societ>'

of London 26(1) :22.

Page 146

THE NAUTILUS, Vol. 113, No. 4

Rehder, H. A. 1944d [24 November]. A new pettinid
shell from the Pacific Ocean, with a note on the genus
Pallium Schroeter The Nautilus 58(2):52-54, pi. 2, figs.
1-2.

Rehder, H. A. 1945a [21 March]. A new genus and spe-
cies of squids from the Philippines. Proceedings of the
Biological Societv' of Wasliington 58:21-26, pi. 3, figs. 1-2.

Bartsch, P and H. A. Rehder. 1945 [2 Juh]. The West
Adantic boring mollusks of the genus Mai~tesia. Smith-
sonian Miscellaneous Collections 104(11): 1-16, pi. 1-3.

Rehder, H. A. 1945b [26 July]. On the status, under
Article 25 of the International Code, of generic names
where the genera concerned are founded upon figs. onlv.
Z.N.(S.) 68. Bulletin of Zoological Nomenclature 1(5):
94-95.

Rehder, H. A. 1945c [10 August]. The measurements of
shells. Mollusca l(6):73-77, figs. 1-3.

Rehder, H. A. 1945d [6 September]. Buccimim zebra
Miiller, the type of Orthalicus. The Nautilus 59(1): 29-
31.

Rehder, H. A. 1945e [20 September]. Two new species
of Cirsotrema (Epitoniidae) from Florida. Proceedings
of the Biological SocietA' of Washington 58:127-130.

Rehder, H. A. 1945f [27 December]. A note of the Bol-
ten Catalogue. The Nautilus 59(2):50-52.

Rehder, H. A. 1945g [27 December]. A note on Mcga-
spira data Gould. The Nautilus 59(2):67.

Rehder, H. A. 1945h [27 December]. A note on Yohella
tulipa (Lamarck). The Nautilus 59(2):67-68.

Rehder, H. A. 1945i. The Chilean species of the mol-
luscan genus Pcronaeiis (Bulimulidae). Revista Chilena
de Historia Natural 48(1944):l-6.

Bartsch. P, H. A. Rehder and B. E. Shields 1946 [30
January]. A bibliograph\- and short biographical sketch
of Wilham Healev Dall. Smithsonian Miscellaneous Col-
lections 104(15):i-95, 1 plate

Rehder, H. A. 1946a [27 June]. Notes on some groups
in the Muricidae of the West Atlantic, with descriptions
of a new subgenus. The Nautilus 59(4):142-143.

Rehder, H. A. 1946b [5 September], .\dditional notes on
the dates of publication of Les Fonds de la Mer. Pro-
ceedings of the Malacological Societv of London 27(2):
74-75.

Rehder, H. A. 1947a [28 February]. Proposed su.spen-
sion of the Regies for the genotvpe of Enjcina Lamarck.
1805 (Class Pelecypoda, Order Heterodonta). Z.N.(S.)
69. Bulletin of Zoological Nomenclature l(9):221-222.

Rehder, H. A. 1947b [14 July]. A new species of Nati-
carius from Florida. The Nautilus 61(l):19-20, pi. 1 (top
figure).

Rehder. H. A. 1947c [14 July]. Cepaea nemoralis in Bos-
ton. The Nautilus 61(1):35.'

Rehder, H. A. 1947d [18 December]. Aiion ater (L.) in
Oregon. The Nautilus 61(2):70.

Rehder, H. A. 1949 [8 June]. Some land and freshwater
mollusks from the coastal region of Virginia and North
and South Carolina. The Nautilus 62(4):121-126.

Rehder, H. A. and R. T. Abbott. 1951a [15 January]. Two
new Recent cone shells from the Western Atlantic (Con-
idae). Journal of the Washington Academv of Science
41(l):22-24, figs. 1-7.

Rehder, H. A. 1951a [9 October]. Comments and list of
modem names, pp. 486-503. In: A. Julia Rogers. The
Shell Book; a Popular Guide to a Knowledge of the
Families of Living Mollusks, and an Aid to the Identi-
fication of Shells Native and Foreign. Revised edition,
Boston: C.T Brantord, xxi + 503 pages, 8 pis.

Rehder, H. A. and R. T Abbott. 1951b [12 November].
Some new and interesting mollusks from the deeper wa-
ters of the Gulf of Mexico. Revista de la Sociedad Ma-
lacologica "Carlos de la Torre" 8(2):53-66, pi. 8-9.

Rehder, H. A. 1952a [22 May]. Objection to M. Gilbert
Ranson's application in regard to the generic name "Gry-
phaca" Lamarck, 1801 (Class Pelec)-poda). Z.N.(S.) 365.
Bulletin of Zoological Nomenclature 6(6):188-189.

Rehder, H. A. 1952b [25 July]. Harold John Finlav
1901-1951. The Nautilus 66(1):30-31.

Rehder, H. A. and L. Forcart. 1952 [29 August]. On die
generic names "Plotia" Roding, 1798, and "Pijramidclla"
Lamarck, 1799, and the proposed validation under ple-
nary powers of the generic name "Pijramklella" La-
marck, 1799 (Class Gastropoda, Subclass Prosobran-
chia). Z.N.(S.) 651. Bulletin of Zoological Nomenclature
6(ll):346-347.

Rehder. H. A. 1952c [17 November]. The publication
dates of Kobelt's " Ulustriertes Conchylienbuch". The
Nautilus 66(2):59-60.

Rehder, H. A. 1953 [2 Februarv]. The position of
"Xcsta" cincta (Lea). The Nautilus '66(3):95-96.

Rehder, H. A. 1954. Mollusks, pp. 469-474. In: Gulf of
Mexico — its origin, waters, and marine fife. Fishery Bul-
letin of the Fish and Wildhfe Senice 89:1-604.

Rehder, H. A. 1955a. A short histon- of the mollusk col-
lection of the U.S. National Museum. Annual Reports
of the Bulletin of the American Malacological Union for
1954:6.

Rehder, H. A. 1955b [30 December]. The genus Tiir-
cictila Diill. Proceedings of the Malacological Societv of
London 31(5-6):222-225, pi. 12, figs. 1-9.

Rehder, H. A. 1956 [May]. Review of: Perry, Louise M.
and Jeanne S. Schwengel. 1955. Marine shells of the

M. J. Sweeney and M. G. Harasewych, 1999

Page 147

western coast of Florida. Paleontological Research In-
sHtution, Ithaca. The Scientific Monthly 82(5):272-273.

Rehder, H. A. 1959a [Septeniher]. Lidentite de llle
Grimwood de Hugh Cuming. Bulletin de la Societe des
Etudes Oceaniennes (Polynesie Orientale) l(2-3):46-48.

Rehder, H. A. 1959b [IS December]. Review of: Keen,
A. MvTa, 1958. Sea Shells of Tropical West American,
Marine Mollusks from Lower California to Colombia.
Stanford Universit\^ Press, Stanford. Science, 130(3390):
1704.

Rehder, H. A. 1960 [1 Januan]. Marine zoogeography,
with special reference to the South Pacific area. (Ab-
stract). Annual Reports of the Bulletin of the American
Maiacological Union for 1959:9.

Rehder, H. A. 1961a [11 January]. Orthalicus in the Cay-
man Islands. The Nautilus 74(.3): 122-123.

Rehder, H. A. 1961b [Dated 1 Januar\-; correct pubU-
cation date is in Febniar)]. Dr Paul Bartsch 1871-1960.
Annual Reports of the Bulletin of the American Maia-
cological Union for 1960:4-6, portrait.

Kellogg, R. and H. A. Rehder 1961 [Dated I January-
correct publication date is February']. Smithsonian In-
stitution (USNM). Annual Reports of the Bulletin of die
American Maiacological Union lor 1960:12.

Rehder, H. A. 1961c. Report on maiacological research
in Antarctica, pp. 89-93. In: Science in Antarctica. Part
I: The life sciences in Antarctica. National Academy oi
Sciences, Natural Resource Council Publication 839, 162
pp.

Rehder, H. A. 1961d [14 July]. Obituary: Paul Bartsch.
Journal of Conchology 25(1):41^3, portrait.

Rehder, H. A. 1961e [10 November]. Comments on the
proposal to place the generic name Gari Schumacher,
1817, on the Official List unemended. Z.N.(S.) 1461.
Bulletin of Zoological Nomenclature 18(5):297-298.

Rehder, H. A. 1962a [9 Januar\']. The status o( Niicclla
Roetbng. The Nautihis 75(3):l69-lll, pi. 10, figs. 1-4.

Rehder, H. A. 1962b [June]. The Pleistocene mollusks
of Grand CaMiian Island, with notes on the geolog)' of
the island. Journal of Paleontology .36(3):58.3-585.

Rehder, H. A. 1962c [15 December]. Atoll news and
comments: Polynesia. Atoll Research Bulletin 94:16.

Rehder, H. A. 1963a [Januarv']. Contribucion al conoci-
miento de los moluscos marinos del Archipielago de Los
Roques y La Orchilla. Memoria de la Sociedad de Cien-
cias Naturales La Salle 22(62)(May-August 1962): 116-
138, 6 figs.

Rehder, H. A. 196.3b [26 April]. Pisania Bivona, 1832
(Mollusca: Gastropoda): its t\pe species, and proposed
addition to the Official List of Generic Names in Zool-
ogy-. Z.N.(S.) 1521. Bulletin of Zoological Nomenclature
20'(3):21.5-216.

Rehder. H. A. 1964 [January]. Personality of the
month — Harald A. Rehder. Hawaiian Shell News 12(3);
4, photograph.

Rehder, H. A. 1964 [March]. To Taliiti for mollusks. Ha-
waiian Shell News 12(5):4-5, map.

Rehder, H. A. 1964 [April]. Notes on Hawaiian Muri-
cidae. Hawaiian Shell News 12(6):4— 5, 8 figs.

Rehder, H. A. 1964 [May]. Marine mollusks of Polyne-
sia— a research project. Hawaiian Shell News 12(7):6-7.

Rehder, H. A. 1964 [August]. A further note on Hoin-
ahcaniha. Hawiiiian Shell News 12(10):2.

Rehder, H. A. 1964 [September] Dr Rehder on Pacific
tour. Hawaiian Shell News 12(11):6.

Rehder, H. A. 1966a [June]. Through the Tuamotus to
Pitcaim for mollusks. Part I. Hawaiian Shell News 14(8):
1, 4-5, 1 figure

Rehder, H. A. 1966b [June]. Let us collect yvith mod-
eration. Hawaiian Shell News 14(8):2.

Rehder, H. A. 1966c [July]. Through the Tuamotus to
Pitcaim for mollusks. Part II (Conclusion). Hawaiian
Shell News 14(9):l-2, 1 figure

Rehder, H. A. 1966d [14 October]. Comment on the
proposal to reject Mitrci pcrlata (Roding), 1798 (Gastro-
poda). Z.N.(S.) 1726. Bulletin of Zoological Nomencla-
ture 23(4): 1.33.

Rehder, H. A. 1966e [14 October]. Comment of the re-
quest for action on the name Valuta mitra Linnaeus,
1758 (Gastropoda). Z.N.(S.) 1728. Bulletin of Zoological
Nomenclature 23(4): 146.

Rehder, H. A. 1966f [1 December]. The non-marine
mollusks of Quintana Roo, Mexico, with the description
of a new species oi' Dn/nidcus (Puimonata: Bulimulidae).
Proceedings of the Biological Society of Washington 79:
27.3-296, figs. 1-20.

Rehder, H. A. 1966g [December]. Cijpraea in the Tu-
buai (Austral Islands). Hawaiian Shell News 14(14):2, 7.

Rehder, H. A. 1967a [February]. Collecting mollusks in
Tongatapu. Hayvaiian Shell News 15(2): 1, 7-8, 2 figs.

Rehder, H. A. 1967b [21 March]. Valid zoological names
of the Portland Catalogue. Proceedings of the United
States National Museum 121(.3579):1-51, 2 figures

Rehder. H. A. 1967c [12 April]. A neyv genus and two
new species in the families Volutidae and Turliineilidae
(Mollusca: Gastropoda) from the Western Pacific. Pacific
Science 21(2): 182-187, 11 figures

Rehder, H. A. 1967d [June]. Review of Morris, Percy
A. 1966. A field guide to shells of the Pacific Coast and
Hawaii including shells of the Gulf of California. Hough-
ton-Mifflin, New York. Atlantic Naturalist 22(2): 14.5-
146.

Rehder, H. A. 1968 [27 December]. The marine mol-

Page 148

THE NAUTILUS, Vol. 113, No. 4

luscan fauna of tlie Marquesas Islands. Annual Reports
of the American Malacological Union for 1968:29-32.

Rehder, H. A. 1969a [1 Januan.]. Voliitocorl)is and Fu-
siioltita, two genera of deepwater Volutidae from South
Africa. The Veliger ll(3):200-209, pis. 40-13.

Rehder, H. A. 1969b [Januarv]. New species and sub-
genera of Volutidae (Fulgorariinae) from the South Chi-
na Sea and Japan. Venus, The Japanese Journal of Mal-
acology 27(4):127-132, pi. 7, figs. 1-7.

Rehder, H. A. 1969c [25 November]. Ditle\ Thaanum,
1867-1963, a memorial sketch. Occasional Papers of the
Bemice P. Bishop Museum 24(3):27— 45, 5 figs.

Rehder, H. A. 1969. Preface, p. 1 and Appendix, pp. 97-
101, pis. 42^5. In: Afshar, F. (ed.) Ta.xonomic revision
of the superspecific groups of the Cretaceous and Ce-
nozoic Tellinidae. Geological Societ\' of America Memoir
119, 215 pages, 45 pis. ''

Rehder, H. A. 1970a [1 April]. Supplementar\' com-
ments on deep water Volutidae from the South China
Sea and South Africa. The Veliger 12(4):415-416, pi. 61,
figs. 1-6.

Rehder, H. A. 1970b [5 June]. Application to fi.\ the
name of the type-species of the genus Ampulla Roding,
1798 (olim Halia Risso, 1826) (Gastropoda: Volutidae").
Z.N.(S.) 1804. Bulletin of Zoological Nomenclature
27(l):41-43.

Rehder, H. A. 1970c [10 August]. Comment on the re-
quest for validation of Miircx lotoriuiu Linnaeus, 1758
(Gastropoda) in its accustomed sense. Z.N.(S.) 1886.
Bulletin of Zoological Nomenclature 27(2):67.

Rehder, H. A. 1971a [9 February]. A molluscan faunule
from 200 meters off Valparaiso, Chile, with descriptions
of four new species. Proceedings of the Biological So-
ciety of Washington 83(51):585-595, figs. 1-8. '

Rehder, H. A. 1971b. Malacological expedition to the
tropical South Pacific. National Geographic Society- Re-
search Reports, 1965 Projects:213-218.

Rehder, H. A. 1972a [1 May]. Comment on proposal to
remove homonNiny of Cassididae and Haqiidae in Mol-
lusca and Arthropoda. Z.N.(S.) 1938. Bulletin of Zoolog-
ical Nomenclature 29(1):2.

Rehder, H. A. 1972b [1 July]. Some notes on the genus
Teramachia (Volutidae: Calliotectinae). The Veliger
15(1):7-10, figs. 1-7.

Rehder, H. A. 1972c [1 July]. Further comments on
deepwater Volutidae from off Southeast Africa, with de-
scriptions of hvo new species of Vohifocorhis. The Ve-
hger 15(1): 11-14, figs. 1-9.

Rehder, H. A. and H. S. Ladd. 1973 [25 February].
Deep and shallow-water mollusks from the central Pa-
cific. Science Reports of the Tohoku University (sen 2,
Geologv), special volume 6:37-49, 2 text-figures, pi. 3,
figs. 1-18.

Rehder, H. A. 1973a [6 July]. Comment on the proposals
concerning family names Cassidae and Harpidae.
Z.N.(S.) 1938. Bulletin of Zoological Nomenclature
30(1):3.

Rehder, H. A. 1973b [6 July], Nipponaphera Habe, 1961
(Gastropoda): proposed designation of the t\pe-species
under the plenan,' powers. Z.N.(S.) 2007. Bulletin of
Zoological Nomenclature 30(l):37-38.

Rehder, H. A. 1973c [20 Julv]. Preface, p. Hi and Paul
Bartsch, 1871-1960, pp. 1-9^ In: Ruhoff F. A. Bibliog-
raphy and zoological taxa of Paul Bartsch. Smithsonian
Contributions to Zoologv 143:1-166.

Rehder, H. A. 1973d [10 October]. Comments on the
problem of the t\pe species of Liicina (Mollusca: Pele-
cxpoda). Z.N.(S.) 2001. Bulletin of Zoological Nomen-
clature 30(2):71-73.

Rehder, H. A. 1973e [27 November]. The family Har-
pidae of the world. Indo-Pacific Mollusca 3(16):20'7-274,
65 pis.

Rehder, H. A. and C. S. Weaver. 1974 [30 April]. A new
species of Vohitocorbis from South Africa. The Nautilus

88(2):31-32, figs. 1-8.

Rehder, H. A. 1974a [30April]. On the genus Vohitocor-
bis with descriptions of two new species from South Af-
rica. The Nautilus S8(2):33-37, figs. 1-8.

Rehder, H. A. 1974b [31 Julv]. Comment on the request
for the designation of a tspe-species of Tuttifa Jousseau-
me, 1881. Z.N.(S.) 2021.' Bulletin of Zoological Nomen-
clature 31(1):11-12.

Rehder, H. A. 1974c [11 September]. Marine biological
research in southeastern PoKmesia. National Geographic
Society Research Reports, 1967 Projects:243-254, 5 figs.

Rehder, H. A. 1974d. Mollusca, pp. 209-214. In: Biyan,
E. H., Jr. (ed.) Panala'au Memoirs. Pacific Science In-
formation Center, Bemice P. Bishop Museum, Honolu-
lu.

Rehder, H. A. and J. E. Randall. 1975 [15 January]. Du-
cie Atoll: Its history, physiography and biota. Atoll Re-
search Bulletin 183, 40 pages, 29'figs.

Rehder, H. A. 1975a [17 Julv]. Corrections to recent
papers on new species of Vohttocorbis from South Af-
rica. The Nautilus 89(3):79.

Rehder, H.A. 1975b [22 September]. Comment on the
request for a ruling on the authorship of Co/k/.s nioluc-
ccnsis. Z.N.(S.) 2059. Bulletin of Zoological Nomencla-
ture 32(3): 133-134.

Rehder, H. A. 1975c [22 September]. Proposed amend-
ment to Opinion 740: correction of name number 2087
on the Official List of Specific Names in Zoology.
Z.N.(S.) 1521. Bulletin of Zoological Nomenclature

32(3): 143.

Rehder, H. A. and B. R. Wilson. 1975 [19 December].

M. J. S\veene\' and M. G. Harasewych, 1999

Page 149

New species of marine mollusks from Pitcaim Island and
the Marquesas. Smithsonian Contributions to Zoology
203:1-16, 10 text-figures, 1 color plate, figs. 1-13.

Rehder, H. A. 1976 [December]. Mollusca, pp. 70-73.
In: Amerson, A. B., Jr. and P. C. Shelton (eds.) The nat-
ural histor\- of Johnston Atoll, central Pacific Ocean.
Atoll Research Bulletin 192, 479 pp.

Rehder, H. A. 1977a [Febmarv]. Collecting mollusks in
Tongatapu. The Mollusk: South Florida Shell Club
15(2): .5-6. [Edited and abbreviated version of Rehder,
1967a]

Rehder, H. A. 1977b. Molluscs (excluding scallops), pp.
14-15, pi. 3. /,i; Carney, J. N. (ed.) Offshore dredging.
South Santo [New Hebrides]. Annual Report of the New-
Hebrides Condominium Geological Survey for the Year
1975.

Rehder, H. A. 1978a [Februar)]. Comments on the pro-
posed designation of a type-species for Picurocera Raf-
inesque, 1818. Z.N.(S.) 83. Bulletin of Zoological No-
menclature 34(4): 198.

Rehder, H. A. 1978b [May]. The coral route. Pacific Is-
lands Monthly, 49(5):8-9.'

Rehder, H. A. 1980a [19 June]. Note on Cemohorskys
application for the use of the pleanar\- powers to des-
ignate a type-species for the genus Dntpclla Thiele,
1925. Bulletin of Zoological Nomenclature 37(2):86-87.

Rehder, H. A. 1980b [5 August]. The marine mollusks
of Easter Island (Isla de Pascua) and Sala y Gomez.
Smithsonian Contributions to Zoology^ 289:1-167, 15
text-figures, 14 pis.

Rehder, H. A. 19Sla. Mollusks (Consulting Editor), pp.
557-619. pis. 94-108. In: Complete Field Guide to
North American Wildlife; Eastern Edition. New York:
Harper & Row Publishers, 714 pp.

Rehder, H. A. 1981b [July]. The Audubon Society Field
Guide to North American Seashells. New York: Alfred
A. Knopf, 894 pp., 705 figs.

Rehder, H. A. 1981c [29 October], A new species of
Vohitocorbis (Volutidae) from Somalia. The Nautilus
95(4):169-170, figs. 1-5.

Rehder, H. A. 1982a [May]. A note on CohiJ^ratia sov-
erbii (Reeve) and a comment on "Notopcpluni translu-
cidum?" Hawaiian Shell News 30(5):5.

Rehder, H.A. 1982b. Marine mollusks of some island
groups of Polynesia. National Geographic Society Re-
search Reports 14:541-548.

Rehder, H. A. 1983a [December]. A revised list of the
marine mollusks of Henderson Island, pp. 34-39. In:
Fosberg, F. R., M.-H. Sachet, and D. R. Stoddard (eds.)
Henderson Island (southern PoKiiesia): summary of cur-
rent knowledge. Atoll Research Bulletin 272, 47 pp., 12
pis.

Rehder, H. A. 1983b [December]. Marine Mollusca, pp.
18-20. In: Sachet, M.-H. (ed.) Natural hi.ston- of Mo-
pelia Atoll, Society Islands. Atoll Research Bulletin 274,
37 pp., 30 figs.

Rehder, H. A. 19S4a [1 April]. Cijpmea thomasi Crosse:
a valid species. Havviiiian Shell News 32(4): 1, 8, figs. 1-
13.

Rehder, H. A. 1984b [27 April]. The genus Brondelia
Bourguignat, 1862, and its t;L\onomic position (Gastro-
poda: Siphonariidae). The Nautilus 98(2):83-84.

Rehder, H. A. 1984c [April]. Cypraca thomasi Crosse: a
valid species. Levantina: A Malacological Newsletter 49:
563-565. [Reprint of Rehder (1984a")]

Rehder, H. A. 1984d [23 August]. Williamia Montero-
sato, 1884 (Mollusca, Gastropoda): proposed conserva-
tion. Z.N.(S.) 2237. Bulletin of Zoological Nomenclature
41(3):159-162.

Rehder, H. A. 1985 [31 October]. A new species of Cor-
aUiophihi (Gastropoda: Coralliophilidae) from southeast-
ern Pol>iiesia. The Nautilus 99(4):97-100, 4 figs.

Rehder, H. A. 1986a [31 Januan]. Joseph Rosewater
(1928-1985), a tribute and bibhographv. The Nautilus
100(1):9-13, 1 figure.

Rehder, H. A. 1986b [31 January]. Natica {Glyphcpithc-
ma) batjeri, new name. The Nautilus 100(1): 38.

Rehder, H. A. 1986c [30 July]. Natico {Glyplwpitlwma)
tcdbai/cii, a replacement name. The Nautilus 100(3):
112. ■

Rehder, H. A. 1986d [July]. Ct/praea thomasi Crosse re-
validated. Hawaiian Shell News .34(7): 12, 9 figs. [Incor-
rectl)' published as vol. .35, no. 6],

Rehder, H. A. 1986e [August]. [Lijiia kiniheii Cotton,
1932] in Shell Letters. Hawaiian Shell News, .34(8):7 [In-
correctly published as vol. 35, no. 7].

Rehder, H. A. 1986f [September]. Neiita picea — again.
Hawaiian Shell News .34(9):6, 1 fig. [Incorrectly pub-
lished as vol. .35, no. 9].

Rehder, H. A. 1987a. [11 June] Obituarv', Joseph RE.
Morrison, December 19, 1906-December 2, 1983. Mal-
acological Review, 19(1986): 121-122, 1 fig.

Rehder, H. A. 19871i. [11 June] Obituan-, Joseph Rose-
water, September 18, 1928-March 22, 1985. Malacolog-
ical Review, 19(1986): 12.5-126. 1 fig.

Rehder, H. A. and R. E. Petit. 1987 [23 March]. Haiya
aiiiculaiis Lamarck, 1822 (Mollusca, Gastropoda): pro-
posed conservation of the specific name. [Case 2563[.
Bulletin of Zoological Nomenclature 44(l):19-20.

Rehder, II. A. 1988. The Audubon Society Pocket Guide
to Familiar Seashells of North America. Knopf, New
York. 192 pp.

Rehder, H. A. 1990a [18 April]. Clarification of the iden-

Page 150

THE NAUTILUS, Vol. 113, No. 4

tity of the snail Margaiites grocnlandicus (Gmelin, 1791)
(Gastropoda: Trochidae). the Nautilus 10.3(4): 117-12.3,
13 figs.

Rehder, H. A. 19yOb. The World of Nature: Seashells.
Gallery Books, W.H. Smith Publishers, New York. 64 pp.

Rehder, H. A. 1992 [21 August]. Harpa cabriti Fischer,
1860, a replacement name for Harpa vcntiicosa La-
marck, 1816 (Gastropoda: Harpidae). The Nautilus
106(3): 123-124, 1 figure.

Rehder, H. A. 1993 [.5 Februan]. A new .species of Har-
pa from the Leeward Islands of Hawaii. The Nautilus
106(4):127-129, figs. 1-2.

At the time of his death, Dr Rehder was working on
a paper recounting die history- of the Division of Mol-
lusks at the National Museum of Natural Histor\', Smith-
sonian Institution.

ACKNOWLEDGMENTS

We are grateful to Richard Johnson, James H. McLean,
Clvde f" E. Roper, and to William Cox and Pamela M.
Henson of the Office of Smithsonian Institution Ar-

chives, for kindlv providing photographs, correspon-
dence and notes on various aspects of Dr. Rehder's life
and works. We also diank Richard E. Petit and Ellis
Yochelson for their comments on this manuscript.

LITERATURE CITED

Anonvmous, 1971. Publications of the United States National
Museum (1947-1970). Bulletin 298 (final volume in se-
ries). Smidisonian Institution Press, Washington DC. 77
pp.

Coan, E. V. and M. G. Harasevwcli. 199.3. Publication Dates
of The Nmitihis. The Nautiliis 106(4): 174-180

Harasewvch, M. G. and Y. 1. Kanton 1991. Rediscovery of Afor-
ginellona gigas (Martens, 1904), with notes on die anat-
omy and systematic position of the subfamily Marginel-
loninae (Gastropoda: Marginellidae) Nemouria 37:1-19

Petit, R. E. and R. Bieler. 1996. On the new names introduced
in the various printings of "Shells of the world in color"
[Vol. I by Tadashige Habe and Kiyoshi Ito; Vol. II by Tad-
ashige Habe and Sadao Kosuge]. M;Jacologia38(l-2):.3.5-46

Turner, R. D. 1966. A sunev and illustrated catalogue of the
Terechnidae (Mollusca: Bivalvia). Museum of Comparative
Zoologv, Cambridge, 26.5 pp.

In Mcmoriain

Constance E. Boone

December 3, 1917

September 13, 1999

THE NAUTILUS 113(4):151. 1999 Page 151

Notices

THE R. T. ABBOTT VISITING CURATORSHIP

The Bailey-Matthews Shell Museum is pleased to iinite applications for the 2000 R. T Abbott Visiting Curatorship.

The curatorship, establishetl origuially in accordance with the wishes of the late Dr R. T Abliott, Ff)unding
Director of the Shell Museum, is awarded annually to enable mollusk systematists to visit the museum for a period
of one to two weeks. Abbott Fellows will be expected, by performing collection-based research, to assist with the
curation of the portions of the museum's Mollusk Collection and to provide one evening talk for the general pubhc.
The museum collection consists of marine, freshwater, and land shells, with emphasis on SW Florida. A large
percentage of the records has been catalogued through a computerized database management system. A substantial
portion of the time will be available for research in the museum collection, but field work in SW Florida can be
arranged. The R. T. Abbott Visiting Curatorship is accompanied by a stipend of $1,500.

Interested malacologists are invited to send a copy of their curriculum vitae togeather with a letter detailing their
areas of taxonomic expertise and research olijectives, and to provide a tentative title ior their talk.
Send materials to:

Dr. Jose H. Leal, Director

The Bailey-Matthews Shell Museum

P O. Box 1580

Sanibel, FL 33957

Apphcations for the 2000 Visiting Curatorship should be sent no later than Januan' 31, 2000. The award will be
announced by February 28, 2000. Questions about the R. T Abbott Visiting Curatorship should be directed to Dr.
Leal at:

(941) 395-2233; fax (941) 395-6706, or e-mail: ileal@gate.net

THE NAUTILUS

Volume 113
1999

REVIEWERS FOR VOLUME 113

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@ This paper meets the requirements of ANSI/NISO Z39.48-1992 (Permanence of Paper).

MRl. WHOJ l.lliUARV

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