THE NAUTILUS
Volume 105
1991
' I
AUTHOR INDEX
Bail, P 159 Marshall, B. A 104
BiELER, R 39 Okutani, T 165
BoucHET, P 159 Paul, A. J 173
Braley, R. D 92 QuiNN, J. F., Jr 81, 166
Clifford, H. F 173 Reid, D. G 1,7
Emerson, VV. K 62 Rosenberg, G 147
GoLiKov, A. N 7 Salisbury, R 147
HouART, R 26 Sergievsky, SO 1
HuLiNGS, N. C 16 Sysoev, a. V 119
Kabat, a. R 39 Toll, R B 116
Kantor, Y.I 119 Vidrine, M F 152
Ledua, E 92 Wilson, J. L. 152
Lucas, J. S 92 Zaslavskaya, N. 1 1
NEW TAXA proposed IN VOLUME 105 (1991)
gastropoda
Calliotropis dentata Quinn, new species (Trochidae) 170
Calliotropis globosa Quinn, new species (Trochidae) 168
Echinogurges tuberculatus Quinn, new species (Trochidae) 170
Gaza olivacea Quinn, new species (Trochidae) 166
Lamellitrochus Quinn, new genus (Trochidae) 81
Lamellitrochus bicoronatus Quinn, new species (Trochidae) 87
Lamellitrochus carinatus Quinn, new species (Trochidae) 84
Lamellitrochus fenestratus Quinn, new species (Trochidae) 85
Lamellitrochus filosus Quinn, new species (Trochidae) 87
Lamellitrochus inceratus Quinn, new species (Trochidae) 88
Lamellitrochus suavis Quinn, new species (Trochidae) 87
Mirachelus acanthus Quinn, new species (Trochidae) 168
Littorina (Littorina) hasatka Reid, Zaslavskaya, and Sergeivsky, new species (Littorinidae) 1
Litturina (Neritrema) naticoides Reid and GoUkov, new species (Littorinidae) 8
Derniomurex (Trialatella) leali Houart, new species (Muricidae) 27
Favartria (Favartia) varimutabilis Houart, new species (Muricidae) 32
Trophon mucrone Houart, new species (Muricidae) 35
Lyria doutei Bouchet and Bail, new species (Volutidae) 160
Vexillum brunneolinea Rosenberg and Salisbury, new species (Costellariidae) 147
Vexillum elliscrossi Rosenberg and Salisbury, new species (Costellariidae) 149
Antiplanes ahijssalis Kantor and Sysoev, new species (Turridae) 125
Antiplanes dendrotiplicata Kantor and Sysoev, new species (Turridae) 127
Antiplanes gabbi Kantor and Sysoev, new name (Turridae) 122
Antiplanes habei Kantor and Sysoev, new species (Turridae) 141
Antiplanes hurilensis Kantor and Sysoev, new species (Turridae) 136
Antiplanes molojimai aquilonalis Kantor and Sysoev, new subspecies (Turridae) 140
Antiplanes obliquiplicata Kantor and Sysoev, new species (Turridae) 128
Antiplanes spirinae Kantor and Sysoev, new species (Turridae) 137
THE NAUTILUS
Volume 105, Number 1
March 14, 1991
ISSS 0028-1344
A quarterly devoted
to malacology.
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EDITOR-IN-CHIEF
Dr. M. G. Harasewych
Division of MoUusks
National Museum of
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CONSULTING EDITORS
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Department of Invertebrates
Field Museum of
Natural History
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Dr. Robert T. Dillon, Jr.
Department of Biology
College of Charleston
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Dr. William K. Emerson
Department of Living Invertebrates
The .American Museum of Natural
Historv
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Dr. Robert Hershler
Division of Mollusks
National Museum of
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Smithsonian Institution
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Dr. Richard S. Houbrick
Division of Mollusks
National Museum of
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Smithsonian Institution
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Mr. Richard I. Johnson
Department of Mollusks
Museum of Comparative Zoology
Harvard Universitv
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Dr. Aurele La Rocque
Department of Geology
The Ohio State Universitv
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% Department of Mollusks
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Department of Geology
Florida Atlantic University
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Museum of Zoologv'
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Dr. Ruth D. Turner
Department of Mollusks
Museum of Comparative Zoology
Harvard Universitv
Cambridge, MA 02138
Dr. Geerat J. Vermeij
Department of Geology
Universitv of California at Davis
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THE€7NAUTILUS
Volume 105, Number 1
March 14, 1991
ISSN 0028-1344
CONTENTS ^ -
j -r. ^J»;JfUtUi>
I LIBRARY ;
' 1
David G. Reid Liitorina kasatka. a New Species from the Kurile IslMdO O c iQQi !
Nadya I. Zaslavskaya and Okhotsk Sea l " /^ ^ '^y ' f 1
Sergei O. Sergievsky | ♦
Wqqcj.s Hole. Macs. — j-
David G. Reid Liitorina naticoides. New Species, with Notes on the
Alexander M. Golikov Other Smooth-shelled Liitorina Species from the
Northwestern Pacific 7
Neil C. Hulings Activity Patterns and Homing of Acanthopleura gemmata
(Blainville, 1825) (Mollusca: Pohplacophora) in the Rocky
Intertidal of the Jordan Gulf of Aqaba 16
Roland Houart The Southeastern Brazilian Muricidae Collected by RV
Marion-Dusfresne in 1987, with the Description of Three
New Species . 26
News and Notices 38
THE NAUTILUS 105(l):l-6, 1991
Page 1
Littorina kasatka, a New Species from the
Kiirile Islands and Okhotsk Sea
David C. Reid
Department of Zoology
The Natural Histor\ Museum
London SW7 5BD, U.K.
Nadya I. Zaslavskaya
Institute of Marine Biology
Academy of Sciences of tlie U.S.S.R.
Vladivostok 690032, U.S.S.R.
Sergei O. Sergievsky
Zoological Institute
Academy of Sciences of the U.S.S.R.
Leningrad 199034, U.S.S.R.
ABSTRACT
A new species, Littorina (Littorina) kasatka. is described from
the Kurile Islands and Okhotsk Sea It is similar in shell char-
acters to a smooth form of L. (S'critrcnw) sitkana. with which
it is sympatric, but is distinguished by the absence of mam-
milliform glands on the penis of the male and by the small
jell) gland of the female. It is predicted that L. kasatka produces
pelagic egg capsules and has planktotrophic development
Kctj words: Littorina. Pacific; Larval development.
INTRODUCTION
The generic name Littorina has in the past been applied
to man\ littorinids with reiativeK large, smooth or spi-
rally sculptured shells, that occur in the littoral zone.
However, a recent phylogenetic analysis of the Littorin-
idae, based largely on anatomical rather than shell char-
acters, has defined Littorit}a more narrowly as a mono-
phyletic clade of about 20 species (Reid, 1989). Species
of Littorina sensu stricto are found only in the northern
hemisphere, where they are characteristic inhabitants of
the eulittoral and littoral fringe on temperate and cold
temperate shores.
During the past two decades close investigation of the
reproductive anatomy of European Littorina species has
revealed the existence of three additional species, pre-
viously unrecognized b\ shell characters (review by Raf-
faelli, 1982; Johannesson & Johannesson, 1990). The bio-
chemical technique of enzyme electrophoresis has
confirmed that these species are genetically distinct (re-
view by Ward, 1990). More recently, a pair of sibling
species has been identified in the northeastern Pacific,
again using the dual approach of reproductive anatomy
and biochemistry (Murray, 1979; Mastro et ai, 1982). In
contrast, the Littorina of the northern and northwestern
Pacific are poorly known. This paper describes a new
species that was discovered during a genetic study of
Littorina kiirila Middendorff, 1848 from the Kurile Is-
lands and Vostok Bay (Zaslavskaya & Sergievsky, in press).
(Littorina ktirila is a junior synonym of L. sitkana Phi-
lippi, 1846, see note added in press in Reid, 1990a). Of
11 loci examined, alleles common to the two species were
foimd at only two, and the genetic identitv was conse-
quently low (Nei's (1978) genetic identity / = 0.065).
The new species was recognized independently during
anatomical studies of Littorina material from the Zoo-
logical Institute, Leningrad, as part of a systematic study
of all Littorina species in the Pacific (Reid, in prep).
Recognition of the new species in the field is difficult,
since shell form is similar to that of L. sitkana and several
other Littorina species, but both males and females dis-
play unique reproductive anatomy.
MATERIAL AND METHODS
The new species is described from six lots in the Zoolog-
ical Institute, Leningrad (ZIL), and one lot in the Zoo-
logical Museum, Moscow (ZMM), all of which are du-
plicated in the Natural History Museum, London
(BMNH). That it is indeed undescribed was confirmed
by comparison with original descriptions of all available
names of Littorina species from the Pacific, and with all
available type specimens (housed in BMNH, ZIL and
National Museum of Natural History, Washington, D.C.);
details will appear elsewhere, in a full account of the
systematics of Littorina (Reid, in prep). Closely related
species are discussed briefly below.
The reproductive anatomy was examined in ten males
and nine females. The spiral pattern of the albumen and
capsule glands of the pallial oviduct can be seen, without
dissection, on the right side of the body adjacent to the
columellar muscle, when a female is removed from the
shell. Serial histological sections were prepared of the
pallial oviduct of two females. Radulae were extracted
from six specimens from three localities, and after clean-
ing in hot concentrated sodium hydroxide solution were
examined by scanning electron microscopy. For com-
parison, 45 specimens of L. sitkana were dissected, from
localities all around the northern Pacific from Hokkaido
to Washington, and four radulae were examined. Details
of other Littorina species investigated are given in Reid
and Golikov (1990).
SYSTEMATIC DESCRIPTION
Littorina (Littorina) kasatka new species
(figures 1-4, 8-11, 12-20, 23)
Page
THE NAUTILUS, Vol. 105, No. 1
Figures 1-4. Litlorina kasatha new species. 1. Holotype (HMNH 1990050), Kasatka Point, Iturup I., Kurile Is. 2. Paratype (BMNH
19900.51 ). 3. Nataliya Bay, Lirup I., Kurile Is. 4. Paramushir I,, Kurile Is. Figures 5-7. Litlorina silkana smooth form, from localities
at which it is s\mpatric with L kasatka. 5. 7. Kasatka Point, Iturup I , Kurile Is. 6. Nataliva Bay, Urup I., Kurile Is. All specimens
in BMNH.
Types: holotype BMNH 1990050, Kasatka Point, Iturup
I., Kurile Is, USSR (figure 1); 9 dry paratypes BMNH
1990051; 38 paratypes in alcohol BMNH 19900.52; 10
paratypes in alcohol ZIL.
Etymology: named after type locality.
Shell (figures 1-4): Dimensions: Adult size range 6.4-
11.0 mm shell height.
Shape: Teleoconch approximately 4 whorls (apices
eroded in available specimens), moderately solid. Glob-
ular to turbinate, whorls smoothly rounded, sutures dis-
tinct. Columella rather narrow, slightly e.xcavated, pillar
concave; sometimes a slight chink between edge of inner
lip and base of body whorl (not a true umbilicus).
Sculpture: Indistinct axial growth lines only, no spiral
grooves; faint spiral striae sometimes visible at high mag-
nification.
Color: Mid-brown, often with 6-16 rows of smudged
whitish dashes, especially at periphery and on base, oc-
casionally forming tessellated pattern (figure 4). Aperture
purplish brow II, Columella white to pinkish brown.
Animal: Head-foot (figures 19, 20): Head black, pale
tips to tentacles and snout, unpigmented stripe across
tentacle base and eye, or unpigmented patch at inside
of tentacle base only. Sides of foot grey to blackish.
Operculum: Normal, paucispiral littorinid (type A of
Bandel & Kadolsky, 1982).
Penis (figures 14-18): Wrinkled base lacking mammil-
liform glands; small, blunt filament not clearly demar-
cated from base; sperin groove open to tip.
Fallial oviduct (figures 12. 13): Complex spiral of 3 loops
oi all)iini('n and capsule glands; final .straight section
D. G. Reid et al., 1991
Page 3
Figures 8-11. Radulae of Littorina kasatka new species Scale bars = 50 fim, 8, 9. Kasatka Point, Iturup I , Kurile Is. 10. 11.
Paramushir 1 , Kurilt- Is 8. 10, N'ieweti ilat 9. I 1. Same radulae as Figures 8, 10, Init viewed troni angle ot 45° to show cusp
shape.
(containing jelly gland) short, not swollen or septate.
Bursa copulatrix small, opening in anterior position.
Raditla (figures 8-11): All cusps moderately pointed.
Rachidian with 3 large, 2 small cusps; outline of tooth
(viewed flat from above) varies from square to oblong
(ratio of length of tooth : width at mid-point 1.00-1.78).
Lateral and inner marginal each 4 large cusps. Outer
marginal 6-8 cusps.
Distribution: Habitat: Rocks and boulders in middle
and upper littoral zone of sheltered shores.
Range (figure 23):
Okhotsk Sea.
Kurile Islands, Gulf of Shelikov in
Records: Kasatka Bay and Kasatka Point, Iturup 1.; Na-
taliya Bay, Urup I.; Ryponkich I.; Paramushir I. (all ZIL
and BMNH); between Nepropusk and River Gank, W.
Kamchatka, Gulf of Shelikov (ZMM, BMNH).
Similar species: Four Littorina species are geograph-
icalK sympatric with L. kasatka over its known range:
L. (Littorina) squalida Broderip & Sowerby, 1829; L.
(Neritrema) sitkana; L. (N.) subrotundata (Carpenter,
1864) and a newly described species (Reid & Golikov,
1990). Of these, no confusion will arise with L. squalida.
which is a much larger (12-44 mm), spirally-grooved
shell (figured by Reid, 1990a). Littorina subrotundata
and the newly described species are also usualK larger
(4-14 mm and 6-16 mm respectively), both are a more
patulous shape, and both have a color pattern of dark
brown spiral lines on a paler brown ground (figured by
Reid & Golikov, 1990). Anatomically, these two species
both show mammilliform penial glands in the male and
a large, swollen, septate jelly gland in the female, unlike
L. kasatka.
Littorina sitkana is the most similar to L. kasatka and
is apparently found in the same habitat, since all but one
of the seven known collections of L. kasatka were sep-
arated from mi.xed lots of the two species. Littorina
sitkana is highly variable. The form that occurs most
commonly in the northeastern Pacific has strong spiral
ribs with striae in the grooves between, but in the north-
western Pacific (and especially in the Kurile Islands and
Okhotsk Sea) the common form is smooth shelled. This
smooth form has been referred to as L. kurila (this name
is actually a synonym of L. sitkana, see Reid, 1990a) in
all recent Soviet literature (e.g., Golikov & Kusakin, 1978).
Since L. kasatka lacks spiral ribs and obvious striations,
it is easily distinguished from sculptured shells of L.
sitkana (figured by Reid, 1990a). However, smooth forms
of L. sitkana are extremely similar. Littorina sitkana is
usually larger (commonly 15 mm, up to 23 mm), has a
Page 4
THE NAUTILUS, Vol 105, No. 1
Figures 12-20. Litiorina kasatka new species. 12, 13. Pallial oviducts. 11-18. Penes. 19, 20. Heads. 12. 17. Nataliya Bay,
Unip I. 13-15, 20. Kasatka Point, Iturup I. 16. Kasatka Bay, Ituriip I 18, 19. Paramushir I. Figures 21, 22. Littorina sitkana,
Seldovia, Alaska. 21. F'allial oviduct. 22. Two views of penis; mammilliform glands shaded .Ml specimens in BMNH Key to figures
12, 13, 21: sparse stipple, albumen gland; dense stipple, capsule gland; dotted lines, septa of jelK gland (visible by transparency);
dashed lines, bursa copulatri.v (visible only by dissection or .serial sectioning).
slightly more inflated last whorl, and the columella is
relatively wider (figures 5-7). Coloration also differs, L.
sitkana being purple brown or blackish, sometimes with
one or two white or orange spiral bands, and never show-
ing the flecked or tessellated pattern of L. kasatka. The
only entirely reliable means of distinguishing the two
species is by their reproductive anatomy: in females, the
pallial oviduct of L. sitkana has two loops in the spiral
pattern, and a large, swollen, septate jelly gland (figure
21 ), whereas there are three loops and a small jelK gland
in L. kasatka; in males, mammilliform penial glands are
present in L. sitkana (figure 22), but absent in L. kasatka.
Shells of the new species do not resemble closely those
of am other Littorina from the northeastern Pacific or
D. G. Reid et al.. 1991
Page 5
Figure 23. Distribution of Litlorina kasaiha new species Rec-
ord from Gulf of Slielikov could not be preciseh localized.
Atlantic. No other Littorina has a pallial oviduct of the
same form (Reid, 1990a), and only in L. scutulata Gould,
1849 and L. striata King and Broderip, 1832 does the
penis also lack mammilliform glands.
DISCUSSION
The diversity of types of spawn and development within
the genus Littorina have been reviewed by Reid (1989,
1990a), who showed that the type of spawn can be pre-
dicted from the relative sizes of the glandular compo-
nents of the pallial oviduct. From its small jelly gland,
lack of brood pouch and presence of a large capsule
gland, it is likely that L. kasatka produces pelagic egg
capsules. In other Littorina species that occur at high
latitudes and reproduce in this way, the capsules are large
(about 1 mm in diameter), biconve.x and contain several
eggs, and the capsules of L. kasatka are probably similar.
It has been suggested that large capsules of this type are
adaptive in relation to high levels of predation on pelagic
propagules in cold seas (Reid, 1990a). The only other
Pacific Littorina with pelagic capsules that occurs at such
a high latitude is L. squalida. All known littorinids with
pelagic capsules show planktotrophic development, so
this can be predicted in L. kasatka. In contrast, the other
four Littorina species in the northern Pacific are non-
planktotrophic, developing within benthic gelatinous egg
masses (Reid & Golikov, 1990).
A cladistic analysis of the 20 or so species of Littorina
then known was given by Reid (1990a), and used as a
basis for subgeneric classification. With three loops in
the spiral of the pallial oviduct, a small jelK gland and
probable pelagic egg capsules, L. kasatka is clearly a
member of the paraphyletic subgenus Littorina. Within
that area, its relationships cannot yet be resolved by
cladistic analysis, because of uncertainty about its spawn
The presence of a bursa copulatri.x in an anterior position
does, however, suggest that its most likely place on the
cladogram is between nodes 9 and 10 (see figure 5 in
Reid, 1990a). As a result of the cladistic analysis the
subgenus Neritrema was defined b\ the synapomorphies
of anterior bursa copulatri.x, large jelly gland and non-
planktotrophic development (Reid, 1990a). In view of
the anterior bursa of L. kasatka, this character must now
be removed from the list of synapomorphies of Neritre-
ma. The cladogram has also been used for a biogeo-
graphical analysis, which showed that in general the
more derived species of northwestern Pacific Littorina
occur at higher latitudes (Reid, 1990b). The probable
position in the cladogram and known geographical dis-
tribution of L. kasatka are consistent with this trend.
ACKNOWLEDGMENTS
A visit to the Soviet I'nion by DGR was supported by
the exchange program of the Royal Society of London
and the Academy of Sciences of the USSR. For permis-
sion to work at the Zoological Institute, Leningrad, DGR
would like to thank the director. Prof. O. A. Scarlato.
Prof. A. N. Golikov gave much valuable help with the
collections there. Photography \\asb\ H. Taylor (BMNH)
and histology by D. W. Cooper (BMNH). '
LITERATURE CITED
Bandel, K and D Kadolsky 1982. Western Atlantic species
of Nodilittorina (Gastropoda: Prosobranchia): compara-
tive morpliology and its functional, ecological, phyloge-
netic and taxonomic implications, X'eliger 25:1-42.
Golikov, A. N. and O. G. Kusakin. 1978. Recent molluscs of
the littoral zone of the U.S.S.R. Nauka. Leningrad, 256 p.
Johannesson, K. and B. Johannesson. 1990. Genetic variation
within Littorina saxatilis (Olivi) and Littorina neglecta
Bean: is L. neglecta a good species? Hsdrobiologia 193:
89-97.
Mastro, E., V. Chow, and D. Hedgecoek. 1982. Littorina
scutulata and Littorina plena, sibling species status of two
prosoliranch gastropod species confirmed by electropho-
resis. Veliger 24:239-246.
Murray, T. 1979. Evidence for an additional Littorina species
and a summary of the reproductive biology of Littorina
from California. Veliger 21:469-474.
Nei, M. 1978. Estimation of average heterozygosity and ge-
netic distance from a small number of individuals. Ge-
netics 89:583-590.
Raffaelli, D. G. 1982. Recent ecological research on some
European species of Littorina. Journal of Mollnscan Stud-
ies 48:342-354.
Reid, D. G. 1989. The comparative morphology, ph\ logeny
and evolution of the gastropod famiK Littorinidae. Philo-
sophical Transacations of the Royal Society of London,
Series B 324:1-110.
Reid, D. G. 199(ta. \ cladistic phylogeny of the genus Lit-
torina (Gastropoda): implications for evolution of repro-
ductive strategies and for classification H\drobiologia 193:
1-19.
Reid, D. G. 1990b Trans-Arctic migration and speciation
induced b\ climatic change: the biogeography of Littorina
Page 6
THE NAUTILUS, Vol. 105, No. 1
(MoUusca: Gastropoda). Bullt-tin of Marine Science 47:35-
49.
Reid, D. G. and A. N. Golikov. 1991. Liltorina nalicoides.
new species, with notes on the other smooth-shelled species
from the northwestern Pacific. Nautilus 105(1 ):7-15.
Ward, R D 1990 Biochemical genetic variation in tlie genus
I^ittorina (Prosofiranchia Mollusca) H\dr()liiologia 193:
53-69.
Zaslavskaya, N, 1 and S. O Sergievsky. In press. Genetic
comparison of Atlantic and Pacific species of Liitorina.
Proceedings of the 9th Malacological Conference of the
L'.S.S.R Leningrad: Zoological Institute,
THE NAUTILUS 105(1):7-15, 1991
Page 7
Littorina naticoides, New Species, with Notes on the Other
Smooth-shelled Littorina Species from the Northwestern Pacific
David G. Reid
Department of Zoology
The Natural History Museum
London SW7 5BD. U.K.
Alexander N. Golikov
Zoological Institute
Academy of Sciences of the L'.S.S.R.
Leningrad 199034, U.S.S.R.
ABSTRACT
A new species. Littorina (Neritrerna) naticoides. is described
from the northwestern Bering Sea, Kamchatka and Kurile Is-
lands. It is believed to be the sister species of L. aleutica and
the\ can be distinguished by penial and shell characters The
two are allopatric, L. aleutica showing an oceanic distribution
and L. naticoides a more continental one, but their ranges are
known to approach to u ithin 2.50 km of each other Five Lit-
torina species from the northwestern Pacific have smooth-shelled
forms, which can be difficult to distinguish. These are L. {N.)
naticoides, L. (N.) aleutica. L. (N.) sitkana. L. (N.) suhrotun-
data and L. (Littorina) kasatka. The diagnostic features of
their shells and reproductive anatomy are reviewed.
Key words: Littorina; Pacific; biogeography.
INTRODUCTION
Systematic studies of the family Littorinidae have been
revolutionised by two developments over the past two
decades. Firstly, anatomical features have been found to
be more reliable than traditional shell characters for
identification of species, and, secondly, there has been a
growing appreciation of the wide range of shell variation
shown by some species, and an understanding of its bi-
ological significance. The first use of anatomical char-
acters to discriminate between sibling species (with sim-
ilar or identical shells) was in the genus Littorina. (The
generic name is here used in the strict sense, as defined
by the cladistic analysis of Reid, 1989a). The most in-
formative of these new characters have proved to be the
shape of the penis, type of egg capsules and development
(Sacchi & Rastelli, 1966; Heller, 1975; Hannaford Ellis,
1979; Murray, 1979) and also the form of the pallial
oviduct (Reid, 1989a, 1990a). In all cases, studies of elec-
trophoretically detectable genetic variation have sup-
ported the status of the sibling species initially recognized
by anatomical differences (Ward, 1990). Although some-
times only one or other sex can be unequivocally iden-
tified by anatomical characters, there are no known cases
of morphologically inseparable Littorina species.
Extreme intraspecific variation in coloration, sculpture
and shape of the shell is a well-known feature of Lit-
torina. and contributes to the difficulties of identification.
Shell variation is especially marked in those species which
show direct development, for here the lack of a widely-
dispersed larval phase enhances the potential for adap-
tation to local environments. Selective factors influencing
morphological and color variation between populations
are believed to include predation, wave action and dam-
age by mobile boulders (e.g., review by Raffaelli, 1982;
Janson, 1982, 1983; Johannesson, 1986; Seeley, 1986).
Other factors such as growth rate can also contribute to
non-genetic shell variation (Kemp & Bertness, 1984).
The genus Littorina is restricted to the northern hemi-
sphere, and most species occur on temperate and cold
temperate shores (Reid, 1990b). So far, modern system-
atic revisions using anatomical details have only been
done in Europe and the northeastern Pacific. Littorina
species are also abundant in the northwestern Pacific,
and the most recent review of the littoral molluscs of the
Siberian region (Golikov & Kusakin, 1978) recognized
six species: L. squalida Broderip & Sowerby, 1829; L.
brevicnla (Philippi, 1848); L. rnandshurica Schrenck,
1861; L. aleutica Dall, 1872; L. sitkana Philippi, 1846;
L. kurila Middendorff, 1848. The identifications were
based only on characters of the shells. The first anatom-
ical work on northwestern Pacific Littorina was done as
part of a cladistic analysis of the 20 species then recog-
nized in the genus (Reid, 1990a). This largely supported
the classification of Golikov and Kusakin (although L.
kurila was synonymized with L. sitkana), but the ana-
tomical material used did not include any from the Soviet
Union.
As part of a wider study of the systematics and dis-
tribution of Littorina species (Reid, in prep), it has re-
cently been possible to examine the extensive collection
of preserved material from the Siberian coast held in the
Zoological Institute, Leningrad. Preliminary results in-
dicate that L. squalida. L. brevicnla and L. rnandshurica
can each be readily identified by shell characters, as
described and illustrated by Golikov and Kusakin (1978)
and Reid (1990a). The remaining three of the species
listed above were distinguished by Golikov and Kusakin
(1978) primarily by their shell sculpture: L. sitkana hav-
ing spiral ribs, L. kurila being smooth or almost so, and
L. aleutica having rows of nodules. This division now
seems to be an artificial one. Differences in reproductive
Pages
THE NAUTILUS, Vol. 105, No. 1
anatomy define five .species in this group: L. sithana, L.
alentica, L. sitbwtundata (Carpenter, 1864), L. hasatka
(recently described by Reid et al., 1990) and L. nati-
coides, which is described herein. The shell characters
of all five species are variable, and in particular L. sit-
kana. L. aleutica and L. stihrottindata can show smooth
or sculptured shells. "Littorina kurila," as defined by
Golikovand Kusakin (1978), embraced smooth or slightly
sculptured forms of all five species.
The nomenclature of these species is rather complex.
During the wider study of Littorina systematics all avail-
able type specimens (housed in Natural History Museum,
London; National Museum of Natural History, Wash-
ington, D.C.; Zoological Institute, Leningrad) have been
examined and all original descriptions consulted, to as-
certain that the new species is indeed undescribed. The
available names include 18 synonyms of L. sitkana and
three of L. subrotundata. Complete synonymies and full
accounts of the described species w ill be given elsewhere.
Here it need only be noted that preserved syntypes of
L. kurila Middendorff, 1848 in the Zoological Institute,
Leningrad, have shown that this taxon is a synonym of
L. sitkana (see note added in proof in Reid, 1990a). As
discussed below, the name "L. kurila" was incorrectly
applied by Reid (1989a, 1990a, b) to what is here con-
sidered an open-coast form of L. subrotundata.
In the present paper the new species will first be de-
scribed, then the four other similar species which some-
times show smooth shells will be discussed and distin-
guished.
MATERIALS AND METHODS
This resolution of the smooth-shelled Littorina species
of the northwestern Pacific is based on examination of
all material in the collections of the Natural History
Museum, London (BMNH), the Zoological Institute,
Leningrad (ZIL) and the National Museum of Natural
History, Washington, D.C. (USNM). The new species is
described from 21 lots in the ZIL, many of which are
duplicated in the BMNH. A total of 27 males and 18
females were dissected, from 8 localities covering the
known range of the species. Five radulae were extracted
from specimens from three localities, and after cleaning
in hot, concentrated sodium hydroxide solution were ex-
amined by scanning electron microscopy. For compar-
ison, 32 specimens of L. aleutica, 82 of L. subrotundata,
45 of L. sitkana and 19 of L. kasatka were dissected, all
from a wide range of localities.
SYSTEMATIC DESCRIPTION
Littorina (Neritrema) naticoides new species
(figures 1-12, 15-23, 28-35, 37)
Types: holotype BMNH 1990053, Milne Bay. Simushir
1., Kurile Is, USSR (figures 1, 16); 2 drv paratypes BMNH
1990054; 6 paratypes in alcohol BMNH 1990055; 2 para-
types in alcohol ZIL.
Etymology: Natica-\ike, a reference to shell shape.
Shell (figures 1-12): Dimensions: Adult size range
6.2-15.9 mm shell height.
Shape: Teleoconch approximately 4 whorls, moderately
solid. Turbinate, often rather patulous, whorls smoothly
rounded, sutures distinct. Ciolumella relatively wide,
slightly excavated, pillar concave.
Sculpture: Protoconch 0.73-075 mm diameter, 1.8
whorls. Teleoconch: indistinct axial growth lines only, no
spiral grooves; faint spiral striae sometimes visible at high
magnification, but no strong striations.
Color: Orange brown to dark purple brown, usually 5-
12 dark brown or black spiral lines; sometimes a pale
spiral band on base. Aperture orange brown to purple
brown. Columella white, sometimes tinged purple brown.
Animal: Head-foot (figures 28-30): Head grey to black,
paler at tips of tentacles and snout; unpigmented patch
over eye and usually another at inside of tenacle base.
Sides of foot usually pale grey speckles or unpigmented,
sometimes blackish.
Operculum: normal, paucispiral littorinid (tvpe A of
Bandel & Kadolsky, 1982).
Penis (figures 15-20): Wrinkled base with 4-15 mam-
illiform penial glands in single row on anterior edge (one
specimen with a single gland); filament short, less than
half length of base, broadly triangular, with simple sub-
epithelial glandular region (more opaque in preserved
specimens); sperm groove open to tip.
Pallial oviduct (figures 21-23, 37): Complex spiral of 3
loops, first of albumen gland, second and third of capsule
gland; final straight section a large, swollen, septate jelly
gland. Bursa copulatrix in anterior position.
Spawn: Benthic, gelatinous egg mass up to 6 mm in
diameter, containing spherical capsules 0.7-0.8 mm in
diameter, each with single egg surrounded by albumen
layer 0.5-0.6 mm in diameter.
Radula (figures 31-34): All cusps bluntly rounded. Ra-
chidian with 3 large, 2 .small cusps; outline of tooth (viewed
flat from above) varies from square to oblong (ratio of
length of tooth: width at mid-point 0.77-2.0). Lateral
and inner marginal teeth each 4 large cusps. Outer mar-
ginal 5-7 cusps.
Distribution: Halntat: A range of intertidal habitats
are reported: on dead shells in sheltered inlets; Mytilus
Figures 1-12. Littorina naticoides new species. 1. Holotype (BMNH 1990053), Milne Hay. Simushir I., Kurile Is. 2, 3. Ozernovskiy,
Kamchatka. 4. 5. 10 km east of Cape Kamchatski\, Kamchatka. 6. Avacha Ba>, Kanu hatka 7. Kronot.skiy Gulf, Kamchatka 8-
10. Egvekinot Inlet, .'\nadyr.skiy (iiiif II, 12. Ozcriiiy River e.stuary, Kamchatka Figure I.'l. Littorina aleutica. Provideniya.
Figure 14. Littorina stilnotundata. Milne Bay, Simushir 1 , Kurile Is (.All specimens in BMNH).
D. G. Reid and A. N. Golikov, 1991
Page 9
Page 10
THE NAUTILUS, Vol. 105, No. 1
15-27 28-30
Figures 15-23. 28-30. Littorina naticoides new species. 15-20. Penes: mamillifdrm j;lands on left edge, sometimes visible by
transparency 21-23. Pallial oviducts 28-30. Heads. 15, 17. 18. Ozernovski\, Kamchatka 16. Penis of holot\pe, Milne Bay,
Simusliir I , Kiirile Is. 18. 19. 23, .30. Krista Gulf, Anadyrskiy Gulf 20. F.gvekinot Inlet, .\nadyrskiy Gulf 21.' 29. Kronotski'y
Ba>, Kamclialka. 22. 10 km east of Gape Kanicliatskiy, Kamchatka Figures 24-27. Littorina alculica. 24-26. Penes 24, 26.
Lihaclieva, Providcniya. 25. Adak I., Aleutian Is. 27. Pallial oviduct, Nizki I., Aleutian Is. Key to figures 21-23, 27: sparse stipple,
albumen gland; dense stipple, capsule gland; dotted lines, septa of jelly gland (visible by transparency); dasfied lines, bursa copulatrix
(visible only by dissection).
zone on exposed shores; on Halosaccion (red alga); in
Alaria (brown alga) bell.
Range (figure 3.5;.Kurile Islands, .south and wesi Kam-
chatka, Anadyrskiy Gulf.
Records: Kiirile Is: Urnp I.; Milne Bay, Simushir I.;
Spaseniya, Simushir I.; Diani Bay, Ketoi I., Paramushir
I.; Kamchatka: Ozerniy River estuary, Okhotsk Sea;
Ozerriovskiy; Avachinskaya Bay, near C^ape Viikova;
Avacha Bay, near Petropavlovsk; Kronotskiv Gulf; 10 km
east of Cape Kamchatskiv; .Anadyrskiy Gulf: Egvekinot
Inlet. Krista Gulf (ail ZIL and BMNH).
D. G. Reid and A. N. Golikov, 1991
Page 11
Figures 31-34. Radiilae of Littorina naticoidcs new species Scale bars = 50 ixm. 31, 32. 10 km east of Cape Kamchatskiy,
Kamchatka. 33, 34. Egvekinot Inlet, Anadyrskiy Gulf. 31, 33. Viewed flat, 32, 34. Same radulae as figures 31, 33, but viewed
from an angle of 45° to show cusp shape.
Similar species: As mentioned above, L. kurila Mid-
dendorff, 1848 is a junior synonym of L. sitkana Philippi,
1846, but the name has been widely used in the literature
for any of the five Littorina species (including L. nati-
coides) in the northwestern Pacific that frequently have
smooth shells. The other four members of the group will
be considered in turn, and the characters of all five spe-
cies are summarized in table 1. Radular characters have
not proved useful in discriminating between them.
L. (Neritrema) .sitkana (figure 39): The typical form of
this species has a strongly carinate shell with spiral stri-
Tabh' 1. Sununar\ of characters of five Littorina species from the northwestern Pacific.
sultro-
aleulica
kasatka
naticoides
sitkana
Inndala
Shell:
adult size (mm)
6-14
fi-11
6-16
6-23
4-14
patulous shape
+
-
+
-
( + )
nodulose sculpture
( + )
—
-
-
-
spiral ribs
{ + )
-
-
( + )
( + )
spiral microstriae
+
—
-
( + )
-
pattern of dark spiral lines
{ + )
—
+
( + )
+
pattern of pale flecks
-
+
-
-
-
Penis:
niamilliform glands
7-12
0
1-15
5-14
5-17
filament > '2 length of base
+
-
-
-
-
Pallia!
oviduct:
spiral pattern
fig. 38
fig. 36
fig. 37
fig. 39
fig. 40
swollen, septate jelly gland
+
—
+
+
+
Spawn
benthic egg mass
+
-
+
+
+
pelagic capsules
-
-
-
—
+ = Present; ( + ) =
= sometimes present: - = absent.
Page i;
THE NAUTILUS, Vol. 105, No. 1
Figure 35. Distriliution of Littorina naticoides new species.
ations between the carinae (e.g., Reid, 1990a: fig. 11)
and is most frequent in the north and northeastern Pacific
from Oregon to the Aleutian Islands, while in northern
Japan it is usually only the base that bears strong grooves.
In either case the shell is thereby distinguished from all
others in the group except rare forms of L. subrotundata.
Problems arise with the smooth forms of this species
(Reid et ai, 1990: figs 5-7), which predominate in the
Kurile Islands and Kamchatka, and also occur in other
parts of the range. These shells may be quite large (up
to 23 mm), are less patulous than L. naticoides and L.
aleutica and are often uniformly black or brown (except
in the Okhotsk Sea, L. sitkana commonly lacks the pat-
tern of narrow black lines seen in L. naticoides and L.
subrotundata). Synonyms based on smooth forms of this
species include L. kurila Middendorff, 1848, L. suhtene-
brosa Middendorff, 1848 and L. sitkana var. atkana (Dall,
1886). Females of L. sitkana can be immediately rec-
ognized by the unique spiral pattern of the pallial oviduct
(figure 39; Reid et ai, 1990: fig. 21) with its long back-
ward loop of capsule gland, but the penis is similar to
that of both L. naticoides and L. sulnutundata.
L. (Neritrema) subrotundata (figures 14, 40): There has
been confusion about the identity of this species, and the
name is here used in a wider sense than by previous
authors. As interpreted here, the species has a wide dis-
tribution around the northern Pacific, from northern Cal-
ifornia thruugli the Aleutian Islands to the soutiiern Ku-
rile Islands.
The name has previously been applied only to the thin-
shelled, tall-spired form (Reid, 1990a: fig. In) initially
described by Carpenter (1864) as a doubtful member of
the genus Assiminea. Synonyms are ?Faludinella cas-
tanea Carpenter, 1865, and Paludinella newcombiana
Hemphill, 1877, and the species has .sometimes been
placed in the genus Algamorda Dall, 1918 (a synonym
of Neritrema Rechiz, 1869, regarded as a subgenus of
Littorina. see Reid, 1989a). The species was poorly known
and lielieved to be restricted to brackish-water salt marshes
from northern California to Washington (MacDonald,
1969; Reid, 1990b). The form from this habitat is anal-
ogous to the form " tenelnosa" of the .Atlantic L. saxatilis
(Olivi, 1792), which is also small, thin-shelled and tall-
spired (Muus, 1967; Janson & Ward, 1985), probably as
a result of similar selection pressures or ecophenotypic
effects in the brackish lagoons in which it occurs. This
form of L. saxatilis contrasts with the lower-spired and
more patulous form on exposed coasts (e.g., Raffaelli,
1979; Janson, 1982).
As a result of the dissection of additional material, it
is now believed that, like L. saxatilis, L. subrotundata
also occurs in habitats covering a wide range of wave
exposures, and has a similarly wide range of variation in
shell morphology. The forms from exposed rocky coasts,
here assigned to L. subrotundata for the first time, have
(like L. saxatilis from similar habitats) a lower spire and
larger aperture than the salt-marsh form, and were pre-
viously incorrectly identified as L. kurila (Reid, 1990a,
b). A shell of this type from the Aleutian Islands was
illustrated b\' Reid (1990a: fig. Im), and figure 14 shows
a slightly more patulous form. A population from the
exposed shores of Tatoosh Island, Washington, has re-
cently been studied by Boulding (1990), who concluded
that it was an undescribed subspecies of L. kurila. How-
ever, examination of specimens from this locality has
revealed no significant anatomical differences from ei-
ther typical L. subrotundata collected in salt marshes in
Washington, or from specimens of this species from the
Aleutian Islands. Furthermore, no diagnostic allelic dif-
ferences between samples from these three areas have
been found by enzyme electrophoresis (E. G. Boulding,
personal communication). The available evidence there-
fore suggests that they are conspecific, but further study
would be desirable. Neither the known geographical dis-
tribution of L. subrotundata, nor its considerable vari-
ation in shell morphology throughout its range, support
its division into subspecies.
One reason why the name L. subrotundata has not
previously been used for the exposed-coast form is that,
at least in Washington, it appears to be completely eco-
logically segregated from the t>pical form in salt marsh-
es, because the species is not known to occur in inter-
mediate habitats. This locally disjunct distribution need
not, however, imply genetic isolation of the two forms.
It could be maintained, for example, by the action of
crab or other predators. It is known that the exposed-
coast form is susceptible to attack by crabs because of
its thin shell, and that crabs are more abundant in pro-
tected rocky habitats than on exposed coasts (Boulding,
1990). It is possible that both exposed coasts and salt
marshes are refuges for this species from crab predators.
Further north in its range, in Alaska (personal observa-
tion), L. subrotundata does occur on both exposed and
sheltered rocky shores and there is continuous variation
in shell shape l:)etween the two extremes of habitat. In-
D. G. Reid and A. N. Golikov, 1991
Page 13
36
37
38
39
40
Figures 36-40. Diagrammatic represfiitation of pallial oviducts of Littorina species from the northwestern Pacific- 36. Litlorina
(Litlorina) kasalka. 37. L. (S'eritrema) naticoides. 38. L. (N.) aleutica. 39. L. (N.) sitkana. 40. L. (iV.) suhroUmdala. Key;
continuous line with spiral loops represents path of egg groove through pallial oviduct; thick section, capsule gland; dense stipple,
seminal receptacle at posterior end; anterior sac, bursa copulatrix; dotted lines, septa of jelly gland. Shape of loops of capsule gland
and its relative size are most important features. Oviducts of L. aleutica and L. naticoides sometimes indistinguishable; c.f figures
21-23, 27,
terestingly, there is again a parallel with L. saxatilis,
which shows a comparable segregation of low-spired and
high-spired forms on exposed coasts and in salt marshes
respectively, at the southern limit of its European range
(Gofas, 1975; personal observation), whereas at more
northerly latitudes it occupies the entire spectrum of
habitats and morphological intermediates are common
(Janson & Ward, 1985; personal observation).
The shell of L. subrotundata is often extremely similar
to that of L. naticoides in size, coloration and shape,
although it does not attain such extreme patulous forms
as the latter (e.g., figures 3-5). Like L. naticoides, it is
usually entirely smooth, but occasional forms have spiral
ribs; in such cases the lack of strong spiral striations in
the grooves distinguishes it from L. sitkana. Once again,
anatomical characters are more helpful; the form of the
pallial oviduct is diagnostic (figure 40) with a relatively
smaller capsule gland than that of either L. sitkana or
L. naticoides, but the penis is similar to those of both
these species.
L. (Neritreryia) aleutica (figures 13. 24-27, 38): Shells
of L. aleutica from the .Aleutian Islands are usually sculp-
tured by four nodulose cords with strong spiral striations
in the intervening grooves (Reid, 1990a: fig. Ik). How-
ever, in the northwestern Bering Sea the shells are not
nodulose; slight spiral ribs may remain, as do the stria-
tions (visible at the periphery in figure 13), which help
to distinguish the shell from the very similarly shaped,
but entirely smooth, L. naticoides. In this case it is the
male anatomy that is diagnostic, the penis showing a
narrower filament at least half as long as the wrinkled
base, whereas that of L. naticoides is shorter (figures 24-
26, c.f. figures 15-20). The pallial oviduct is similar only
to that of L. naticoides. but the jelly gland is usually
relatively larger and the capsule gland a little smaller
(figures 22, 38, c.f. figures 21-23, .37).
L. (Littorina) kasatka: This newly-described species
(Reid et ai, 1990) is the smallest of the five smooth-
shelled Littorina species (6-11 mm) and has the most
restricted distribution, being recorded only from the Ku-
rile Islands and Gulf of Shelikov. The shell is closest to
small, smooth specimens of L. sitkana, although its shape
is subtly different, with a relatively narrower columella
and smaller aperture. Coloration also differs, the shell of
L. kasatka being brown, often with indistinct white flecks
or tessellation, unique among the five species discussed
here. Despite the superficial similarity of the shell, this
species is probably not closely related to the others in the
group, since its reproductive anatomy is markedly dif-
ferent. Mamilliform penial glands are absent in the male,
while the pallial oviduct lacks a swollen, septate jelly
gland and shows a unique spiral pattern of albumen and
capsule glands. These characteristics of the oviduct in-
dicate that the spawn consists of pelagic egg capsules,
probably with planktotrophic development, and for this
reason it is classified in the subgenus Littorina (Reid et
ai, 1990). In contrast, the four other species have a large
jelly gland, benthic egg masses and direct development,
and are therefore members of the subgenus Neritrema
(Reid, 1989a, 1990a).
DISCUSSION
An earlier review of living Littorina species with a cla-
distic analysis of their relationships (Reid, 1990a) must
be revised in the light of the new information on material
Page 14
THE NAUTILUS. Vol. 105, No. 1
from the Siberian coast. At least three changes are nec-
essary. Firstly, "L. kurila" {non Middendorif, 1848) of
the earlier publication is now believed to be the same as
L. sulrrotundata (no differentiating characters were found
in the earlier analysis). Secondly, L. kasatha must be
added to the cladogram (Heid 1990a: fig. 5), in a position
between nodes 9 and 12 (as yet unclear because of lack
of information about egg capsules, but between 9 and
10 most likely, Reid et al. 1990). Thirdly, L. naticoides
must be added; this is clearly the sister species of L.
aleutica, since with the exception of shell sculpture, all
characters used in the cladistic analysis are the same in
both species.
This sister-group relationship is supported by the sim-
ilar oviducts, egg masses, penial glands and patulous shells.
Consistent differences in the relative length of the penial
filament and in shell sculpture, and small differences in
relative sizes of oviducal glands, are, however, sufficient
to separate L. naticoides as a new species. So far, the
two species have not been found sympatrically. Littorina
aleutica is recorded from the Aleutian, Komandor and
Pribiloff Islands, and from St Lawrence Island and Provi-
deniya in the northern Bering Sea. Littorina naticoides
occurs from Anadyrskiy Gulf to the Kurile Islands, ap-
proaching within 350 km from the closest record of L.
aleutica in the northern Bering Sea, and within 250 km
in eastern Kamchatka. In part this apparent allopatry
could be explained by lack of collections from the remote
northwestern shores of the Bering Sea. However, the
separate distributions in the Aleutian, Komandor and
Kurile Islands and in Kamchatka are probably real, be-
cause there are extensive collections of Littorina from
all these areas. Furthermore, the two distributions suggest
a possible ecological separation. Littorina aleutica has
the more "oceanic" distribution, occurring only on is-
lands and at the tip of the Chukotskiy Peninsula, whereas
L. naticoides is more "continental," occurring on main-
land coasts, even in inlets and estuaries, as well as in the
Kurile Islands. There are similar examples elsewhere in
the Littorinidae of closely related congeners with con-
trasting "oceanic" and "continental" distributions (Lit-
toraria, Reid, 1986; Peasiella, Reid, 1989b), but hitherto
the phenomenon has been demonstrated only in the trop-
ics.
The biogeography of Littorina has been analysed by
Reid (1990b). These modifications to the cladogram and
distributional data on new species do not alter the earlier
conclusion that in general in the northeastern Pacific
more apomorphic species occur at higher latitudes, per-
haps as a result of climatic cooling which induced spe-
ciation. Indeed, recognition of "L. kurila" (sensu main
text of Reid, 1990a and Reid, 1990b) as conspecific with
L. sulrrotundata removes the anomaly of the latter s
northeastern Pacific distribution. However, L. naticoides
and L. aleutica are the only known example in the genus
of sister species that occur in the same ocean and show
allopatric distributions; in other ca.ses distributions show
broad overlap This could indicate that a different spe-
ciation mechanism has been involved here.
ACKNOWLEDGMENTS
The visit of DCR to the Zoological Institute, Leningrad,
was supported by the exchange program of the Ro\al
Society of London and the Academy of Sciences of the
USSR. DGR would like to thank the Director of the
Institute, Prof. O. A. Scarlato, for the opportunity to work
there. Photography of shells was b\ P R Crabb(BMNH).
LITERATURE CITED
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of Nudilittorina (Gastropoda: Prosobranchia): compara-
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Boulding, E. G. 1990. Are the opposing selection pressures
on exposed and protected shores sufficient to maintain
genetic differentiation between gastropod populations with
high intermigration rates? Hydrobioiogia 193:41-52
Carpenter, P. P. 1864. Supplementary report on the present
state of our knowledge with regard to the Mollusca of the
west coast of North .America. Report of the British .Asso-
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Ciofas, S. 1975. Sur I'extension de Littorina saxatitis (Olivi)
(Moll. Gaster.) dans le Detroit de Gibraltar. Bulletin de la
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95-99.
Golikov, A. N. and O. G. Kusakin 1978. Recent molluscs of
the littoral zone of the I'.S.S R Nauka, Leningrad, 256 p.
Hannaford Ellis, C J. 1979, Morphology of the oviparous
rough w'ink\e Littorina arcana Hannaford Ellis. 1978, with
notes on the taxonomy of the L saxatilis species-complex
(Prosobranchia: Littorinidae). Journal of Conchology 30:
43-56.
Heller, j. 1975, The taxonomy of some British Littorina spe-
cies with notes in their reproduction (Mollusca: Proso-
branchia). Zoological Journal of the Linnean Societv' 56:
131-151.
Janson, K. 1982. Phenotypic diftereritiatioii in Littorina sax-
atilis Olivi (Mollusca, Prosobranchia) in a small area on
the Swedish west coast. Journal of Molluscan Studies 48:
167-173.
Janson, K. 1983. Selection and migration in two distinct phe-
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THE NAUTILUS 105(l):16-25, 1991
Page 16
Activity Patterns and Homing of Acanthopleura gemmata
(Blainville, 1825) (Mollusca: Polyplacophora) in the
Rocky Intertidal of the Jordan Gulf of Aqaba
Neil C. Hulings'
Marine Science Station
PO Box 195
Aqaba, Jordan
ABSTRACT
Acanthopleura gemmata forages and homes irrespective of the
time of da\\ Feeding excursions begin when the water depth
of the ebbing tide over the individual is around 10 cm or less
and homing occurs prior to the level of the flooding tide reach-
ing about the same depth. The patterns of behavior differ
significantly from those of A. gemmata previously reported at
Heron Island, Australia and in Somalia. Reduced predation
pressure is considered to be one of the factors accounting for
this difference in behavior
Key words: Chiton; ecology; predation; Red Sea.
INTRODUCTION
Ferreira (1986) revised the genus Acanthopleura Guil-
ding, 1829 to include Acanthozostera Iredale and Hall,
1926. The revision also included placement of Acantho-
pleura haddoni Winckworth, 1927 and A. spiniger (Sow-
erby, 1840) into the synonymy of A. gemmata (Blain-
ville, 1825). The material examined by Ferreira (1986)
included specimens from Israel, Egypt, Djibouti, Oman,
Somalia, Heron Island, Australia as well as numerous
other localities in the Indo-Pacific region. Of the inter-
tidal chitons, A. gemmata has the widest geographic
distribution, from 32°E to 140°W and 33'N to 6°48'S
(Ferreira, 1986). The synonymy and definition of A.
gemmata proposed by Ferreira (1986) is followed in the
present paper.
Reports on the activity and homing of Acanthopleura
gemmata include those of Thorne (1967, 1968 as Acan-
thozostera gemmata) at Heron Island, Australia and
Chelazzi et at. (1983a,b, 1987) and Chelazzi and Par-
pagnoli (1987) along the coast of Somalia. Of the reports
of A. gemmata (cither as A. haddoni or A. spiniger) in
the Red Sea, including the Gulf of Aqaba (Sykes, 1907;
' Present address: Ranunkelgatan 7D, 431 32 Miilndal, Swe-
den.
Leioup, 1937, 1960, 1980; Rees & Stuckey, 1952; Safriel
& Lipkin, 1964; Eibschiitz e/ ai. 1967; Fishelson, 1971;
Mergner & Schuhmacher, 1974; Mastaller, 1978, 1979;
Pearse, 1978, 1983; Ayal & Safriel, 1980; Soliman &
Iskander, 1982; Hulings, 1986, 1987; Al-Hajj, 1987), none
contain information on foraging or other patterns of ac-
tivity, nor on homing behavior.
This paper describes the activity patterns and homing
of Acanthopleura gemmata (figure 1), the only chiton
in the rocky intertidal of Jordan. The vertical distribution
of this chiton ranges from the mid-midlittoral to the
midlittoral-infralittoral fringe contact. The northern Gulf
of Aqaba is the northernmost geographic occurrence of
A. gemmata according to Ferreira (1986). Voucher spec-
imens of A. gemmata have been deposited in the Na-
tional Museum of Natural Historv Smithsonian Institu-
tion (USNM 858104) and in the reference collection of
the Marine Science Station, Aqaba, Jordan.
STUDY AREA
The characteristics of the rocky intertidal zone along the
coast of Jordan have been described bv Hulings (1986,
1987). Briefly, the tides are mixed, having a spring range
of around 1.0 m and a neap range of 0.5 m. The diurnal
inequality of the high tides averages 4.2 cm, that of the
low tides 4.7 cm. The tide levels are often influenced by
atmospheric pressure (L'ziel, 1968; Hulings, 1989). The
tide levels also fluctuate w ith changes in sea level of up
to 1 m that occur in the northern Gulf. During the period
July through October, sea level is higher while from
December through May, it is lower; the intervening pe-
riods are transitional (Uziel, 1968; Fishelson, 1973; Hul-
ings, 1989). Although the vertical ranges and variations
in the tide are small, the prevailing near horizontal sub-
strata of the midlittoral zone are subjected to extended
submergence during the periods of higher sea le\el and
emergence during periods of lower sea level (Hulings,
1986).
The climatic conditions in the northern Gulf of Aqaba
are very significantK influenced bv the much greater
N. C. Hulings, 1991
Page 17
-•- 3
Figure 1. Acantho])lcura gemmata from the rock\ midlittnral
zone of Jordan, northern Gulf of Aqaba, (f.O x ).
desert land area surrounding the very limited marine
area. The land to sea winds are northerly 93*^' of the
time with Beauforts 3 (7 to 10 kn) and 4 (11 to 16 kn)
occurring 63% of the time. Mean air temperatures range
from 16 °C in January to 32 °C in July-August; minimum-
ma.ximum means ranges are 12 to 20 °C in January' and
26 and 37 °C in July-August (Jordan Meteorological De-
partment). Anati (1976, 1980) estimated evaporation in
the Gulf to be 4.2 m/yr. The area is further characterized
by strong solar insolation (Assaf & Kessler, 1976; Leva-
non-Spainer et al.. 1979).
The locality (figure 2) investigated in detail for the
activity of Acanthopleura gemmata is located at the
Marine Science Station (34°.59'20"E, 29°29'30"N), ap-
pro.ximately 10 km south of the city of Aqaba. The mid-
littoral zone is characterized by two types of substrata,
landward beachrock and below, a platform of near hor-
izontal calcareous fossil reef matrix. The beachrock area
is biologically recognizable b>' the presence of the giant
barnacle Tetraclita squamosa rufotincta Pilsbry, 1916,
which colonizes the surface of the beachrock. This area
is the Tetraclita zone of Safriel and Lipkin (1964) and
is considered herein as the middle portion of the mid-
littoral. The zone averages 5 m in width and consists of
discontinuous strips of beachrock, underlain by fossil reef,
parallel to the shore; between the strips are depressions
and tide pools as well as numerous and varied refuges.
The surface configuration of the fossil reef in the area
of the beachrock is ver\ irregular. In addition, the un-
derlying reef substrate is hard compared to the reef ma-
trix of the platform.
The platform of nearly horizontal, calcareous reef ma-
trix is 10 to 18 m wide, extending from the seaward edge
of the beachrock to the midlittoral-infralittoral fringe
(Safriel & Lipkin, 1964) contact and is also referred to
herein as the lower midlittoral. The surface topography
of the platform is mostly even and uniform. In the area
iS^
Figure 2. A section oi tlie stud\ area tiuruig ebbing tide. The
arrows indicate: 1. .■\pproxiniatesupraiittoral fringe-midlitorrai
contact zone; 2. The Tetraclita or mid-niidlittorai zone; 3.
lower midlittoral-infralittoral fringe contact; 4. outer edge of
the fringmg reef, See text for details.
of the midlittoral-infralittoral fringe contact, however,
the substratum is eroded, providing refuges such as over-
hangs, crevices and pits. Beyond the contact is a 35 to
85 m wide back reef lagoon of a fringing reef. The
lagoon, having a maximum depth of around 2 m de-
pending on the tide and sea level, terminates at the outer
edge of the reef. It is here that wind-generated waves
break and their energ\ is further reduced across the
lagoon. Thus the intertidal area has minimal exposure to
wave action. During high tides, however, wind-induced
choppy conditions may prevail, but the wave amplitude
is small because of the shallow depth.
.•\t the localitx' in\estigated, the mean length of Acan-
thopleura gemmata is 53.0 mm (SD = ±8.6 mm; range,
30 to 75 mm; n = 160) and the mean density is 4.4
individuals/ m- (SD = ±3.1; n = 80 m-). According to
Hulings (1986), A. gemmata becomes sexualK' mature
at a length of between 24 and 27 mm.
METHODS AND MATERIALS
The data on the activity and homing of Acanthopleura
gemmata were obtained entireK from detailed obser-
vations made during the period 14 October through 16
December 1986. Observations were made on groups of
A. gemmata within a 100 m stretch parallel to the beach
and 10 to IS ni wide. The area included the midlittoral
range of the vertical distribution of A. gemmata as well
as its occurrence in beachrock areas, on fossil reef capped
by beachrock and on the platform below the beachrock.
The shell plates and the home area or scar of Acan-
thopleura gemmata were color-coded using different
colors of enamel paint. Subsequent repainting of the
plates was often necessary and there was no detectable
negative effect of the painting. Forty-five individuals
comprising eight diflerent groups, each with three to
eight individuals, were color-coded One group was on
beachrock (six individuals), three on fossil reef capped
by beachrock (17 indi\iduals) and four (22 individuals)
Page 18
THE NAUTILUS, Vol. 105, No. 1
on the platform. During the study perioii, one specimen
was lost at the beachrock site, two from the beachroek-
fossil reef substrata and three from the platform area.
Thus 39 individuals, wliich were present during the en-
tire two month period of investigation, were used in the
analyses of activity and behavior. Various combinations
of groups and or indi\ iduals were used for the analysis
of specific t\ pes of acti\ it\ and l)ehavior.
During the October-December period, daytime ob-
servations were made on 46 days. The total number of
days included an initial two consecutive days of obser-
vations follow ed by eight weeks of five consecutive days
per week and a hnal four consecutive days. On each day,
each individual within the eight groups was observed
one or more times (all groups observed each time being
a set of observations). The number of observations on
each group averaged 5/day (range, 1 to 11 /day). Over
the 46 days, the number of sets of observations per group
ranged from 139 to 161 with the total numb)er being
5,712. Nighttime observations were made on two groups
totaling eight individuals during October (2), November
(2) and December (1). There were two sets of observa-
tions each night, a total of 80 sets or 320 observations.
Six additional groups of six to 11 individuals, a total of
53, within the area w ere checked once a day during each
five consecutive days of observations. In addition, peri-
odic observations were made during January, February,
June and August 1987 and February and June 1988.
The data recorded during each observation of each
color-coded individual included the position in relation
to the color-coded home scar. If the individual was in
the home scar, it was so noted as well as whether there
was a reversal in the orientation. If the individual was
away from the scar, the distance in cm from the home
scar and the direction was noted. Occasionally, actual
movement of an individual was observed. In most cases,
however, movement was inferred based on the position
of the individual. The depth of water over the individual
was measure in approximate cm intervals. If the indi-
vidual was partially to completely submerged dorso-
ventrally, it was designated as awash. In the absence of
water cover, the substratum was recorded as wet if water
was visible or dry if no water was visible.
The number of sets of observations made in relation
to the state of the tide during the daytime over the two
months were as follows: eight at flood tide, 39 at flooding
tide, 27 at ebb tide, 18 at ebbing tide and six at ebbing
to ebb tide. During the period, there were 19 days of
spring tide, 10 of transitional spring to neap, 10 of neap
and seven of transitional neap to spring. During night-
time observations, the tide was at flood on one occasion,
at ebb on another, ebbing on a third and flooding on two
others. During two of the nights, there were spring tides,
during one neap and during two transitional spring to
neap.
Two additional groups of Acantlwplcura gcmniata
were monitored for vertical migration with changes in
sea level from low during October (10 days) to high
during December ( 1 1 days) with November (22 days)
being transitional. One group of 12 individuals was in
the Tr/rflr/i7a -beachrock area of the midlittoral and the
other group, 11 individuals, was near the midlittoral-
infralittoral triiige contact. During the October-Decem-
ber period, a total of 91 observations during 46 days were
made on the Te^rar/i7a-beaclirock group and 24 obser-
vations during 16 days on the midlittoral-infralittoral
fringe group. Subsequent observations on each group
were conducted during January (2 days), February (1),
June (1) and August (1) 1987 and February (1), June (1),
August (1) and September (1) 1988. These observations
included four days of higher sea level and five of lower.
The direction of movement from the home scar during
foraging forays was measured in 17 color-coded Acan-
thopleura gemmata in the beachrock area and on the
platform over a period of 2 to 29 days. The range of
foraging (in cm) was measured among the same indi-
viduals over the same period. The range was considered
to be the distance from the home scar to the most distant
point within the individual grazing area, the latter iden-
tifiable by the color being lighter than the surrounding
area.
RESULTS
Activity, in terms of movement, of Acanthopleura gem-
mata from mid-October to mid-December 1986 was
found to be dependent on the depth of water over the
individual (table 1). Based on 19 color-coded individuals
in four groups on the platform and five in one group in
the beachrock area, when there was movement, it oc-
curred predominantly from when the substratum was
wet to when the water depth of the tide over the indi-
vidual was 10 cm or less irrespective of the time of day.
During the day, 88% of the instances of movement oc-
curred within the wet to 10 cm range, while 90% of
nocturnal movement occurred within this range. A sim-
ilar pattern prevailed in relation to the t\pe of tide day
and night (table 2). However, movement did not always
occur during the wet to 10 cm depth interval (tables 1,
2). Among the same 19 color-coded individuals, day and
night movement in relation to the type of tide (table 2)
was as follows: 34% of the instances of movement oc-
curred during spring tide, 38% during transitional spring
to neap, 16% during neap and 12% during transitional
neap to spring.
It is recognized that the number of observations at
night was very limited. However, additional evidence
tor nocturnal activity is derived from comparing the
position of an individual at the end of one day with that
at the beginning of the next. Among five groups of 24
color-coded individuals, a total of 318 observations re-
vealed 113 (36%) instances of positions different from
that of the last observation, usually late afternoon, on the
previous day. Among the total different positions, 34
(30%) were in the home scar at the beginning of the
second day. By comparison, 193 (94%) of the 205 in-
stances of the position being the same were in the home
scar at the beginning of the second day.
N. C. Hulings, 1991
Page 19
Table 1. Movement of Acaiitliupleuru geiuiuata in relation
to water depth during day and night (u = number of obser-
\atioiis; — = no obser\ation; A = awash).
Table 3. Home (■.s. awa\ position of Aamlhoplcuru gcmmata
in relation to water depth (n = number ol observations)
Water
Movement
No
iiovement
depth
Da\
N'ighl
Day
N'ight
(cm)
n
n
n
n
Dry
Wet
0
97
15
26
264
54
A-5
172
43
171
49
5-10
137
48
128
60
10-15
45
12
104
39
15-20
8
—
102
0
20-25
0
—
48
—
25-30
0
—
32
—
30-35
0
—
10
—
35-40
0
—
5
—
Total
459
118
890
202
The home vs. away position of one group of five in-
dividuals with respect to water deptli during the day
over 41 days is show n in table 3. In 70'^o oi the instances,
the individuals were at home over a range of conditions
from dry substratum to a water depth of 50 cm, while
85% of the away instances were during the wet to around
10 cm water level period.
The movement of Acanthoplcura gcmmata with the
ebbing tide was either rotation within the scar or the
initiation of a feeding excursion. The latter generally
occurred from when the water depth over the individual
was around 10 cm until the time the individual was
completely exposed. Following exposure, the animal ei-
ther returned to the homing scar before complete drying
of the substratum or became inactive and remained in
situ. There was no movement when the substratum was
dry (table 1). Movement of the in situ individual started
upon wetting by the flooding tide and the individual
returned to the homing scar immediately before the wa-
ter depth of the flooding tide reached around 10 cm over
Water depth
(cm)
Home
n
Away
n
Dry
Wet
A-5
5-10
10-15
15-20
20-25
25-30
30-35
35-40
50
Total
10
0
96
46
42
95
85
45
74
21
75
4
46
3
43
3
25
1
19
1
5
0
20
219
the scar. In a few cases, return occurred at a water depth
of up to 20 cm (table 1). Fecal pellets similar to those
described by Taylor and Way (1976) were released dur-
ing the period of occupation of the homing scar. The
general pattern of activity of A. gemmata with the ebb-
ing and flooding tide, based on a total of 4,445 obser-
vations of 24 individuals during the October-December
period, is summarized in table 4.
Other responses to the flooding tide among 39 chitons
based on 2,769 observations were as follows: no move-
ment at all was detected (62'^.); movement restricted to
one or more 1S0° rotations witliin the homing scar (15%);
short-term excursion, usually not exceeding 5 cm (23%).
In addition, 124 observations on 24 individuals during
25 days revealed no long excursions during subsequent
tide cycles within the same day in 41 cases (33%).
The direction of movement on long excursions from
the homing scar over a period of 2 to 29 days among 17
individuals was within an average angle of 177° (SD =
±98°; n = 179) and ranged from 45 to 360° The distance
of the excursions of the same individuals within the graz-
Table 2. Movement of Acanthupleura gcmmata ni relation to water depth and type type of tide day and night (in parentheses).
S = spring tide; SN = transitional spring to neap; N = neap; NS = transitional neap to spring; n = number of observations; — = no
observation; A = awash.
Water
Mo\
ement
No mo\
ement
depth
S
SN
N
N8
S
SN
N
NS
(cm)
n
n
n
n
n
n
n
n
Dry
O(-)
O(-)
O(-)
0(-
26(— )
O(-)
O(-)
O(-)
Wet
51(0)
27(12)
12(0)
7(—
184 (0)
59(21)
1(0)
20(— )
A-5
53(14)
58(16)
38(17)
23(—
72(19)
61(17)
21(16)
17(-)
5-10
39 (20)
60(14)
7(20)
31(-
48(14)
42(19)
17(14)
21(-)
10-15
12(4)
29(1)
0(0)
4(-
25 (25)
62 (28)
10(0)
7(-)
15-20
3(0)
2(0)
2(0)
2(-
21 (29)
26(0)
23(0)
32(— )
20-25
O(-)
O(-)
O(-)
0(-
2(-)
13(-)
14(-)
19(-)
25-30
O(-)
O(-)
O(-)
0(-
O(-)
5(-)
7(-)
20(— )
30-35
O(-)
0(— )
O(-)
0(-
l(-)
l(-)
l(-)
7( — )
35-40
O(-)
158(38)
O(-)
176(43)
O(-)
58 (37)
0(-
l(-)
380(87)
l(-)
270(85)
2(-)
l(-)
Total
67 (—
96 (30)
144 (-)
Page 20
THE NAUTILUS, Vol. 105, No. 1
ing area averaged 21 cm (SD = ±15 cm; n = 179). On
the near horizontal reef matrix, the average angle of the
excursions was 149° (SD = ±106°; n = 100) and the
mean distance 21 cm (SD = ±13 cm; n = 100) while
on beachrock and associated substrata, 233° (SD = ±72°;
n = 79) and 23 cm (SD = ±17 cm; n = 79).
Observations on 12 Acanthopleura gemmaia in the
rc/rar/ifa -beachrock area and 11 near the mitllittoral-
infralittoral fringe contact revealed no vertical migration
from the period of lower sea level in October through
the transitional period during November to higher sea
level in December. The absence of vertical migration
was subsequentK confirmed by four observations during
high sea level and five during low sea level in 1987 and
1988. This is contrary to the report of vertical migration
of A. gemmaia in Hulings (19S7) which was based on a
more limited number of observations.
The investigation of the homing behavior of Acan-
thopleura gemmata during mid-October to mid-Decem-
Table 4. Activity of Acanthopleura gemmata in relation to
the tide (n = number of observations).
Activitv II '>i
ber revealed that 79% (SD
:205
= 1,561) of 39
color-coded individuals in eight groups exhibited definite
homing behavior, i.e., returning to the same color-coded
homing scar following foraging excursions. The obser-
vations included day and night as well as during spring,
transitional and neap tides. The homing behavior and
foraging excursions occurred irrespective of the time of
day and the type of tide. The remaining 21% (SD =
±20%; n = 466) occupied new sites for a variable period
of one to several days. Observations on other groups
totaling 53 individuals as well as periodic observations
through June 1988 essentially confirmed these findings.
Within groups in which the individuals were clumped
or in close proximity, exchange of home scars was found
to be common. The average number of homes occupied
by each of 39 individuals over the two month period was
3.2 (SD = ±1.5; n = 2,027). The period of consecutive
occupation of a home other than the original among 16
individuals (n = 242) varied from 1 to 19 days and
averaged 5.8 days (SD = ±5.0 days). Among 39 indi-
viduals, four occupied the original home scar for the
entire two month period of observation while seven did
not return to the original or new home scar. Among the
remaining individuals, there were 20 cases in which there
was immediate or eventual return to the original home
scar and 49 in which the individual moved to one or
more home scars and returned immediately or eventu-
ally.
Within one group of five individuals, there was only
one case of reciprocal swapping on a daily basis over the
two month period (n = 760). In six cases, there was initial
reciprosity followed by suKsequent continued occupation
of up to five da\s. Within the same group and over the
same time span, the total number of sites occupied was
47. There were 27 reoccupations of a site following oc-
cupation by another individual.
The most recognizable homing scar of Acanthoph'ura
gemmata in the rockv intertidal of Jordan is a distinct
depression that is concave in cross and longitudinal sec-
tions. The scar is about 1 mm deep with the deepest area
\o movement from home during
ebbing tide nor during exposure.
\lo\emenl from home with the ebb-
ing tide and return home with the
subsequent flooding tide.
Movement from home with the ebb-
ing tide and return home prior to
the flooding tide.
Movement of those previously away
at ebb tide to home with the flood-
ing tide
1.709
1,584
576
.5,(1
38
36
13
13
coinciding with that region occupied by the foot. The
surface ot the scar is smooth compared to the surface of
the surrounding substratum and has a lighter color than
the surrounding substratum, v\ hich is covered with epi-
growth. Though abandoned scars become overgrown,
they remain recognizable. Such readily recognizable
homing scars are found on the nearly horizontal, rela-
tively soft, homogenous, fine-grained calcium carbonate
fossil reef matrix, a dominant rocky intertidal substratum
along the coast of Jordan. The scars are also found on
the sides of refuges such as depressions, crevices and small
caves. The scars of A. gemmaia lack the algae occurring
in the homing scars of the patellid limpet Cellana radiata
(Born, 1778) and the pulmonale Siphonaria laciniosa
(Linnaeus, 1758) reported by Hulings (1985).
Other less recognizable scars may be identified on the
basis of substrate color. Apart from this difference, the
surface configuration of the homing area is not noticably
different from that of the surrounding substratum. These
scars are also found on calcium carbonate substrata, one
type of which underlies beachrock and is harder than
the matrix noted above. Another type of substratum with
such scars includes fossil coral heads. Recognizable hom-
ing scars have not been detected on substrata such as
granitic and similar rock and beachrock although homing
on these substrata has been observed. Endean et at. (1956)
noted the absence of excavations by Aeanthoph^itra gem-
mata in hard rock on the coast of Australia.
The orientation of both homing scar and chiton varies
irrespective of the substratum. The scar may be near to
horizontal on such substrata or on the sides of refuges.
In the refuges, the anterior-posterior axis of the chiton
may be parallel to the horizontal or inclined to vertical
with the anterior end up. Semi-quantitative data (Hul-
ings, in preparation) indicates that the density of Acan-
thopleura gemmata in areas with refuges is 4.1 m- (SD
= ±3.2/m-; n = 183) while that in areas without refuges
is 5.5/m- (SD = ±5.4/nr; n = 79).
Among 37 chitons and based on 2,027 observations,
the mean percent homing of 22 indiv iduals on relatively
.soft, nearly horizontal reef matrix v\ here the majority of
the readily recognizable homing scars occur was 85%
(SD = ±14%). B\ comparison, the mean homing of 15
N. C. Hulings. 1991
Page 21
individuals on hard substrata such as beachrock and the
underlying substrata was 69% (SD = ±24%). The dif-
ference between the means is statistically significant at
P = 0.01 (n — 2 = 35). While the mean number of homes
occupied by the individuals on soft and hard substrata
was almost the same, 3.4 (SD = ±1.4) vs. 3.1 (SD =
±1.6), there were eight instances among the individuals
on soft substrata of return to the original home vs. 12
among those on hard substrata. In the case of movement
to a different home and return, there were 34 instances
on soft substrata vs. 10 on hard.
DISCUSSION
There are limitations that must be imposed on the in-
terpretation of the data obtained during the investigation
of the acti\it\ patterns and homing of Acanthopleura
gemmala. First, the data are based entireK on obser-
vations; no e.xperimental manipulations were conducted
although the value of such is recognized. Second, statis-
tical treatment is limited, of a general nature and is used
for indicative purposes only. FinalK , the data appK' onh-
to adult indi\ iduals. Other limitations are indicated be-
low
The homing behavior of Acanthopleura gemmata in
the northern Gulf of .\qaba is similar to that reported
for this species by Thorne (1967, 1968) at Heron Island,
Australia, and by Chelazzi et al. ( 1983a, b, 1987) and
Chelazzi and Parpagnoli (1987) on the coast of Somalia.
The homing of 79% of the individuals recorded herein
is the same as reported by Thorne (1967) while Chelazzi
et al. (1983b:17) reported a 91%' return to the home of
the "previous owner." In addition, the same percentage
was found for individuals occupying new homes at Heron
Island and on the coast of Jordan. Chelazzi et al. (1983a)
found periodic movement to a new home a common
behavioral pattern as did Throne (1967) and found here-
in. In terms of reciprocal swapping of homes, the low-
incidence found herein is in agreement with that found
by Thorne (1967). The 57% reoccupation of a home
following occupation b}' another individual is higher than
the 42% reported by Chelazzi et al. (1983b).
The orientation in the homing scar, including 180°
rotation, is essentialK the same in the three geographic
areas. There is also considerable similarit\ in the mor-
phology and appearance of the homing scar. The oc-
cupation of scars in exposed (platform) sites as well as
in refuges (pits, crevices, overhangs) is a common be-
havior (Thorne, 1967; Chelazzi et al.. 1983b).
Chelazzi et al. (1983b) reported aggressive behavior
between a chiton returning to its home scar and an in-
truder occupying the same homing scar. The returning
chiton attempts to oust the intruder, but, if unsuccessful,
the owner either mo\es elsew here or the two individuals
occupy the same scar. In the present study, aggressive
behavior was not observed, but the co-occupation of a
single scar by two individuals, side by side, was observed
on several occasions.
Onl\ a few periods of observations during one day
included a combination of a long enough time span,
suitable water level conditions and movement by Acan-
thopleura gemmata to indicate the return to the home
scar following grazing. During such a combination, 43
observations on eight indi\ iduals suggests return by the
same route as that of the excursion. This is consistent
with the reports of Thorne (1967) and Chelazzi et al.
(1983b, 1987).
Thorne (1967) suggested chemosensitivit\ and topo-
graphic memory as potential mechanisms whereby
Acanthopleura gemmata returns to the homing scar.
This author (Thorne, 1968) later added celestial navi-
gation, sight, random movements, olfactory clues, to-
pographic memory and detection of previous trails as
possible mechanisms. For a \ariety of reasons, he ruled
out the first three and, after subjecting the last three to
experimental treatment, considered only topographic
memory and detection of trails as possible homing mech-
anisms in A. g£>mma?a. Eibschiitz p< a/. (1967) discounted
the magnetic properties of the radula of A. gemmata at
Eilat as a factor in navigation because of the low level
of residual magnetism.
Of note is the occurrence of homing even when the
scar and the surrounding area was completely covered
with sand during periods of sedimentation, or with shell
debris follow ing a storm. The frequenc\ of homing by
individuals with well-de\ eloped scars was significantK'
higher, suggesting that a chemical trail may be involved
in the homing of A. gemmata. as indicated by Chelazzi
and Parpagnoli (1987). However, reciprocal swapping of
scars and the reoccupation of sites following occupation
by one or more indi\iduals suggests that there is not a
trail specific for each individual but one common to all
individuals of the species.
The mean length of the feeding excursions of Acan-
thopleura gemmata in this stud\' was 21 cm. By com-
parison, Chelazzi et al. (1983b) reported 30 cm and
Thorne (1967, 1968) 47 cm. Thorne (1967, 1968) found
that not all individuals forage every night and Chelazzi
et al. (1983b:16) noted that nocturnal activity depended
"on the position in the spring-neap cycle." In the present
study, not all individuals w ere found to be active during
the conditions under which foraging occurs (tables 1, 4)
nor during consecutive tide cycles.
Chelazzi et al. (1983a) found the feeding excursions
of Acanthopleura gemmata to be the longest between
neap and spring tides and minimal around neap tides.
In the present study, the length of feeding excursions in
relation to the t\ pe of tide was not determined. However,
the number of instances of movement in relation to the
type of tide was highest during transitional spring to
neap tides followed by spring, neap and transitional neap
to spring tides (table 2). Based on the assumption that
mo\ements during ebbing conditions represent foraging
excursions, an analysis of one group of five individuals
on 26 days (n = 157) revealed 41% of the instances of
movement occurred during ebbing tides. Of this per-
centage, 43% of the instances occurred during transi-
Page 22
THE NAUTILUS, Vol. 105, No. 1
tional spring to neap tides, 32% during spring, 13% dur-
ing neap and 5% during transitional neap to spring.
Movement of Acarithopleura gemtnata in the north-
ern Cult of .\qaba is more directl) related to \\ ater depth
over the indi\iduai than to the t\pe of tide. During
periods of higher sea level or portions of the neap tide
cycle, the water level over an individual may not drop
to 10 cm or less, especially in the lower midlittoral zone.
Thus feeding excursions may not occur during one or
more consecutive days. The fewer instances of movement
during neap and neap to spring tides over that of spring
and spring to neap (table 2) may also be indicative of
reduced activit\'.
Many Acanthopleura gemmata in the northern Gulf
have a well-defined grazing area, recognizable by dif-
ferences in color from that of the surrounding area. This
area is used repeatedly (Chelazzi ei al., 1983a, b, 1987),
but appears to be exclusive, unlike that reported by Che-
lazzi et al. {1983b). The feeding areas are more dispersed
among groups of individuals than the homing scars as
also found by Chelazzi et al. (1983a). Grazing appears
to be most common at the outer limits of the area, as
also reported by Thorne (1967).
On the coast of Jordan, the range of the feeding ex-
cursions of Acanthopleura gemmata was found to differ
with the surface topology of the substratum. On the more
regular surface of the near horizontal platform of cal-
careous reef matrix, the range averaged 149° while on
the very irregular surface of the beachrock and under-
lying substrata, the average range was 233°. The differ-
ence in the range may reflect the amount of colonization,
distribution and abundance of algae as well as recolon-
ization following grazing on the two different substrata.
It would appear that the conditions for recolonization
are more favorable on the soft, porous reef matrix due
to the nature of the substratum and decreased tidal ex-
posure than on the more exposed hard, impervious
beachrock and associated substrata. As a result, more
algae are more evenly distributed more of the time on
the platform. Accordingly, a greater range of foraging
would be necessary on the beachrock and associated sub-
strata in order to obtain the necessary food requirements.
The mean length of A. gemmata along an exposure gra-
dient within the vertical distribution may also reflect food
availability as well as a response to exposure (Hulings,
in preparation). In the mid-midlittoral beachrock area,
the most tidally exposed within the range, the mean
length was found to be 47.0 mm (SD = ±7.6 mm; n =
60) while in the area of the midlittoral-infralittoral fringe
contact of the platform, the most benign, the mean was
59.8 mm (SD = ±9.8 mm; n = 50). On the platform
just below the beachrock, an area intermediate in ex-
posure, the chitons had a mean length of 53.3 mm (SD
= ±6.2 mm; n = 50). The difference between the means
is statistically significant at P = 0.001.
No aggressive behavior within the feeding area of
Acanthopleura gemmata has been ob-served. This is con-
sistent with the report of Chelazzi et al. (1983b). Among
the chitons on the Jordanian coast, there was no pre-
vailing up- or downslope nor lateral direction of the
feeding excursions. .\s a result of orientation and length
excursion analyses of A. gemmata and the sympatric A.
hrevLspinosa (Sowerby. 1840), Chelazzi et al. (1983a) and
(Chelazzi and Parpagnoli (1987) found the prevailing di-
rection of movement during the feeding excursions of
A. gemmata to be downw ard and that of A. brevispino.sa
upward. They considered this behavior to be an example
of minimizing food competition and maximizing the uti-
lization of algae in the midlittoral.
There is a possible example of minimizing competition
in the rocky intertidal of Jordan. Based on density and
presence-absence data, the correlation between the spa-
tial distribution of the limpet Cellana radiata and Acan-
thopleura gemmata was found to be negative (Hulings,
in preparation). The limpet is considered to be the major
grazing competitor of the chiton, especialK in the lower
midlittoral, where the two are the numericalh dominant
intertidal grazers. However, the conditions under which
each forages differs. Cellana radiata forages only when
submerged, becoming active soon after submergence by
the flooding tide to just before or just after emergence
(Hulings, 1985). Acanthopleura gemmata forages with
the ebbing tide.
The timing and the conditions under which feeding
excursions of Acanthopleura gemmata occur at Heron
Island, Australia and on the coast of Somalia differ very
significantly from those in the northern Gulf of Aqaba.
Thorne (1967, 1968), Chelazzi et al. (1983a,b, 1987) and
Chelazzi and Parpagnoli (1987) found A. gemmata to
be active only during nocturnal low tides and not during
the day, whether exposed by the tides or not. Chelazzi
et al. (1983b) reported that A. gemmata remains in the
homing scar during the following high tide and diurnal
low tide.
By comparison, Acanthopleura gemmata in the north-
ern Gulf has been found to be active during day and
night. Foraging excursions commence when the water
depth of the ebbing tide over the individual is around
10 cm or less. Return to the homing scar occurs prior to
or with the flooding tide; in the latter case, before the
water level is around 10 cm over the individual (table
4).
The mechanism(s) triggering the initiation of feeding
excursions and homing in Acanthopleura gemmata in
the northern Gulf is not known. Other investigators have
suggested various mechanisms, including a biological
clock (Thorne, 1967) and the release of chemical infor-
mation during high tide (Chelazzi & Parpagnoli, 1987).
Though not investigated, the presence of photoreceptors
such as aesthetes and ocelli in the shell plates and the
possibilitv of the girdle being photosensitive (Bo\le, 1977)
does not appear to be involved in the triggering of move-
ment in A. gemmata as it occurred da\ and night. Of
particular interest is the finding of Moulton (1962) of the
initiation of movement aw ay from clusters of the cerith-
iid Chjpeomorus monilijcrum Kiener, 1841 at Heron
Island when the water depth reaches 10 cm (the same
species occurs in the rock\ intertidal of Jordan according
N. C. Hillings, 1991
Page 23
to Hillings, 1989), Moulton (1962) suggested a hydrostatic
mechanism and, behaviorally, positive barokinesis and
thigmotaxis as being involved in clustering and dispersion
of the cerithiid. Based on laboratory experiments, Rohde
and Sandland (1975) discounted a hydrostatic mecha-
nism as the onl) factor in dispersion and clustering of
C. moniliferum.
It is suggested that a difference in predation pressure
on Acanthoplciiro gemmata between the Heron Island-
Somalia region and the northern Gulf of Aqaba may
account, in part, for the nocturnal behavior in the former
region vs. the diurnal-nocturnal behavior in the latter
area. Both Thorne (1967) and Chelazzi et al. (1983a,b)
considered homing in A. gemmata to be of significance
in the reduction of and protection against predation.
Among the predators listed by Thorne (1967, 1968)
are the grapsoid Grapsus strigosus (Herbst, 1799) which
is synon\mous with G. albolineatiis Lamarck, 1818 ac-
cording to Banerjee (1960); the xanthids Eriphia leavi-
mana Latrielle, 1817 and Leptodius exaratus (H. Milne
Edwards, 1834); the anomuran Clibanarius virescens
(Krauss, 1834); birds including herons, waders and gulls
and elasmobranchs, Chelazzi el al. (1983b) reported the
xanthid crabs Eriphia smilhi McLear\, 1838 and Ozius
guttatus H. Milne Edwards, 1834 as less important pred-
ators and the tetradontid fish Arothron immaculatiis
(Block and Schneider, 1801) as the most important pred-
ator (also see Chelazzi et al., 1987; Chelazzi & Parpagnoli
1987).
In the northern Gulf of Aqaba, the grapsoid Grapsus
granulosus H. Milne Edwards, 1853 is a common mid-
midlittoral slab occupant while G. alholineatus most
commonly inhabits boulders (Hulings, 1989). Of the above
xanthid crabs, onlv Leptodius exaratus has been record-
ed in the Jordanian Gulf (D. Guinot, personal commu-
nication). Other xanthids including Lijhia leptochilis
(Zehntner, 1894) and Polijdectus cupulifcr (Latrielle,
1825) have been found in the rocky "eulittoral' b\ Ma-
staller (1979). The anomuran Clibanarius signatus Hel-
ler, 1861 is a mid-midlittoral species on slab along the
coast of Jordan (Hulings, 1989).
Among the fishes, Arothron immaculatus has not been
reported from the Gulf of Aqaba (Dor, 1984). However,
other tetradontids have been reported in the Gulf at Eilat,
including Arothron hispidus (Linnaeus, 1758) by Ben-
Tuvia and Steinitz (1952) and Tortonese (1968) and a
questionable Tetradon aff. diadcmata Riippell, 1829 by
Steinitz and Ben-Tuvia (1955). There is. however, no
information on the habitat occurrences of these tetra-
dontids. Among the species of fish iinading the rocky
intertidal during high tide according to Fishelson (1977),
potential predators on Acanthopteura gemmata include
the labrid Cheilio inermis (Forsskal, 1775) and the ba-
listid Sufflan}en albicaudatus (Riippell, 1829) (R. Or-
mond, personal communication). As for marine birds,
there is not a resident population of species in the north-
ern Gulf. However, the area is on the migratory route
of gulls, herons and waders (Safriel, 1968).
It is inferred that the predation pressure on Acantho-
pleura gemmata in the northern Gulf of Aqaba is con-
siderably less than at Heron Island and in Somalia. The
absence of a resident marine bird population as well as
rare occurrences during migratory periods indicates the
absence of a diurnal predator that would feed while A.
gemmata is exposed and active at low tide. The stomach
contents of the grapsoids Grapsus albolineatus and G.
granulosus include macerated fragments of algae, hy-
droids and detrital particles but no chitons (Hulings, un-
published data). In addition, the structure of the chelae
of the two grapsoids (Schiifer, 1954) is not suitable for
dislodging or preying upon A. gemmata. The hermit
crab Clibanarius signatus in the rock> intertidal of Jor-
dan is small (maximum length around 2 cm) as are the
xanthid crabs and are thus considered to be of minor, if
any, importance as predators. Evidence lor fishes as pred-
ators is lacking, including the absence of "scars" on the
plates of A. gemmata similar to those caused by Arothron
immaculatus in Somalia as reported bv Chelazzi et al.
(1983b).
The homing scars of Acanthopleura gemmata along
the coast of Jordan are only a mm or so deep as opposed
to those on the coast of Somalia being 1 to 10 cm deep
(Chelazzi et al., 1983b). The shallow scars may, in turn,
reflect the absence of one or more significant predators
during high tide when compared to the situation in So-
malia. In the latter case, Chelazzi et al. (1983b) consid-
ered that the homing behavior of A. gemmata protects
it from the most important predator, Arothron immac-
ulatus, during high tide when predation pressure is the
highest.
It is concluded, therefore, that the diurnal-nocturnal
pattern of behavior in Acanthopleura gemmata in the
northern Gulf of Aqaba is a result of the reduced ter-
restrial and submarine predation pressure.
That periodicity of acti\ity can change with predation
pressure has been demonstrated b\ Fricke (1974), who
found that locomotor and feeding activity of the sea
urchin Diadema setosum (Leske, 1778) in the northern
Gulf of Aqaba was nocturnal in the presence of predatory
fishes. In another area, where the predators w ere absent,
the urchins were active irrespective of the time of day.
Fricke (1974) hypothesized the evolution of the day-
night activity as an adaptation in a habitat free of pred-
ators.
The timing of the acti\it\ patterns in relation to water
depth over Acanthopleura gemnuita in the northern Gulf
does, however, suggest a response to predation. The be-
ginning of the feeding excursions at around the 10 cm
water depth of the ebbing tide, and homing when the
depth of the flooding tide reaches 10 cm, as well as the
lack of movement during periods when water depth ex-
ceeds 10 cm (table 1) indicates an adaptation whereby
movement is minimal at times when it might attract
submarine predators. In most of the cases where the
water depth over the individual exceeded 10 cm, the
chiton was in the home scar (table 3).
By occupying a concave scar on the exposed, soft,
horizontal slab, the chiton blends in with the surface
Page 24
THE NAUTILUS, Vol. 105, No. 1
topograpliy and coloration of the surrounding .substra-
tum. Additional caniouOage results from the spreading
of the girdle around the internal periphery of the scar.
The latter contrasts to the form of the bod\ during ex-
posure at the low tide. During exposure, the body is raised
and arched longitudinalK- as a result of what appears to
be contraction of the girdle. Chitons homing in the hard
substrata of the beachrock areas most commonly occur
under overhangs or in crevices or caves rather than on
the surface of the substrata.
The response to submarine predation among other
intertidal homing species in the northern Gulf, Cellana
radiata and Sipbonaria laciniosa. is different. Both for-
age and home only when submerged and the activity
may continue through maximum flood tide. However,
C. radiata is active day and night whereas S. laciniosa
is nocturnal and neither is active when exposed during
low tide (Hulings, 1985). These patterns may also be a
response to predation pressure.
In addition to protection against predation, homing in
Acanthopleiira gemmata has been attributed to protec-
tion against environmental stresses (Chelazzi et al.,
1983a, b, 1987). Certainly, the nocturnal foraging and
homing of A. gemmata at Heron Island (Thorne, 1967,
1968) and on the coast of Somalia (Chelazzi et al., 1983a, b,
1987; Chelazzi and Parpagnoli, 1987) reduces exposure
to environmental extremes. However, in the case of the
rocky intertidal in the northern Gulf of Aqaba, which is
subjected to severe climatic and environmental extremes,
the foraging and homing activity of A. gemmata is tide-
dependent and independent of the time of day or year.
And while other intertidal species migrate vertically with
changes in sea level in the northern Gulf, a behavior
considered, in part, to be an avoidance of environmental
extremes (Hulings, 1987), A. gemmata does not. As a
result, the chiton is active during periods of environ-
mental extremes, including the combined warmer-lower
sea level period, as well as during benign periods. As
noted above, homing following feeding excursions dur-
ing periods of maximum exposure to climatic extremes
does not always occur. Thus the homing of A. gemmata
in the northern Gulf as a response to protection against
environmental stresses appears to be of less significance
than elsewhere.
It is recognized that differences in the behavior pat-
terns of Acanthopleura gemmata between the northern
Gulf of Aqaba and Somalia-Heron Island may be due to
differences in physical conditions. The tides are of the
same type in all three areas, although the range is greater
than in Aqaba. Other conditions, especially the geomor-
phology, appears to be similar, based on information
given in Endean et al. (1956), Vannini et al. (1977) and
Chelazzi and Varmini (1980). Geographic differences in
activity- patterns and behavior among the same rocky
intertidal species occurring in the northern Gulf of Aqaba
and in other geographic areas outside the Red Sea, in-
cluding Heron Island, have been noted by Hulings (1989).
Differences exist even within the Red Sea. The repro-
ductive periodicity of Acanthopleura gemmata in the
northern (Julf of Aqaba (Hulings, 1986) is similar to that
in the Gulf of Suez (Pearse, 1978; Soliman & Iskander,
1982), while both differ from that in the Red Sea proper
(Pearse, 1978).
ACKNOWLEDGEMENTS
The author wishes to thank Mr Burhaii Gharaibeh,
Huw wara, Jordan and Mr. Percy Linder and Mr. Jonne
Lysell, Kallered, Sweden for assistance. The constructive
criticism of two anonymous reviewers is greatfully ac-
knowledged.
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THE NAUTILUS 105(l):26-37, 1991
Page 26
The Southeastern BraziUan Muricidae Collected by
RV Marion-Dusfresne in 1987, with the
Description of Three New Species
Roland Ilouart
Research Associate Iiistitut Royal
B-3400 Latiden (Ezeinaal),
BELGIUM
ABSTRACT
Twenty-five species of Muricidae are recorded from south-
eastern Brazil, including eight species of Muricinae, nine of
Muricopsinae, one of Ergalataxinae, one of Ocenebrinae, four
of Trophoninae, and two of Typhinae. A short discussion is
gi\en for each species. Three new species are described: Der-
mornurex leali n.sp. differs from related taxa by the presence
of 5 varices on its last teleoconch whorl and by its different
intritacalx; Favartia varimtitabilis n.sp. is compared with F.
alveata, from which it differs by the erratic placement of the
varices and by its multispiral protoconch; Trophon mucrone
n.sp. differs from T. lacuneUiis. T. dabneiji. and T. limicola
by its spiral sculpture, rounded teleoconch whorls, paucispiral
protoconch, and number of axial lamellae. Two species remain
unidentified and are listed as Muricopsis (?Murexsul) sp. and
Favarlia (Favartia) sp. Drupa didyma Schwengel, 1943 is syn-
onymized with TrachijpoUia turricula (von Maltzan, 1884).
Kcij words: Muricidae; Brazil; Dermomurcx: Favartia. Tro-
phon; new species.
INTRODUCTION
The material examined was dredged or trawled off the
southeastern coast of Brazil during the Expedition MD55
carried out by the R.V. Marion-Dufresne in May 1987,
as a joint project of Museum National d'Histoire Natu-
relle (MNHN), Paris, and Universidade Santa Ursula, Rio
de Janeiro. Drs. A. Guiile and F. Roumos were cruise
leaders. Drs. P. Bouchet, J. Lea! and B. Metivier sorted
the gastropod material on board. Muricidae were col-
lected from 34 to 1,575 m, mostly along a series of sea-
mounts running perpendicular to the Brazilian coast,
between 18°50' and 23°47'S. Most of species collected
are already known to occur off the Brazilian coast; Chi-
coreus (Siratus) consuelae (Verrill, 1950), Poirieria (Pa-
zinotus) stimpsonii (Dall, 1889), Pygrnaepterys ger-
mainae Yokes and D'Attilio, 1980, and Trophon vrrrillii
Bush, 1893, however, represent new Brazilian records.
A report on the deep-water Volutidae collected during
the same cruise has been already published (Leal &
Bouchet, 1989).
For references to original descriptions not indicated
here, the reader is referred to Radwin and D'Attilio
(1976).
The number of dead collected specimens is followed
by "dd"; the number of live collected specimens is iden-
tified by "Iv ".
The following institutional abbreviations are used:
MCZ, Museum of Comparative Zoology, Cambridge;
MNHN, Museum d'Histoire Naturelle, Paris; MORG,
Museu Oceanografico de Rio Grande; USNM, National
Museum of Natural History, Smithsonian Institution.
Prefixes for station data denote collecting equipment:
CB, Blake trawl; CP, beam trawl; DC, dredge.
SYSTEMATICS
Family MURICIDAE Rafinesque, 1815
Subfamily MURICINAE Rafinesque, 1815
Chicoreus (Siratus) coltrorum Yokes, 1990
Type locality: Ilha de Itaparica, Bahia, Brazil.
Chicoreus (Siratus) coltrorum \'okes, 1990:127, figs. 7-13.
(figure 3)
Material examined: Southeastern Brazil, stn. DC26, Yi-
toria Bank, 20°21'S, 36°59'W, 97 m (2 dd); stn. DC27,
Montague Bank, 20°26'S, 36°42'W, 81 m (1 dd); stn.
DC40, Davis Bank, 20°40'S, 34°41\V, 60 m (5 dd); stn.
DC43, Dogaressa Bank, 20°51'S, 33°45'W, 63 m (2 dd);
stn. DC47, Columbia Bank, 20°42'S, 32°13'W, 94-105 m
(5 dd); stn. DC48, Columbia Bank, 20°44'S, 32°08"V\',
250-300 m (1 dd); stn. DC50, Coluinbia Bank, 20°44'S,
31°50'W, 95-120 m (3 dd); stn. DC82, 18°56'S, 37°52'W,
85-105 m (1 dd).
A recently de,scribed and often misidentified species,
C. coltrorum is superficially similar to C. consuelae (Ver-
rill, 1950) and nia\ be distinguished mainK b\ the dif-
ferent protoconch (figures 3, 4). The protoconch of C.
coltrorum has l'/2 rounded whorls while C. consuelae
has a conical protoconch consisting of 2Vi whorls; other
differences are minimal. Onl\' empty shells were col-
lected. Rios (1985:82) reports this species from Brazil (as
R. Houart, 1991
Page 27
Stratus consuelae), from Atol das Rocas, Fernando de
Noronha; Rio Grande do Norte to Espirito Santo.
Chicoreus (Siratiis) consuelae (Verril, 1950)
(figure 4)
Material examined: Southeastern Brazil, stn. DC52,
Martin Vaz Island, 20°29'S, 28°51'W, 64-80 m (3 dd);
stn. DC59, Trindade Island, 20°30'S, 29°19'W, 52-60 m
(1 dd).
Similar to C. coltrorum (see above), the species is re-
ported from off the coast of Te.xas to Curasao and in the
Lesser Antilles (Yokes, 1990:129).
Chicoreus (Siratus) tenuivaricosus
(Dautzenberg, 1927)
(figures 18, 19)
Material examined: Southeastern Brazil, stn. CB90,
19°34'S, 39°34'W, 34 m (33dd) (6 Iv); stn. CB92, 19°34'S,
38°55'W, 340-360 m (1 dd) (1 Iv).
Many specimens, mostly juveniles, were collected from
station CB90 in dead and live condition. Two specimens,
of which one was alive, were taken from station CB92,
in 340-360 m; this record is probabK based on specimens
that had remained entangled in the net of a previous
haul (stn. CB90), and this bathymetrical range requires
confirmation.
The radula is muricine, with a lateral sickle-shaped
tooth, and a central tooth with 3 major and 2 minor cusps
(figures 18-19).
Aspella morchi Radwin and D'Attilio, 1976
(figures 9, 10, 29)
Material examined: Southeastern Brazil, stn. DC22, Vi-
toria Bank, 20°32'S, 38°11'W, 52 m (4 Iv); stn. DC40,
Davis Bank, 20°40'S, 34°41'W, 60 m (1 Iv); stn. DC41,
Davis Bank, 20°39'S, 34°43'W, 58-70 m (2 Iv).
The paucispiral protoconch (figures 9, 10) had not
previouslv been figured. The cancellate intritacal.x is well
illustrated by Radwin and D'Attilio (1976:223, fig. 167).
A specimen with 6 teleoconch whorls, and broken pro-
toconch, (MNHN, station DC40) (figure 29) is 10 mm
long, while the maximum length given by Radwin and
D'Attilio is of 6.6 mm. PreviousK known from Natal Bay
(type locality), Fortaleza (Cerea) (Radwin & D'Attilio,
1976) and Kapmoan (Bahia) (coll. R. Houart).
Aspella senex Dall, 1903
Material examined: Southern Brazil, stn. DC22, Vitoria
Bank, 20°32'S, 38°11"W, 52 m (1 dd).
The geographical range reported by Radwin and
D'Attilio (1976:25) includes North Carolina and the Gulf
of Mexico. Rios (1985:87) reported this species from the
-ggSB
Figures 1, 2. Intritacalx on last teieoconcli whorl (scale bars:
0 5 inni) 1. Dcrmomurex (Trialalclla) oxum Petuch. 2. Der-
iiKimiirex (TriahitcUa) Icali n.sp.
Abrolhos Islands. The present record extends the geo-
graphical range further southward.
Dermomurex (Trialatella) oxum Petuch, 1979
(figures 1, 26)
Dcrmomurex (Trialatella) oxum Petuch, 1979:517, fig, IE, F.
Type locality: 2 km east of Santa Barbara Island, Abrol-
hos Archipelago, Bahia State, Brazil, 17°57'S, 38°41"W,
in 25 m.
Material examined: Southeastern Brazil, stn. DC22, Vi-
toria Bank, 20°32'S, 38°11'W, 52 m (1 dd); stn. DC40,
Davis Bank, 20°40'S, 34°41'W, 60 m (2 Iv); stn. DC82,
18°56'S, 37°52'W, 8.5-105 m (1 Iv).
The largest shell is 15.5 mm long (figure 26), while
Petuch mentions 13 mm for his largest specimen (ho-
lotype).
Dermomurex (Trialatella) leali n.sp.
(figures 2, 5, 27, 28)
Type material: Holotype MORG 26457, 9.5 x 5.6 mm,
southeastern Brazil, stn. DC22, Vitoria Bank, 20°32'S,
38°11"W, .52 m (dd).
Description: Shell 9.5 mm in length, roundly-fusiform.
Spire moderately high, with 1.5 protoconch whorls and
4 convex teleoconch whorls (subadult). Suture deeply
impressed. Protoconch whorls rounded. Axial ornamen-
tation of first teleoconch whorl consisting of 5 ribs, second
whorl of 6 ribs, third whorl of 7 ribs. Last teleoconch
whorl with 4 weak, lightly rounded varices. No other
Page 28
THE NAUTILUS, Vol. 105, No. 1
y
Figures 3-17. Proloconchs (scale bars: 0.5 mm). 3. Chiroreus {Siratus) coUrorum Yokes, stn. DC26. 4. Chtcorcus (Siratus)
conmelae{\\-u\\\). stn DC52. 5. nermomurex {Trialatella)leali n.sp , Ik.IoIn pe MOKG 26457, stn D(:22 6. 7. Poirieria (Pa^"'"'"s)
stimpwnii (Dall), stn. DC82, 8. Muricopsis {?Murexsul) sp., stn. DC35 9, 10. Aspella morchi Radwiii & D .\ttilu., stn L)^"!"; ' '•
Muricopsis {Risomurex) necocheana (Pilsbrv), stn. DC22. 12. Favartia (CMrilnella) varimutaMis n.sp , holotspe MORG 26458,
sin. DC;i5. 13. Favartia {Favartia) sp , stn. 'dC35. 14. Trophon aculeatus Watson, stn. CB104. 15, 16. Trophon mucrone n.sp.,
paratype MORG 26459, stn. IX:70 17. Siphonochelus (Siphonochchis) riosi (Bertsch & D'Attilio), stn, CB104.
R. Hoiiart, 1991
Page 29
21
23
^ 25
Figures 18-25. Radulae (scale bars figures 20, 22, 24: 20 Mm; figures 21, 23, 25: 10 Mm; figure 18: 100 Mm; figure 19: 50 Mm),
18, 19. Chicoretis iSirattis) tcnuivaricosus (Dautzenberg), stn. CP90, 20, 21. Favartia (CarilneUa) varimutabilis nsp., paratype
MNHN. 22, 23. Trachypollia turricula (von Matzan), stn. DC34. 24, 25. Siphonochelus (Siphonochelus) riosi (Bertsch & DWttilio),
stn. CB104.
Page 30
THE NAUTILUS, Vol. 105, No. 1
R. Hoiiart, 1991
Page 31
axial sculpture. No apparent spiral sculpture except thick,
whitish intritacalx, strongly reticulate when thin surface
laver partially removed, otherwise minutely sculptured
with sharp axial striae (figure 2).
Aperture broad, roundly-ovate. Columellar lip smooth,
adherent. Anal notch broad, shallow. Outer lip weakly
erect, with 4 low denticles within. Siphonal canal short,
open, bent adaperturally at tip. Shell whitish.
Etymology: Named after J. H. Leal, School of Marine
and Atmospheric Science, Miami, Florida, member of
the scientific staff on board of the Marion-Dtifresne dur-
ing the MD55 cruise.
Discussion: The Dermonmrex of the Western Atlantic
region have been revised by Yokes (1975, 1976, and 1985)
and include the following Recent species: Dermomurex
(Dermonmrex) panperctiliis (C. B. Adams, 1850); Der-
nwmurex (Dermomurex) alabastrum (A. Adams, 1864);
Dermomurex (Dermomurex) pacei Petuch, 1988; Der-
momurex (Gracilimurex) elizabethae (McGinty, 1940);
Dermomurex (Trialatella) abtjssicola (Crosse, 1865);
Dermomurex (Trialatella) oxum Petuch, 1979; Der-
momurex (Trialatella) glicksteini Petuch, 1987; Der-
momurex (Trialatella) kaicherae Petuch, 1987; Der-
momurex (Trialatella) cuna Petuch, 1990.
In addition there are several fossil species, discussed
and illustrated in Yokes (1975).
The holotype of D. abyssicola has not been located
(Yokes, 1975:148) but the original illustration shows a
shell with 3 strong varices and intervaricial ridges, while
D. leali has 5 varices on the last teleoconch whorl. Der-
momurex abyssieola has a brown band at the base of its
last whorl and a longer siphonal canal.
Dermomurex leali differs from D. oxum in having 5
varices on its last whorl instead of 3 strong varices and
intervaricial ridge between pairs of varices as in D. oxum.
Dermomurex leali has more rounded whorls, covered by
a more finely sculptured intritacalx (figure 2), its aperture
is relatively larger, the siphonal canal is shorter and the
spire is lower.
Dermomurex leali differs from D. glicksteini and D.
kaicherae by its more globose shell, its more numerous
varices on last teleoconch whorl, its absence of intervar-
icial axial ridges, and shorter siphonal canal.
In addition, D. oxum, D. glicksteini and D. kaicherae
have more or less obvious spiral sculpture when the in-
tritacalx is removed, whereas it is absent in D. leali.
Dermomurex leali differs from D. cuna by its more
rounded varices, shorter siphonal canal, comparatively
larger aperture and by its different and more sharply
striate intritacalx.
Poirieria (Pazinotus) stimpsonii (Dall, 1889)
(figures 6, 7, 32)
IVIaterial examined: Southeastern Brazil, stn. I)C22, Vi-
toria Bank, 20°32'S, 38°11'W, 52 m (2 dd); stn. CB79,
19°02'S, 37°48'W, 1,500-1,575 (1 dd); stn. DC82, 18°56'S,
37°52'W, 85-105 m (1 Iv).
Four juveniles, easily recognizable shells. The illus-
trated specimen (figure 32) is pinkish. The species is
known from western Florida to Barbados (Radwin &
D Attilio, 1976:87), but has not previously been recorded
south of Barbados.
Subfamily MURICOPSINAE Radwin & D'Attilio, 1971
Muricopsis (?Murexsul) sp.
(figures 8, 30, 31)
Material examined: Southeastern Brazil, stn. DC35, Ja-
seur Bank, 20°42'S, 35°22'W, 82-105 m, (1 Iv) (juvenile);
stn. DC43, Dogaressa Bank, 20°51'S, 33°45'W, 63 m (1
dd, 3 Iv) (juveniles).
The largest of the Brazilian specimens is 5.2 mm long.
All are pinkish. Examination of adult specimens is nec-
essary for positive identification.
Muricopsis (Risomurex) necocheanus (Pilsbry, 1990)
(figure 11)
Material examined: Southeastern Brazil, stn. DC15,
2r37'S, 40°18'W, 37 m (1 dd) (damaged); stn. DC22,
Yitoria Bank, 20°32'S, 38°11'W, 52 m (3 Iv); stn. CP25,
Yitoria Bank, 20°36'S, 37°27'W, 65 m (1 dd).
Recorded by Rios (1985:84) from Espirito Santo, Bra-
zil, to Bahaia Blanca, Argentina. Yokes and Houart (1986:
84) did not observe a complete protoconch, but, from
the material then available, they presumed the proto-
conch to be rounded instead of keeled as in most Muri-
copsis. The protoconch here illustrated (figure 11) con-
firms their assumption.
Favartia (Favartia) cellulosa (Conrad, 1846)
Material examined: Southeastern Brazil, stn. DC29,
Montague Bank, 20°27'S, 36°41'W, 310-350 m (1 dd).
Recorded by Rios (1985:85). The only specimen col-
lected during the expedition is an empty, juvenile, dam-
aged shell, obviously carried downslope from shallow
water.
Figures 26-33. 26. Dermomurex (Trialatella) oxum Petuch, stn. DC22, 15.5 x 8 mm. 27, 28. Dermomurex (Trialatella) leali
n.sp , holotype MORG 26457, 9.5 x 5.6 mm. 29. .\spella morchi Radwin & D'Attilio, stn DC40, 10 x 5 mm, 30, 31. Muricopsis
(?Murexsui) sp., stn. DC35, 5.2 x 3.2 mm. 32. Poirieria {Pazinotus) stimpsonii (Dall), stn. DC82, 6.5 x 4,2 mm, 33. Favartia
(Favartia) sp., stn. DC22, 8.2 x 5 mm. Figures 34-36. Favartia (Caribiella) varimutabilis n sp 34, 35. Holotype MORG 26458,
9x5 mm. 36. Paratype MNHN, stn DC22, 9 x 5.1 mm Figure 37. Pygmaepterys germainae Yokes & D'Attilio, stn. DC34,
5.5 X 3.4 mm.
Page 32
THE NAUTILUS, \ol. 105, No. 1
Favartia (Favartia) nucea (Morch, 1850)
Material examined: Southeastern Brazil, .stn. D(]15,
2r37'S, 40°18'W, 37 m (1 dd); stn. DCMO, Davis Bank,
20°40'S, 34''41'W, 60 m (1 Iv) (juvenile).
A species that has previously been recorded from Bra-
zil hv Rios (1985:85).
Favartia (Favartia) sp.
(figures 13, 33)
IVIaterial examined: Southeastern Brazil, stn. DC22, Vi-
toria Bank, 20°32'S, 38°11'W, 52 m (1 dd); stn. DC35,
Jaseur Bank, 20°42'S, 35°22'W, 82-105 m (3 iv) (juve-
niles).
The shell is apparently different from any known west-
ern Atlantic Favartia species. The shell and the proto-
conch of a juvenile specimen are here illustrated. The
colour is light pink to light brown.
The material collected during the expedition consists
only of an adult but damaged shell (figure 33) and of 3
juveniles. There is presently insufficient material to com-
pare in more detail with other Favartia.
Favartia (Favartia) varimutabilis n.sp.
(figures 12, 20, 21, 34, 35, 36)
Type material: Holotvpe MORG 26458, 9x5 mm,
southeastern Brazil, stn. DC15, 21°37'S, 40°18'W, 37 m
(dd); 3 paratypes MNHN: stn. DC22, Vitoria Bank,
20''32'S, 38°11"W, 52 m (1 dd); stn. DC27, Montague
Bank, 20°26'S, 36°42'W, 81 m (1 Iv); stn. DC40, Davis
Bank, 20°40'S, 34°41'W, 60 m (1 dd).
Other material examined: Stn. DC24, Vitoria Bank,
20°42'S, 37°50'W, 48-52 m, coll. R. Houart (1 dd).
Description: Shell 8.5-9 mm in length, stout, fusiform.
Spire high, with 2.5 protoconch whorls and 5 weakly
convex, teleoconch whorls. Suture impressed. Protoconch
whorls rounded, ending with a deeply indented varix.
Last whorl with 3-5 irregular, rounded, erratically placed,
squamous varices. Penultimate whorl bearing 8 or 9 var-
ices. No intervaricial axial sculpture. Spiral sculpture of
7 cords, occasionally with 1 thread between each pair of
cords, 2 or 3 adapical cords sometimes extending as small
spinelets on varices.
Aperture roundly-ovate, outer lip erect and smooth,
weakly lirate within. Columellar lip rim erect, with 2 or
more small and narrow folds. Anal notch broad, shallow.
Siphonal canal short, narrowly open, weakly bent adap-
erturally, ornamented with 2 small open spines. Shell
pale orange to pale brown, with darker spiral cords.
Radula typically muricopsine with a prominent cen-
tral cusp (figures 20, 21).
Etymology: Vari: from varix; nmtalnlis: variable. Named
for its variable shape of shell and varices.
Discussion: It .seems doubtful that the shells illustrated
in figures 34, 35 and 36 are conspecific. In fact, the last
teleoconch whorl of F. varimutabilis may be weakly
convex or rounded, with erratical placement and variable
form of the varices. Early teleoconch whorls are identical
in all specimens examined.
Favartia (Carihiella) alveata (Kiener, 1842), known
from Florida to Brazil has a shell with fewer, larger,
scabrous cords, regularly placed varices and a paucispiral
protoconch of 1.5 whorls, denoting nonplanktotrophic
larval development. Favartia varimutabilis is also related
to the Pacific species F. jeanae Emerson and D'Attilio,
1979, known from the Philippine Islands.
Favartia (Murexiella) hidalgoi (Crosse, 1868)
IVIaterial examined: Southeastern Brazil, stn. DC55,
Martin Vaz Island, 20°32'S, 28°52'W, 780-795 m (1 dd)
(fragment); stn. DC81, 19°00'S, 37°48'W, 120-135 m (1
dd).
Known from the Gulf of Mexico to Brazil. In Brazil
recorded from Rio Grande do Sul by Rios (1985:85). Only
2 damaged and dead specimens were collected during
this expedition.
Favartia (Murexiella) glypta (M. Smith, 1938)
Material examined: Southeastern Brazil, stn. DC15,
21°37'S, 40°18'W, 37 m (1 dd, 1 Iv); stn. DC22, Vitoria
Bank, 20°32'S, 38°11'W, 52 m (3 dd) (juveniles); stn.
DC73, 19°00'S, 37°48'V^, 607-620 m (1 dd) (juvenile).
Comparison of the photograph of the holotype of Mu-
rex glypta Smith, 1938 illustrated in Vokes (1968:117),
of a very similar shell from Ilha do Pai, Brazil (coll. R.
Houart), and of the type material of Murexiella iemanja
Petuch, 1979 (holotvpe USNM 780652 and paratype
USNM 780653) confirms the conclusion of Rios (1985:
85) who synonymised Murexiella iemanja Petuch, 1979
with Murex glypta Smith, 1938.
The other shell illustrated as Favartia glypta in Vokes
(1968: pi. 7, fig. 5) from the post-Caloosahatchee For-
mation, Florida (specimen USNM 645895) is Favartia
celltilosa (Conrad, 1846).
Rios (1985:85) records F. glypta from Amapa to Rio
de Janeiro and Abrolhos Islands.
Fygmaepterys germainae Vokes and D'Attilio, 1980
(figure 37)
'^Fygmaepterys germainae Wikes and D'.\ttilio, 1980:50, pi. 1,
figs. 1-4.
Type locality: Off Punta Higuero, northwestern Puerto
Rico, 90 m.
Material examined: Southeastern Brazil, stn. DC24, Vi-
toria Bank, 20°42'S, 37''50'W, 48-52 m (1 dd); stn. DC27,
Montague Bank, 20°26'S, 36°42'W, 81 m (1 dd): stn.
DC34, Jaseur Bank, 20°28'S, 35°54'W, 54 m (1 dd).
Described from fossil specimens (Miocene and early
Pleistocene) from the Dominican Republic and Costa
R. Houart, 1991
Page 33
Rica and from Recent specimens from the Caribbean
Sea (Puerto Rico to Panama, in 12-90 m). The presence
of this species off southeastern Brazil extends consider-
ably its geographical range.
this synonymy. The multispiral protoconch indicates
planktotrophic development, which accounts for the
broad distribution of this species in the western and east-
ern Atlantic.
Subfamily ERGALATAXINAE
Kuroda and Habe, 1971
Cytharomorula graiji (Dall, 1889)
Material examined: Southeastern Brazil, stn DC28,
Montague Bank, 20°27'S, 36°42'W, 525-600 m (1 dd);
stn. DC48, Columbia Bank, 20°44'S, 32°08'W, 250-300
m (2 dd); stn. DC55, Martin Vaz Island, 20°32'S, 28°52' W,
780-795 m (14 dd); stn. DC73, 19°00'S, 37°48'W, 607-
620 m (3 dd); stn. CB77, 19°41'S, 37°48'W, 790-940 m
(2 dd); stn. DC81, 19°00'S, 37°48'W, 120-135 m (2 dd);
stn. DC82, 18°56'S, 37°52'W, 85-105 m (3 dd); stn. CB92,
19°34'S, 38°55'W, 340-360 m (1 dd); stn. CB103; 23°36'S,
42°02'W, 200-217 m (1 Iv).
Mentioned by Radwin and D'Attilio (1976:44) from
Barbados and by Rios (1985;86) from Rio de Janeiro to
Rio Grande do Sul in 80-175 m.
The species probably lives throughout the Caribbean
to Brazil. Also known in the Eastern Atlantic in the Ca-
nary Islands, Madeira and St. Helena (Bouchet & Waren,
1985).
Subfamily OCENEBRINAE Cossmann, 1903
TrachypoUia turricula (von Maltzan, 1884)
(figures 22, 23)
Cantharus (Pollia) turricula von Maltzan, 1884:67.
Type locality: Goree, Senegal, West Africa.
Material examined: Southeastern Brazil, stn. DC15,
21°37'S, 40°18'W, 37 m (1 dd); stn. DC22, Vitoria Bank,
20°32'S, 38°11'W, 52 m (4 dd, 1 Iv); stn. DC27, Montague
Bank, 20°26'S, 36°42'W, 81 m (1 dd, 2 Iv); stn. DC28,
Montague Bank, 20°27'S, 36°42'W, 525-600 m (1 dd);
stn. DC30, Jaseur Bank, 20°27'S, 36°17'W, 60 m (1 Iv);
stn. DC34, Jaseur Bank, 20°28'S, 35°54'W, 54 m (2 Iv);
stn. DC35, Jaseur Bank, 20°42'S, 35°22'W, 82-105 m (9
dd); stn. DC40, Davis Bank, 20°40'S, 34°41'W, 60 m (1
dd); stn. DC42, Dogaressa Bank, 20°55'S, 34°01'W, 60 m
(6 dd); stn. DC43, Dogaressa Bank, 20°5rS, 33°45'W, 63
m (1 dd, 3 Iv); stn. DC47, Columbia Bank, 20°42'S,
32°13"W, 94-105 m (1 dd); stn. DC61, Trindade Island,
20°29'S, 29°18'W, 63 m (1 dd); stn. DC82, 18°56'S,
37°52'W, 85-105 m (3 dd); stn. DC83, 18°50'S, 37°57'W,
60 m (1 dd).
The shell dredged at station DC28 (525-600 m) was
empty and damaged, and most likely carried downslope.
Drupa didyma Schwengel, 1943 described from off
Palm Beach, Florida, is similar; comparison with the
original diagnosis and with specimens from West Africa,
Florida (coll. R. Houart), and Brazil (MNHN) confirms
Subfamily TROPHONINAE Crossmann, 1903
Trophon aculeatus Watson, 1883
(figures 14, 40)
Trophun aculeatus Watson, 1882:390.
Type locality: Off Pernambuco, 09°05'S, 34°50'W, 640 m.
Material examined: Southeastern Brazil, stn. CB77,
19°41'S, 37°48'W, 790-940 m; stn. CB104, 23°42'S,
42''07'W, 430-450 m; stn. CB105, 23°47'S, 42°10'W, 610
m (many live and dead collected specimens).
Rios (1985:88) reported this species to occur off Brazil
(off Pernambuco) and Guadeloupe and included the type-
locality of T. lacunellus (Dall, 1889), a different species
that Rios synonymised with T. aculeatus. Live-taken
specimens were collected by R.V. Marion-Dufresne in
430-450 m. All specimens collected in 610-940 m were
empty shells.
Trophon pelseneeri E. A. Smith, 1915
Trophon pelseneeri E. A. Smith, 1915:92, pi. 2. figs. 6-7.
Type locality: West of Falkland Islands, 229 m.
Material examined: Southeastern Brazil, stn. CBlOl,
22''58'S, 42°06'W, 50 m (1 dd); stn. CB102, 23°07'S,
42"'04'W, 100 m (1 dd) (damaged).
Recorded by Rios (1985:88), from the state of Rio de
Janeiro, Brazil, to the Falkland Islands, Argentina. Tro-
phon orbignyi Carcelles, 1946 is a synonym (Rios, 1985:
88), Trophon pelecetiis Dall, 1902 is probably an earlier
name, but study of more specimens is required to confirm
this.
Trophon verrillii Bush, 1893
(figures 41-43)
Trophon verrillii Bush, 1893:214, pi. 1, fig. 16.
Type locality: Off Cape Fear, North Carolina, 1,183 m.
Material examined: Southeastern Brazil, stn. CB76,
18°59'S, 37°50'W, 637 m (3 dd, 4 Iv).
Trophon verrillii differs from T. aculeatus in having
a larger protoconch and totally lacking spiral ornamen-
tation. The illustrated specimen (figures 41, 42) is very
similar to the holotype (figure 43). To m\ knowledge T.
verrillii was previously known only from the type lo-
cality
Page 34
THE NAUTILUS, Vol. 105, No. 1
R. Houart, 1991
Page 35
30"
15"
ATLANTIC
OCEAN
Trmdade
0 100 200 km
20
24
Figure 46. Map of investigated area with location of cited stations. Reprodnced, with permission, from Leal and Bouchet (1991)
with some additions.
Trophon miicrone n.sp.
(figures 15, 16, 38, 39)
Type material: Holotype MNHN, 26.5 x 11 mm, south-
eastern Brazil, stn. CB79, 19°02'S, 37°48'W, 1,500-1,575
m (Iv); 7 paratypes MORG 26459; stn. DC70, 18°59'S,
37°48'W, 1,540-1,550 m (4 dd) (juveniles); stn. CB77,
19°41'S, 37°48'W, 790-940 m (1 dd) (damaged); stn. CB79,
(2 dd).
Other material examined: Stn. CB78, 18°59'S, 37°48' W,
1,200 m, coll. R. Houart (1) (damaged).
Distribution: Known only from the material examined.
Description: Shell 26.5-28.0 mm in length, roundly-
elongate. Spire high with 1.5-1.75 protoconch whorls and
6 or 7 teleoconch whorls. Suture impressed. Protoconch
whorls rounded. Axial ornamentation on last teleoconch
whorl consisting of 12 sharp, low lamellae and numerous
growth striae. Spiral sculpture of 4 or 5 low, weak cords;
shoulder smooth, except for axial lamellae.
Aperture broadly ovate. Columellar lip smooth. Anal
notch obsolete. Outer lip thin, smooth, glossy within.
Siphonal canal moderately long, straight, broadly open.
Shell whitish.
Etymology: Mwcrone-from mucro (tip, sharp end).
Named for its arrow or javelin tip shape.
Discussion: Three species, Trophon lacunellus (Dall,
1889), described from off Guadeloupe, in 1,406 m, T.
dabneyi Dautzenberg, 1889, described from the Azores,
in 1,287 m, and T. limicola Verrill, 1885, described from
the New England Coast, in 1,542-3,718 m may be com-
pared.
Trophon lacunellus differs from T. mucrone by its
higher spired and smaller shell without spiral sculpture,
narrower teleoconch whorls, and more rounded proto-
conch whorls.
Trophon dabneyi has more rounded teleoconch w horls
than T. mucrone, spiral cords that are absent or very
weak, and a multispiral protoconch of 2.5 whorls (prob-
Figures 38-40. ,38, 39. Trophon nuicrone n.sp., holotype MNHN, 26.5 x 11 mm. 40. Trophon aculealus Watson, stn. CB104,
13.9 X 7 mm Figures 41-43. Trophon verrillii Bush. 41, 42. Stn. CB76. 10 x .5 5 mm 43. Holotype, MCZl 19157, 13.9 x
6.8 mm Figures 44, 45. Siphonochehts (Siphonochelus) riosi Bertsch & D'Attilio, stn CB103, 11.5 x 5 2 mm.
Page 36
THE NAUTILUS, Vol 105, No. 1
able planktotrophic development) (Hoiiart, 1981: pi. 3,
fig. 14; Bouchet & Warra, 1985: fig. 305).
The holotype of T. limicola Verrill, 1885, a 10.5 mm
high shell, was also compared with a 9.7 mm high ju-
venile specimen of T. mucrune. Trophon limicola has
20 axial lamellae on its last teleoconch whorl (only 9 in
T. mucrone), a longer siphonal canal, more rounded
protoconch whorls, and lacks spiral sculpture.
Additional notes on West Atlantic Trophon species:
Trophon aculeatus Watson, 1883 and T. verrillii Bush,
1893 differ in many features such as spiral sculpture,
number of axial lamellae, shape of whorls and proto-
conch. Trophon abijssorum Verrill, 1885, described from
a juvenile specimen of 8 mm, is more closely related to
T. verrillii. and has angulate and smooth teleoconch
whorls, prominent spines at the shoulder and relatively
longer siphonal canal.
Subfamily TYPHINAE Cossmann, 1903
Typhis (Typhina) belcheri Broderip, 1833
Material examined: Southeastern Brazil, stn. CB102,
23°07'S, 42°04'W, 100 m (1 dd).
The holotype of Typhis belcheri was illustrated by
Kaicher (1980: card 2514). Comparison with the holotype
of Typhis cleriji Petit, 1842 (MNHN), and with many
specimens from Brazil (coll. R. Houart) and from West
Africa (MNHN and coll. R. Houart) lead to the conclu-
sion that these taxa are conspecific. The species is re-
corded by Rios (1985:89) as Riigotyphis cleryi (Petit,
1842), from Rio de Janeiro to Torres, Rio Grande do Sul.
It lives usually in shallow water.
Typhis belcheri is also known from the Cape Verde
Archipelago and Senegal (West Africa) (MNHN).
Siphonochelus (Siphonochelus) riosi
(Bertsch & D'Attilio, 1980)
(figures 17, 24, 25, 44, 45)
Typhis (Typhina) riosi Bertsch & D'Attilio, 1980:135, figs. 6, 7.
Type locality: Off Tramandai, southern Brazil, 30°04'S,
47°55'W, 280 m.
Material examined: Southeastern Brazil, stn. CB92,
19°34'S, 38°55'W, 340-360 m; stn. CB102, 23°07'S,
42°04'W, 100 m; stn. CB103, 23°36'S, 42°02'W, 200-217
m; stn. CR104, 23°42'S, 42°07'W, 430-450 m; stn. CB105,
23°47'S, 42°10'W, 610 m (many live and dead speci-
mens).
The specimens collected in 100-217 m are larger than
tho.se dredged in 430-450 m, but no other differences
arc apparent between the two samples. As already men-
tioned by Vokes (1984:215), the specimen illustrated as
Typhis longicornis Dall in Bayer (1971: fig. 31) is S.
riosi.
The radula (figures 24, 25) consists of rachidian teeth
and a pair of curved lateral teeth. Rachidian teeth with
3 main cusps and asymmetrical lateral denticles.
ACKNOWLEDGMENTS
I am very grateful to Dr. P. Bouchet (Museum National
d'Histoire Naturelle, Paris) and to Dr. J. H. Leal (Ro-
senstiel School of Marine and Atmospheric Science, Mi-
ami) for giving me tfie opportunity to examine this ma-
terial and for their much appreciated suggestions, to Dr.
R. S. Houbrick, D. M. Tyler, S. S. Greenhouse and F. J.
Collier (National Museum, of Natural History, Smithson-
ian Institution, Washington, DC) and to Dr. S. P. Kool
(Museum of Comparative Zoology, Cambridge) for the
loan of type material, to Dr. P. Bouchet and Dr. A. Waren
(Swedish Museum of Natural History, Stockholm) for
preparation and SEM work of the radulae, and to Dr. P.
Bouchet and Prof. E. H. Vokes (Tulane University, New
Orleans) for critical review of the manuscript.
LITERATURE CITED
Bertsch, H. and A D'.4ttilio. 1980. New species of Muricidae
(Gastropoda) from the Indian Ocean, the Philippines and
Brasil. Venus 39(3);131-138.
Bouchet, P and A. Waren. 1985. Revision of the Northeast
.'Ktlantic bathyal and abyssal Neogastropoda excluding
Turridae (Moliusca, Gastropoda). Bolletino Malacologico
supplement 1:123-296.
Bush, K. J. 1893. Reports on the results of dredgings, under
the supervision of Alexander Agassiz, in the Gulf of Mexico
(1877-78), and the Caribbean Sea (1879-80). and along
the Atlantic coast of the United States (1880) by the United
States coast survey steamer "Blake . . . Report on the
Moliusca dredged by the "Blake" in 1880, including de-
scriptions of several new species. Bulletin of the Museum
of Comparative Zoology 23:197-244
Clench, W J and I Perez Farfante. 1945. The genus Murex
in the western .Atlantic Johnsonia 1(17): 1-56.
Houart, R. 1981 Revision des Trophoninae d Europe. Infor-
mations de la Societe Beige de Malacologie 9(l-2):3-70.
Kaicher, S. D. 1980. Card catalogue of world-wide shells,
Muricidae V. Privately published, St. Petersburg, Florida.
Leal, J. H. and P. Bouchet. 1989. New deep-water Volutidae
from off southeastern Brazil (Moliusca: Gastropoda). The
Nautilus 103(1):1-12.
Leal, J. H. & P. Bouchet. 1991, Distribution patterns and
dispersal of prosobranch gastropods along a .seamount chain
in the Atlantic Ocean. Journal of the Marine Biological
Association of the United Kingdom 71(1) (February 1991).
In press.
Maltzan, H. F. von. 1884. Diagnosen neuer Senegambischer
gastropoden. Nachrichtsblatt der deuteschen Malakozool-
ogischen Gesellschaft 16:65-73.
Petuch, E J. 1979. New gastropods from the .\brolhos Ar-
chipelago and reef complex, Brazil Proceedings of the
Biological Society of Washington 92(3):5 10-526.
Petucli, E. J 1987. New Caribbean molluscan faunas. CERF,
Charlottesville, Virginia: 1-154, addendum .M-A4.
Kadwin, G E. and A. D'Attilio. 1976. Murex shells of the
world, an illu.strated guide to the Muricidae. University
Press, Stanford, 284 p
Rios. E. C. 1985. Seashells of Brazil Rio Grande, 328 p.
R. Houart, 1991
Page 37
Smitli, E. A, 1915. Mnllusca. Pt. 1, Gastropoda prdsohraiR-liia,
Scaphopoda and Pelec\poda -British Antarctic ( "Terra
Nova") expedition, 1910. Zoology 2(4):61-112.
Yokes, E. H. 196S. Cenozoic Muricidae of the western At-
lantic region. Ft 4, Hexaplex and MurexicUa. Tulane Stud-
ies in Geology 6(2-3):85-126.
Yokes, E. H. 1975. Cenozoic Muricidae of the western At-
lantic region, Ft 6, Aspclla and Dermomurcx. Tulane Stud-
ies in Geology and Paleontology 11:121-162.
Yokes, E. H. 1976. Cenozoic Muricidae ot the western At-
lantic region. Dcrmonuircx — addendum Tulane Studies
in Geology and Paleontolog\ 1245-46.
Yokes, E. H. 1984. Comparison of the Muricidae of the east-
ern Pacific and western Atlantic, with cognate species.
Shells and Sea Life 16(11):210-216.
Yokes, E. H. 1985. Review of the west coast aspelloids Aspclla
and Dermomurex (Gastropoda: Muricidae), with the de-
scriptions of two new species. The Yeliger 27(4):430-439.
Yokes, E. H. 1990 Two new species of Chicoreitu subgenus
Sirfl^i.? ((Gastropoda: Muricidae) from Northeastern Brazil.
The Nautilus 103(4):124-130.
N'okes, E. H. and A. D'Attilio. 1980. I'ygmaepterys, a newly
described taxon of Muricidae (Mollusca: Gastropoda), with
the description of three new species from the Onozoic of
the western Atlantic Tulane Studies in Geology and Pa-
leontolog) 16:45-54
Yokes, E. H. and H Houart. 1986. An evaluation of the taxa
Miiricopsis and Ri.s(jmurcx (Gastropoda. Muricidae), with
one new species of Risoiiiiircx. Tulane Studies in Geology
and Paleontology 19:63-88.
Watson, R. B. 1882. Mollusca of H.M.S. Challenger expedi-
tion. Journal ot the Linnean Society London Part 14372-
392.
THE NAUTILUS 105(1):38, 1991
Page 38
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Museum of Comparative Zoology
Harvard University
Cambridge, MA 02138
Dr. Aurele La Rocque
Department of Geology
The Ohio State University
Columbus, OH 43210
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 Zoology
Harvard University
Cambridge, MA 02138
Ms. Paula M. Mikkelsen
Harbor Branch Oceanographic
Institution, Inc.
Ft. Pierce, FL 33450
Dr. Donald R. Moore
Division of Marine Geology
and Geophysics
Rosenstiel School of Marine and
Atmospheric Science
University of Miami
4600 Rickenbacker Causeway
Miami, FL 33149
Mr. Richard E. Petit
P.O Box 30
North Myrtle Beach, SC 29582
Dr. Edward J. Petuch
Department of Geology
Florida Atlantic University
Boca Raton, FL 33431
Dr. David H. Stansbery
Museum of Zoology
The Ohio State University
Columbus, OH 43210
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 ("alifornia at Davis
Davis, CA 95616
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TH Et7NAUTI LUS
CONTENTS
Volume 105, Number 2
April 10, 1991
ISSN 0028-1344
Riidiger Bieler
Alan R. Rabat
William K. Emerson
Malacological Journals and Newsletters, 1773-1990
39
First records for Cymatium mundiim (Gould) in the
eastern Pacific Ocean, with comments on the
zoogeography of the tropical trans-Pacific Tonnacean and
non-Tonnacean prosobranch gastropods with Indo-Pacific
faunal affinities in western American waters
62-
Or>
LIBRARY
APR 2 3/991 j
.i!!!^l_^. Mass. /
THE NAUTILUS 105(2):39-61, 1991
Page 39
Malacological Journals and Newsletters, 1773-1990
Riidiger Bieler'
Delaware Museum of Natural History
P O. Box 3937
Wilmington, DE 19807, USA
Alan R. Kabal-
Museum of Comparative Zoology
Harvard University
Cambridge, MA 02138, USA
ABSTRACT
This compilation lists the 286 malacological journals and news-
letters (representing 374 titles), arranged by continent and
country The place, duration, and status of each publication is
given and changes in titles are cross-referenced. Of these serials,
157 are still published It is determined that about 25?o of
current malacological publications are published in these jour-
nals devoted solely to mollusks. The geographical distribution
(31 countries represented) and the usage of malacological jour-
nals are analyzed in terms of the broader context of the scientific
journal literature field.
Key words: Bibliography, journals; malacology; serials.
INTRODUCTION
Malacologists all over the world are often overwhelmed
b\' the enormous diversity of journals and newsletters in
our field. It is virtually impossible for any one library to
possess all these journals, let alone keep up with the new
serials arising every year. The purpose of this paper is
to introduce some order to this chaos by providing com-
plete bibliographical data on the corpus of malacological
serials. It has been over three decades since the useful,
albeit cursory, compilation of Jutting and Altena (1958)
who listed only 36 titles. More recently, Biirk and Jung-
bluth (1985) provided a detailed index to most of the
German malacological publications; this valuable refer-
ence essentially supersedes previous German efforts such
as Buschmeyer's (1938). A brief treatment of malacolog-
ical serials as part of the history of conchology was pre-
sented by Dance (1986:145).
Scientific journals began in 1665 with the appearance
of the Philosophical Transactions [London] and the
Journal des Sqavans [Paris]. It would take another cen-
' Current address: Division of Invertebrates, Field Museum
of Natural Historv, Roosevelt Road at Lake Shore Drive, Chi-
cago, IL 60605, USA.
^Current address: Division of Mollusks NHB-118, National
Museum of Natural History, Smithsonian Institution, Wash-
ington, DC 20560, L SA
tury for the start of the first malacological journals: the
late 1700's German serials of J. S. Schroter who published
four short-lived serials dealing primarily with fossil mol-
lusks (Friess, 1982:93-95). However, "modern" mala-
cological journals did not come into full bloom until the
mid 1800's, with several European titles (all now de-
funct). The oldest currently published malacological
journal is the Archiv fiir Mollushenkunde [1868, under
an earlier title]. The Zoological Record, Mollusca Section
[1864] is an abstracting source and not a contributed
journal.
As the historian Derek Price (1986:5-6, 18-19) has
noted, the total number of scientific journals has doubled
every fifteen years, for a 5% increase per year. Yet, ob-
viously this exponential rate of increase cannot continue
infinitely; eventually a saturation point representing lo-
gistic growth will be reached. The proliferation of new
journals in the last decade does not seem to indicate that
the field of malacology has reached its carrying capacity,
although the recent demise of certain journals may ini-
tiate this trend.
Equally important is the usage of scientific articles:
how widely they are read and cited by others. Price
(1986:118) used Lotka's law and the Pareto distribution
to determine that ". . . about 10 percent of all published
papers have never been cited, about 10 percent have
been cited once, about 9 percent twice, and so on, the
percentages slowly decreasing . . .". This widely quoted
statement [usually paraphrased as "10 percent of all pub-
lished papers never subsequently cited" (Wheeler, 1989:
1 1)] is the inevitable consequence of what the sociologist
Robert Merton (1968:61) diagnosed as "insanabile scri-
bendi cacoethes [= the itch to publish].
MATERIALS AND METHODS
We undertook this project over a two year period,
entailing considerable bibliographic research and cor-
respondence. We compiled a master list based on the
serial holdings of seven major malacological libraries:
Academy of Natural Sciences, Philadelphia; Delaware
Museum of Natural History; Field Museum of Natural
History; Museum National dHistoire Naturelle, Paris;
Museum of Comparative Zoology; The Natural History
Museum, London; and National Museum of Natural His-
Page 40
THE NAUTILUS, Vol. 105, No. 2
torv, Smithsonian Institution. Then, this hst was aug-
mented by extensive correspondence with various col-
leagues who checked certain geographical sections, as
well as with the editors of shell club newsletters (ad-
dresses obtained from the American Malacological Union
membership list and various other sources); these indi-
viduals are listed in the acknowledgements. Unfortu-
nately, no replies were received from some shell clubs
and certain other clubs are no longer extant and we were
unable to find any complete sets of their newsletters.
While most of the major malacological research li-
braries have complete sets of the principal journals in
the field, the same is certainly not true for the numerous
shell club newsletters. It is to be regretted that some
museums extend little effort to keep even the newsletter
of their own national malacological society. We are aware
of the difficulties in treating this "grey" literature and
we strongly recommend that malacological libraries en-
deavor to maintain holdings of these newsletters (es-
pecially those published in their country) to the greatest
extent possible. The development of online computerized
catalogues of library holdings will allow researchers to
readily locate these serials.
SCOPE
Herein we have attempted as complete a compilation as
is possible of all the malacological serials (journals and
newsletters) that have come to our attention. As the read-
er may be aware, it is not always easy to differentiate
between malacological journals and newsletters. Gen-
erally, scientific journals publish original research articles
and are intended as part of the permanent scientific
record. They typically have an editorial review board to
maintain scientific standards, and they are often pub-
lished by a scientific institution or scholarly society.
Newsletters usually serve as a vehicle for news and in-
formation within the respective organization, and in the
case of shell club newsletters they disseminate knowledge
in layman's terms, offer advice for beginners in the field
and are often simply meant to entertain.
Shell club newsletters are not only of interest to the
scientific community because they occasionally provide
such original data as biogeographical or habitat infor-
mation, but also they become permanent scientific record
once they (sometimes unintentionally) publish taxonomic
statements, especially new species names and type des-
ignations. There is obviously no distinct line between
journals and newsletters, seeing that some of today's jour-
nals started in newsletter format, while some self-pro-
claimed "journals " hardly deserve that label. We made
no attempt to sort the malacological serials into these
two categories. We have usually excluded those shell club
newsletters which are completely restricted to internal
communications, such as membership lists and an-
nouncements of upcoming events.
We do not include multi-volume malacological trea-
tises that were issued "in parts " over long time periods
but represent a single encyclopedic compilation and not
a diverse journal. For example, Kiener's Iconographie
des Caquilles Vivantes; the Martini-Chemnitz and Kiis-
ter editions of the Systematisches Conchylien-Calnnet;
Phihppi's Abbildungen und Beschreibiingen . . . ; Reeve's
Conchologia Iconica; the Rossmassler-Kobelt Iconogra-
phie der Land- und Siisswasser-Molliisken, Sowerby's
Thesaurus Conchyliorum; and the Tryon-Pilsbry Man-
ual of Conchology are all excluded from this paper. We
also do not include symposia volumes, including those
of regular meetings, unless specifically issued in serial
format.
Several of the French titles in this list actually rep-
resent compiled reprints, by one author, of malacological
articles from various journals. We have included them
since they are obscure and are sometimes cited as if they
were serials themselves.
We noted several interesting aspects regarding the
etymology of the titles of these various serials. Obviously,
most are based on "mollusk", "malaco-", "conch-", or
"shell "; others from generic names or famous malacol-
ogists. One should note several titles that have been used
more than once: Argonauta, The Conchologist, Journal
of Malacology and Tide Lines.
For completeness, and in appreciation of the fact that
there is also some intended humor in our field of science,
we have included five satirical "serials" in our listings.
Three of them, titled Conchologica Nonsensica, The
Double Helix and Slug Newsletter, are easily spotted,
but The Slug and especially the cleverly produced Jour-
nal of Molluscan Behaviour are less obvious. These par-
odies were not included in our statistics.
In compiling this list, we also noted several journals
with "malacological" titles, although they do not qualify
for inclusion herein as they are not about mollusks at all,
or are not exclusively about mollusks. We list them here
to avoid future confusion: Nautilus: A monthly miscel-
lany . . . [1845-1846; Sag Harbor, New York]; Astarte,
A Journal of Arctic Biology [1951-1983; Tronis0]; The
Conch Shell [1953-1966; Bishop Museum, Honolulu];
The Conch: A Biafran Journal of Literary and Cultural
Analysis [1969-current; Paris]. An exception is made for
the larger shellfisheries journals (which also cover crus-
taceans). Here we draw an arbitrary line between those
that are included {e.g.. Journal of Shellfish Research).
and others that are not {e.g.. Proceedings of the ... th
National Shellfish Sanitation Workshop). Coche (1983)
lists numerous serial publications in shellfisheries and
aquaculture.
We would greatly appreciate information regarding
any additions or emendations to this list, since it is our
intention to publish future "Addenda". There are several
shell club newsletters for which we were unable to obtain
full data, despite repeated enquiries. In some cases, the
shell clubs themselves do not know the full history of
their own newsletter(s). These titles are indicated by an
asterisk; the information needed is noted. It is imperative
that the Zoological Record also be informed of new-
publications; editors should send copies (or announce-
ments) of their publications to: MoUusca Section, The
R Bieler and A. R. Rabat, 1991
Page 41
Natural History Museum, Cromwell Road, London SW7
5BD, England The authors will gladly provide editors
with a list of the major malacological research museums
that should also be informed of new malacological
publications.
DISCUSSION
It is worthwhile to consider the role of malacological
journals, not onl\' within the field of malacology, but
more broadh within the biological sciences. We have
found that there are 157 currentK published malacolog-
ical serials. According to The Serials Directory, 3rd Edi-
tion, 1988-89, there are over 118,000 current serial titles
in all fields of knowledge (excluding newspapers). More
specifically, according to the 1988/89 Zoological Record
Serial Sources (1990:vi), there are 5,540 current journals
in the zoological sciences that are recorded for the Zoo-
logical Record. Obviously, the malacological journals are
a very small part of the whole of scientific literature.
Yet, their importance in the field of malacology is far
greater than might have been suspected. In order to
evaluate this, we have analyzed the titles recorded in the
Zoological Record, Mollusca Section, to determine the
proportion of malacological articles published in mala-
cological serials. The results are as follows:
% published
Number
Number
in malac
of malac.
Year
of titles
serials
serials
1880
181
23.2%
12
1930
1,02.3
24.7%
8
1980
3.904
26,0%
120
[The results for 1880 are based on the entire sample; those for
1930 and 1980 are each based on the average of three sub-
samples of 100 entries].
It is a remarkable consistency that over the last cen-
tury, about 25% of all malacological titles covered by the
Zoological Record have been published in malacological
journals. The malacologist is (or should be) cognizant
that malacological journals are rarely read by non-mala-
cologists, while non-malacological journals usually reach
a wider audience. Hence it is not surprising that the bulk
of malacological journals are devoted to papers on sys-
tematics, evolution, and organismic-level studies; more
reductionist malacological studies {e.g., biochemistry,
molecular and cell biology) are invariably found in the
appropriate non-malacological journals.
It is also worthwhile to consider the geographic dis-
tribution of malacological serials in relation to the dis-
tribution of malacologists and more broadly in terms of
the overall scientific literature. It is obvious that most
journals are published in Europe and the United States;
a more detailed analysis reveals several interesting as-
pects. Altogether, thirty-one countries are represented
by malacological serials. We have counted 286 serials
(374 including changes in titles) of which 157 (54.9%)
are still published. Of the 157 current serials, we infor-
mally consider 45 (28.7%) to be professional journals and
the remainder as newsletters. The countries with the most
serials (and the numbers of serials — not counting changes
in titles) are: USA (117); Great Britain (22); France (19);
Japan (19); Australia (17); Italy (14); and Germany (13).
The distribution by world region is as follows:
Total
ber of
titles
121
6
29
39
171
374
Number
of serials*
94
5
25
29
125
8
286
Number
currently
published
51
3
14
20
65
4
157
Region
Europe (12)
Africa (1)
Asia (5)
Oceania (4)
North .America (3)
Central and South
America (6)
[world totals] (31)
Numbers in parentheses indicate the number of countries.
* Not counting changes in titles.
More specifically, it is obvious that no one malacologist
can read, let alone make reference to, all these mala-
cological serials. In his scientometric analyses. Price (1986:
67 ff.) noted that the usage of the scientific literature is
a function of the inverse square law. That is, given the
total number of serials in a field, half the reading will
be done from the square root of that total number. Ap-
plying this valuable heuristic concept to the field of mal-
acology, we note that there are 157 currently published
malacological serials. The square root of 157 is 12.5.
Therefore, one can conclude that despite the overabun-
dant diversity of current titles, only about 11-12 will
account for half the total usage of malacological serials
by malacologists.
An informal poll ["Of the following malacological
journals, which eleven do you consider to be the most
important?"] of nine professional malacologists revealed
a remarkable consensus of opinion: (a) 5 journals were
listed by all 9 respondents; (b) 5 journals got 6, 7 or 8
votes; (c) 3 journals got 3, 4 or 5 votes; and (d) 5 journals
got only 1 or 2 votes. Altogether, a total of thirteen [13]
journals received at least 3 votes; this represents only
8.2% of the total number of current malacological serials.
It does not seem to be essential for a malacological
research library to subscribe to every current malaco-
logical publication. Indeed, a compilation of the sub-
scription prices for these 13 journals comes to almost
$350 (at individual or member rates), which is reasonable
when compared to other fields of science. No malaco-
logical journals are issued by commercial (for-profit) pub-
lishing houses, which are often much more expensive
than institution or society journals (Feldmann, 1989). In
the field of systematics, it is obviously important to have
complete runs of the major journals, including those that
are now defunct. It is difficult to estimate the current
cost of obtaining a "complete" malacological journal li-
brary; certainly in excess of $50,000 as one rarely sees
these items in book dealer catalogues and there are rel-
Page 42
THE NAUTILUS, Vol. 105, No. 2
atively few comprehen.sive private malacological librar-
ies that are potentially available.
Another element of interest is the overall numerical
abundance and growth of the malacological literature,
including all publication sources. Solem (1974:7-8) has
estimated the size of this literature and his results are
comparable to ours; Schopf (1967) provided similar cal-
culations for the Ectoprocta [= Bryozoa]. Based on direct
counts and estimates from the Zoological Record and
Ruhoff (1980), we have compiled data on the total num-
ber of malacological publications, on an annual basis. As
of 1990, we have determined that approximately 167,000
such publications have appeared. The cumulative totals
from antiquity to the following years are: 1875 (9,000);
1900 (19,330); 1925 (34,965); 1950 (57,115); 1975
(106,060); and 2000 (estimate 210,000). This literature
is doubling every 25 years which seems rather dramatic;
however. Price (1986:5-6) documented that the overall
scientific literature is doubling every 15 years and certain
fields even faster, thus malacology is actually increasing
at a slower rate than the overall scientific literature field.
A matter of considerable nomenclatural importance is
the description of new species and higher taxa in news-
letters or non-professional journals. All too often, these
non-refereed papers are not widely available, and the
type specimens are commonly not deposited into rec-
ognized museums (for further discussion, see Lilllco, 1990
and Loch, 1990). We hope that the forthcoming revision
of the International Code of Zoological Nomenclature
will resolve these problems.
In conclusion, these analyses reveal several interesting
aspects of the field of malacological journals and news-
letters, including their importance in the field of mala-
cology and their geographic distribution. It seems that
the quantity (if not quality) of malacological journals
will continue to increase arithmetically (if not exponen-
tially), especially as new computer desk-top publishing
techniques tempt private individuals to start up their
own journals.
For each entry, we have included the full title along
with the place(s), duration and status of the publication.
For the journals and the newsletters of general interest,
we have included the subscription or membership ad-
dress (which is not always the same as the place of pub-
lication). For shell club newsletters of only local interest,
we have not included their mailing address; these are
listed in Rice (1990). We also list supplements or special
editions. References to cumulative indexes or other ex-
ternal sources of information about the journals are in-
cluded. The periodicals have been arranged alphabeti-
cally within each country, according to the first word
that is not an article. Major changes in title are listed
separately; minor changes in wording of the subtitle are
indicated by brackets, with alternate versions separated
by a slash (/).
Europe/Near East 42
Africa 47
Asia 48
Oceania 49
North America 50
Central and South America 57
EUROPE/NEAR EA.ST
Belgium
Achatina. A Newsletter of African non-marine malacol-
ogy. [Tervuren; Leiden (The Netherlands)]. 1 [1970]-13
[1985].
Annales de la Societe Malacologique de Belgique.
[Bru.xelles]. 1 [1863 ]-15 [1880]. (Comprising "Memoires"
and "Bulletin [des Seances]".) Continued as Annales de
la Societe Rovale Malacologique de Belgique. See Back-
buys (1985);'Dhondt (1989).
Annales de la Sociele Royale Malacologique de Bel-
gique. [Bru.xelles]. 16 [1881]-37 [1902]. Continuation of
Annales de la Societe Malacologique de Belgique. Con-
tinued as Annales de la Societe Royale Zoologique et
Malacologique de Belgique.
Annales de la Societe Royale Zoologique et Malaco-
logique de Belgique. [Bruxelles]. 38 [1903]-52 [1921].
Continuation of Annales de la Societe Royale Malaco-
logique de Belgique.
Apex. [Bruxelles]. 1 [Feb. 1986]-current. Continuation
of Informations de la Societe Beige de Malacologie. |M.
J. Buyle, Av. Maurice Maeterlinck 56, B-1030 Bruxelles,
Belgium r
Arion. Bulletin de contact [trimestriel] de la Societe Beige
de Malacologie. [Bruxelles]. 1 [Jan. 1977]-current, {ad-
dress: see Apex I
Bulletin Mensuel d'lnformation de Malacologie Les
Naturalistes Beiges [variously titled]. [Bruxelles]. 1 [1966]-
4 [1971]. Continued as Informations de la Societe Beige
de Malacologie.
Bulletin Periodique, Fondation Conchyliologique de
Belgique/Belgische Concholiologische Vereniging/
Conchological Foundation of Belgium [Rhode-Saint-
Genese]. 1 [1961]. [Single number: English, Flemish and
French editions].
La Conchioline. Bulletin mensuel du Groupement Beige
d'etude des Coquilles et Mollusques. [Bruxelles]. 1 [1976]-
15 [1977].
Gloria Maris. Tijdschrift uitgegeven door de Belgische
Vereniging voor Conchyliologie [, Malacologie en Pa-
leontologie]. [Antwerpen]. 1 [1967]-current. Also, "Me-
dedelingenblad' : local members newsletter and meeting
announcements, 1 [1986]-current. Miscellaneous supple-
ments, "Bijvoegsel", loose pages. {Ferd. Verbieststr. 9/6,
B-2030 Antwerpen, Belgium}
lnforination»9 de la Societe Beige de Malacologie. [Brux-
elles], 1 [Jan. 1972]-I3[Nov. 1985]. Continuation of Bul-
letin Mensuel d'lnformation de Malacologie Les Natura-
listes Beiges. Continued as Apex. New taxa listed by
Duchamps (1986).
Proces-Verbaux des Seances de la Societe Malacolo-
gique de Belgique. [Bruxelles] 1 [1872]-9 [1880]. Con-
R. Bieler and A. R. Kabat, 1991
Page 43
tinued as Proces-V^erbaux des Seances de la Soeiete Roy-
ale Malacologique de Belgique. Largely overlaps
with the "Bulletin" of the Annales de la Soeiete Royale
Malacologique de Belgique; see that entry.
Proces-Verbaux des Seances de la Soeiete Royale Ma-
lacologique de Belgique. [Bruxelles], 10 [1881}-27 [1898].
Continuation of Proces-Verbaux des Seances de la Soeiete
Malacologique de Belgique.
France
[for French Polynesia and New Caledonia, see under
Oceania]
Amenites IVlalacoIogiques par M. J. R. Bourguignat.
[Paris]. 1 [1853-1856]-2 [1856-1860]. Collected reprints
from Revue et Magasin de Zoologie.
Annales de Malacologie. Soeiete Malacologique de
France. [Paris]. 1 [1870-1884]-2 [1884-1886].
Archives Malacologiques par M. Jules Mabille. [Paris].
1 [1867]-5 [1869].
Bibliotheque Conchyliologique (J. C. Chenu). [Paris],
{serie 1) 1 [1845]-4 [1846]; (serie 2) 1 [1845]. The 6rst
series is a translation of seven English-language mala-
cological monographs; the second series is a translation
of the malacological articles from the Transactions of the
Linnean Society (London), volumes 1-17, 1791-1835.
Bulletins de la Soeiete Malacologique de France. [Paris].
1 [Jan. 1884]-7 [1890].
Contributions a la faune malacologique fran^aise (A.
Locard). [Paris]. 1 [1889]-16 [1891]. [Some are reprinted
from several French journals].
Elona. Bulletin de liaison de Malacologie continentale
fondamentale et appliquee. A Newsletter of French non-
marine Malacology. Soeiete Fran^aise de Malacologie.
[Paris]. 1 [1974]-6[1980].
Haliotis. Soeiete Fran^aise de Malacologie. [Paris]. 1
[1971]-current. (Lab. Biol. Invertebres Marins et Ma-
lacologie, Museum National d'Histoire Naturelle, 55 rue
de Buff on, F-75005 Paris 05, France!
Journal de Conchyliologie. Comprenant lEtude des
[Animaux, des Coquilles vivantes et des Coquilles fos-
siles/Mollusques vivants et fossiles]. [Paris; Mosman (Aus-
tralia)]. 1(1) [Feb. 1850]-1 15(4) [Dec. 1979]. For dates of
early issues, see Fischer-Piette (1937) and Winckworth
(1936, 1937). History: Fischer (1978). Index issues: 1850-
1872 (1878); 1873-1892 (1897); 1893-1936 (1940). Dance
(1986:145) erroneously stated that this journal "has con-
tinued publication to this day".
Magasin de Conchyliologie. [Paris]. 1 [1830]-2 [1845].
Reprints of molluscan sections of the Magasin de Zoolo-
gie, d'Anatomie Comparee et de Paleontologie, Deux-
ieme Section, Mollusques et Zoophytes.
Mappa. Club fran^ais des collectionneurs de coquillages.
[Paris]. 1 [1974]-9 [1980]. Continued as Xenophora.
Miscellanees Malacologiques, par A. de Saint-Simon.
[Toulouse]. 1 [1848]-2 [1856].
Le Nouveau Pes-Pelicani. [Nice]. 1 [1973]-8 [1974]. Con-
tinuation of Pes-Pelicani.
Nouvelles Miscellanees Malacologiques par M. le Doc-
teur Paladilhe. [Paris]. 1 [1866]-4 [1869]. [Compiled re-
prints from Revue et Magasin de Zoologie].
Pes-Pelicani. [Nice]. 1 [1968]-17 [1972]. Continued as
Le Nouveau Pes-Pelicani.
*Pyrum. Bulletin de liaison et d etudes des collection-
neurs de coquillages Region Provence Cote d'Azur. [Tou-
lon]. 1 [1975/76?]-7 [post 1977?]. [Issues 1-3 and 6-7
were undated; 4 and 5 dated 1977].
Revue Biographique de la Soeiete Malacologique de
France. [Paris]. 1 [1870]-2 [1886].
Series Conchyliologiques comprenant TEnumeration
de Mollusques terrestres et fluviatiles recueillis pendant
le cours de differents voyages, ainsi que la description
de plusieurs especes nouvelles (A. Morelet). [Paris] 1
[1858]-4 [1875].
Les Spicileges Malacologiques par M. J. R. Bourguig-
nat. [Paris]. 1 [1860]-15 [1862]. [Mostly collected reprints,
from Revue et Magasin de Zoologie].
Vertigo. Association Frangaise pour I'Etude des Mol-
lusques continentaux. [Chartres]. 1 [1990]-current. {12
rue Saint-Michel, F-28000 Chartres, France!
Xenophora. Bulletin du Club Franfais des Collection-
neurs de Coquillages/Bulletin de I'Association Fran9aise
de Conchyliologie. [Paris]. 1 [Jan. 1981]-current. Con-
tinuation of Mappa. {1 impasse Guemenee, F-75004 Par-
is, France}
Germany
[See Biirk and Jungbluth (1985) for an overall review
and index of most of these journals]
Abhandlungen des Archiv fiir Molluskenkunde. See en-
try under Archiv fiir Molluskenkunde.
Acta Conchyliorum. Club Conchylia e.V. [Darmstadt;
Stuttgart]. 1 [1981]-current. Number 2 [May 1990] was
also sold as a book, "News on Conidae . . .". JNeckar-
anlage 6, W-6930 Eberbach am Neckar, Germany!
Archiv fiir Molluskenkunde. [Frankfurt am Main]. 52(1)
[Jan. 1920]-current. A supplement series, Abhandlungen
des Archiv fiir Molluskenkunde, 1(1) [Jan. 1922]-2(2)
[July 1929]. Continuation of Nachrichtsblatt der
Deutschen Malakozoologischen Gesellschaft. | For-
schungsinstitut Senckenberg, Senckenberganlage 25,
\V-6000 Frankfurt-am-Main 1, Germany}
Beilage zum Nachrichtsblatt. See entry under Nach-
richtsblatt der Deutschen Malakozoologischen Gesell-
schaft.
*Club Conchylia Informationen. [Eberbach]. 1 [19 ?]-
current. Continuation of the Concha.
Concha. Informationen des e[ingetragenen] V[ereins] Club
Conchylia. [Wilnsdorf-Obersdorf]. 1 [1969]. Continued
as Club Conchvlia Informationen.
Page 44
THE NAUTILUS, Vol. 105, No. 2
Conchologische Mitlheilungen. [Kassel] I [1881 ]-3
[1894], Continuation of Novitates Conchologicae.
Fiir die Litteratur und KenntniO der INaturgeschichte,
sonderlich der Conchylien und der Steine. [Weimar].
1 [1781]-2 [1782]. Continuation of Journal fiir die Lieb-
haber des Steinreichs und der Konchyliologie. Continued
as Neue Litteratur und Beytrage zur KenntniB der Na-
turgeschichte, vorziiglich der Conchylien und FoDilien.
Heldia. Miinchner Malakoiogische Mitteilungen. [Miin-
chen]. 1(1) [May 1984]-current. Sonderheft 1 [1987]
"Land- und SiiBwassermollusken der Maltesischen In-
seln", K.-H. Beckmann. {Postfach 26-01-23, W-8000
Miinehen 26, Germany f
Jahrbiicher der Deutschen Malakozoologischen Ce-
sellschaft. [Frankfurt am Main]. 1 [1874]-14 [1887].
Journal fiir die Liebhaber des Steinreichs und der Kon-
chyliologie. [Weimar]. 1 [1773]-6 [1780]. Continued as
Fiir die Litteratur und KenntniC der Naturgeschichte,
sonderlich der Conchylien und der Steine.
Malakoiogische Abhandlungen. Staatliches Museum fiir
Tierkunde in Dresden. [Dresden]. 1 [1964]-current. | Au-
gustusstraBe 2, O-8010 Dresden, Germany}
Malakozoologische Blatter. [Kassel]. 1 [1854]-25 [1878];
new series 1 [1879]-11 [1891] (reprinted, 1969, Otto
Koeltz, Koenigstein-Taunus). Continuation of Zeitschrift
fiir Malakozoologie.
Mitteilungen der Berliner Malakologen. [Berlin]. 1
[1953]-19 [1967].
Mitteilungen der Deutschen Malakozoologischen Ge-
sellschaft. [Frankfurt am Main]. 1(1) [June 1962]-cur-
rent. [Volume numbers discontinued after 3(36); only as
issues thereafter.] Supplement 1 [1982] "Bericht iiber den
ersten Workshop Malakozoologie in der Bundesrepublik
Deutschland vom 09. bis 11. Oktober 1981 in Miinster
i.W.". (address: see Archiv fiir Molluskenkunde(
Nachrichtsblatt der Deutschen Malakozoologischen Ce-
sellschaft. [Frankfurt am Main]. 1 [Dec. 1868]-51(4)
[Oct. -Dec. 1919]. A supplement series, Beilage zum
Nachrichtsblatt, Beitrage zur Kenntniss der mitteleuro-
paischen Najadeen, 1 [1908]-4 [1910]. Continued as the
Archiv fiir Molluskenkunde. Index to volumes 1-45: Ko-
belt (1914).
Neue Litteratur und Beytrage zur KenntniB der Natur-
geschichte, vorziiglich der Conchylien und FoBilien.
[Leipzig]. 1 [1784]-4 [1787]. Continuation of FUr die
Litteratur und KenntniB der Naturgeschichte, sonderlich
der Conchylien und der Steine. Continued as Unterhal-
tungen fiir Conchylienfreunde und fiir Sammler der Mi-
neralien.
Novitates Conchologicae. Series prima. Mollusca extra-
marina. Beschreibung und Abbildung neuer oder kri-
tischer Land- und Siis.swasser Moliusken . . . (ed. L. Pfeif-
fer). [subtitles varied]. [Kassel]. 1 [1854-186G]-5 [1876-
1879]. Collation: Johnson (1969).
Novitates (Conchologicae. Abtheilung II; Meeres-Con-
chylien. Abbildung und Beschreibung neuer Conchylien
. . . (ed. W. Dunker). [subtitles varied]. [Kassel]. 1 [1858]-
16 [1878]. Supplement 1 [1862] " Monographic der Mol-
luskengattung Dosinia, Scopoli {Artemis, Poli)", E. R6-
mer. Supplement 2 [1853] "Index Molluscorum quae in
itinere ad Guineam inferiorem collegit Georgius Tams",
G. Dunker. Supplement 3 [1864-1862] "Monographic
der Molluskengattung Venus Linne", E. Romer. Sup-
plement 4 [1869-1874] "Japanische Meeres-Conchilien",
C. E. Lischke. Supplement 5 [1874] "Ueber Vorderasia-
tische Conchylien nach den Sammlungen des Prof. Haus-
knecht", E. von Martens. Supplement 6 [1877]. "Clau-
silienstudien", O. Boettger. Supplement 7 [1882] "Index
Molluscorum Maris Japonici", G. Dunker. Collation:
Johnson (1969).
Schriften zur Malakozoologie aus dem Haus der Natur.
[Cismar]. 1 [1989]-current. IHaus der Natur-Cismar,
Hinter dem Kloster 42, W-2433 Cismar, Germany!
UNITAS Malacologica Newsletter. [Tubingen]. 1 [1987]-
current. {Tropenmed. Inst., Univ. Tiibingen, Wilhelm-
straBe 31, W-7400 Tiibingen, Germanv '
Unterhahungen fiir Conchylienfreunde und fur Samm-
ler der Mineralien. [Erlangen]. 1 [1789]. Continuation
of Neue Litteratur und Beytrage zur KenntniB der Na-
turgeschichte, vorziiglich der Conchylien und FoBilien.
Zeitschrift fiir Malakozoologie. [Hannover; Kassel]. 1
[Jan. 18441-10(12) [Dec. 1853]. Continued as Malako-
zoologische Blatter.
Great Britain
The British Shell Collectors' Club Newsletter. [Not-
tingham]. 1 [1972]-14 [1983]. Continued as Pallidula.
Index, 1982, numbers 1-50 (D. Feld).
Bulletin of the Malacological Society of London. [Lon-
don]. 1 [Sept. 1983]-current. Continuation of the Mala-
cological Society of London Newsletter, {address: see
Journal of Molluscan Studies}
The Cephalopod Newsletter. [London]. 1 [1977]-current.
{Dept. of Geological Sciences, LIniversity College, Gower
Street, London WCIE 6BT, England} '
CIAC [Cephalopod International Advisory Council]
Newsletter — Cephalopod International Advisory Board.
[London; Cambridge]. 1 [1985]-current. {British Antarc-
tic Survey, NERC, High Cross, Madingley Road, Cam-
bridge CB3 OET, England}
[Conchological Leaflets] (F. P. Marrat). [Liverpool] (1)
[1876J-11 [1880/1883]. See McMillan (1961).
The Conchological Society of Great Britain and Ireland.
Papers for Students. [London]. 1 [1964J-18 [1981].
Conchological Society of Great Britain and Ireland Spe-
cial Publication. [London]. 1 [Feb. 1980] "Marine Mol-
lusca described by John Gwyn Jeffreys, with the location
of the type material", A. Waren.
The Conchologist: A quarterly Magazine for Conchol-
ogists. [London]. 1(1) [March 1891]-1(4) [Dec. 1891].
Continued as The Conchologist: A Journal of Malacology.
R. Bieler and A. R. Kabat, 1991
Page 45
Originally printed as a quarto in 58 pages; reprinted in
1903 in octavo, and repaginated with 88 pages; see Toni-
lin & Fisher (1935). Index: McClelland (1923).
The Conchologist: A [quarterly] Journal of Malacology.
[London]. 2(1) [March 1892]-2(8) [Oct. 1893], Continu-
ation of The Conchologist: A quarterly Magazine for
Conchologists. Continued as The Journal of Malacology.
Index: McClelland (1923).
The Conchologists' Newslelter. The Conchological So-
ciety of Great Britain and Ireland. [London]. 1 [1961]-
current. {address: see Journal of Conchologyl
The Cowry. [St. Austell, Cornwall; Port Macquarie (Aus-
tralia)]. 1 [1960]-2(1) [1968]. [Vol. 1 nos. 1-3 reprinted
by Philip L. Hano, New York],
Digesta Malacologia. [London]. 1 [1901]-2 [1903]. Num-
ber 1 is an index to the American Journal of Conchology
iq.v.); number 2 is an index to the malacological articles
in the Annals and Magazine of Natural History (a journal
not exclusively about mollusks).
Handlists of the Molluscan Collections in the Depart-
ment of Zoology, National Museum of Wales. Series 1,
The Melvill-Tomlin Collection. [Cardiff]. 1 [1981]-cur-
rent. | National Museum of Wales, Cathays Park, Cardiff
CFl 3NP, Wales}
Helix. Land Shell Studies Group. [Nottingham]. 1 [1981].
The Journal of Conchology. Conchological Society of
Britain and Ireland. [London]. 2(1) [Jan. 1879]-current.
Continuation of The Quarterlv Journal of Conchology.
Index [1879-1922, vols. 2-16]': McClelland (1923). (A.
Seccombe, 36a Selbourne Road, Ilford, Essex IGl 3AJ,
England 1
The Journal of Malacology. [London]. 3(1) [March 1894]-
12(4) [Dec. 1905]. Continuation of The Conchologist.
Index: McClelland (1923).
Journal of Molluscan Behaviour, The International
Journal devoted to the Behaviour of Molluscs. [East
Orkney]. 12(6):48 [1966]; only issue (a parody).
The Journal of Molluscan Studies. Malacological Soci-
ety of London. [London]. 42(1) [April 1976]-current.
Continuation of Proceedings of the Malacological Society
of London. Supplements 1 [1976], 3 [1977], 5 [1978], 6
[1978; reprinted 1988], 7 [1980], 9 [1981], 11 [1982], 15
[1985], 16 [1986], 17 [1990]: "The Prosobranch Molluscs
of Britain and Denmark, Parts 1-10", V. Fretter & A.
Graham. Supplement 2 [1977] "On the genus Tornatina
and related forms ", E. Marcus. Supplement 4 [1977] "An
annotated checklist of the western Atlantic warm water
opisthobranchs ", E. Marcus. Supplement 8 [1980] "Re-
vision of the North-East Atlantic bathyal and abyssal
Turridae (Mollusca, Gastropoda) ', P. Bouchet & A. Wa-
ren [see Bollettino Malacologico Supplemento 2, 3 for
continuation]. Supplement 10 [1982] "Systematics of the
genera of the Order Ascoglossa (Gastropoda)", E. Mar-
cus. Supplement 12 [1983] "Catalogue of Recent and
fossil Lithophaga (Bivalvia)", K. H. Kleeman. Supple-
ment 12A [1983; reprinted 1988] "Proceedings of the
Second Franco-British Symposium on Molluscs", ed. A.
Bebbington. Supplement 13 ["1983" 27 February 1984:
fide Bollettino Malacologico (1986) Supplemento 2: 561;
reprinted 1988] "A generic revision of the family Eulim-
idae (Gastropoda, Prosobranchia)", A. Waren. There was
no Supplement 14. {D. Roberts, Dept. of Biology, Queen's
Univ. of Belfast, Belfast BT7 INN, Northern Ireland)
The Littorinid Tidings. Occasional Newsletter of the
Littorinidae Research Group. [Manchester]. 1 [April
1974J-8 [June 1979].
The Malacological and Conchological Magazine. [Lon-
don]. 1 [1838]-2 [1839].
Malacological Society of London Newsletter. [London].
1 [1976]-16 [April 1983]. Issues 3-7 not numbered; issues
3-4 not dated [both 1977]. Continued by the Bulletin of
the Malacological Society of London.
*[Newsletter]. Northamptonshire Natural History So-
ciety (Conchology Section). [Northampton]. ?-current.
{The Humfrev Rooms, Castilian Terrance, Northampton
NNl ILD, England}
Pallidula. The Magazine of the British Shell Collectors'
Club. [Carlisle]. 15 [1984]-current. Continuation of The
British Shell Collectors' Club Newsletter. {12 Grainger
Road, Isleworth, Middlesex TW7 6PQ, England)
Proceedings of the Malacological Society of London.
[London]. 1(1) [Oct. 1893]-41(6) [Dec. 1975]. Continued
as The Journal of Molluscan Studies. Index [1893-1923,
vols. 1-15]: McClelland (1923).
The Quarterly Journal of Conchology. [London]. 1(1)
[Feb. 1874]-1(17) [Nov. 1878]. Continued as The Journal
of Conchology. Index: McClelland (1923).
Species Conchyliorum. [London]. 1(1) [Nov. 1830]. [Al-
though intended as an multivolume treatise, only one
number was ever published].
*Subscriber's Bulletin. The Snail Centre. [Colwyn Bay].
1 [198 ?]-current.
Tentacle. lUCN/SSC Mollusc Specialist Group. [Glas-
gow]. 1 [1989]-current. {Natural History Dept., Art Gal-
lery and Museum, Kelvingrove, Glasgrow G3 8AG, Scot-
land)
The Zoological Record. Mollusca Section. [London]. 1
[1864]-current. {Biosis, 2100 Arch Street, Philadelphia.
PA 19103-1399)
Hungary
Malakologiai Tajekoztato. Matra Miizeum. [Gyongyos].
1 [1980]-current. {Kossuth Lajos utca 40, H-3200 Gyon-
gyos, Hungary)
Soosiana. [Budapest]. 1 [1973]-current. Supplement I
["1977" 1979] "A Magyarorszagi Recens Puhatestiiek El-
terjedese" [= "Distribution of the Recent Mollusca of
Hungary "], L. Pinter. ITermeszettudomanyi Muzeum,
Baross utca 13, H-1088 Budapest, Hungary}
Page 46
THE NAUTILUS, Vol. 105. No. 2
Israel
.Argamanit. The Hebrew Newsletter. [Haifa; Jerusalem;
Kibbutz Netzer Sereni], [1970H1976]. (Continued as Le-
vantina.
Argamon. Journal of the Israel Malacological Society
[and the (Municipal) Malacological Museum, Nahariya].
[Haifa; Jerusalem; Kibbutz Netzer Sereni]. 1 [I970]-7(5)
[1981]. Continued as Argamon, Israel Journal of Mala-
cology.
Argamon. Israel Journal of Malacology. [Kibbutz Netzer
Sereni]. 7(6) [1988]-current. Continuation of Argamon,
Journal of the Israel Malacological Society. {Kibbutz
Netzer Sereni, 70-395 Israel}
Levantine. A Malacological Newsletter. [Kibbutz Netzer
Sereni]. 1 [1976J-53 [1984]. Continuation of Argamanit.
Continued as Levantina, A Journal of Malacology.
Levantina. A Journal of Malacology. [Kibbutz Netzer
Sereni]. 54 [1984]-current. Continuation of Levantina, A
Malacological Newsletter, [address: see Argamon]
Italy
Argonauta,, Journal of Malacology. Associazione Mala-
cologica Internazionale (A.M. I.). [Roma]. 1 [May 1985]-
current. Supplementos [1987-current] "Notiziario A. M.
I." [reviews and announcements of malacological serials
and books], fCasella Postale, P.O. Box 45, 00125 Acilia,
Roma, Italy]
Bollettino Malacologico. Societa Italiana di Malacologia.
[Milano]. 15(1-2) [Jan. -Feb. 1979]-current. Continuation
of Conchiglie, Notiziario Mensile della Unione Malacolo-
gica Italiana. Indice specifico generale (1965-1986), 1986.
Supplemento 1 [1985] "Revision of the Northeast Atlantic
bathyal and abyssal Mollusca, Part 2: Neogastropoda ex-
cluding Turridae", P. Bouchet & A. Waren. Supple-
mento 2 [1986] "Revision of the Northeast Atlantic bathy-
al and abyssal Mollusca, Part 3: Aclididae, Eulimidae,
Epitoniidae", P. Bouchet & A. Waren [continuation of
Journal of Molluscan Studies, Supplement 8]. Supple-
mento 14 [1979]-22 [1986] "Guida illustrata
all'identificazione delle Conchiglie del Mediterraneo",
B. Sabelli & G. Spada. Continued from the supplements
in Conchiglie. Additional miscellaneous supplements, such
as membership lists. {Acquario Civico, Viale Gadio 2,
20121 Milano, Italy}
Bulletino della Societa Malacologica Italiana. [Pisa]. 1
[18751-20[1897]. Continuation of Bulletino Malacologico
Italiano. Both reprinted by Antiquariaat Junk, Lochem,
Netherlands (1973). Index for both by Lamme (1974)
Antiquariaat Junk, Lochem, Netherlands, 53 pages.
Bulletino Malacologico Italiano. [Pisa]. 1(1) [Jan. -Feb.
1868]-7 [1874]. Continued as Bulletino della Societa Ma-
lacologica Italiana.
La Conchiglia/The Shell. Rivista [Mensile/Internaziona-
le] delle Meraviglie del Mare. [Roma]. 1 [March 1969]-
current. The issues of volume 1 and 2(1-4) were in Ital-
ian, with an English translation distributed in mimeo-
graph; number 2(5) is presumably the first "International
Edition" in Italian and English, which has continued to
date. Supplemento 1 [Nov.-Dec. 1970]-4 [Nov. -Dec.
1972] "Van Nostrand's Standard Catalog of Shells Month-
ly Supplement", R. J. L. Wagner and R. T. Abbott [de-
spite its title, these four supplements were not issued
monthly with this journal]. Supplemento [Jan. 1983]
"Indice Analitico Generale (1969-1982)" [supersedes
several previously issued indexes]. {Via C. Federici 1,
00147 Roma, Italy}
Conchiglie. Notiziario Mensile [del "Conchiglia Club"/
della] Unione Malacologica Italiana. [Milano]. 1 [Jan.
1965]-14(11-12) [Nov-Dec. 1978]. Continued as Bollet-
tino Malacologico. Supplemento 1 [1969]-5 [1975], "Ca-
talogo illustrato delle Conchiglie marine del Mediter-
raneo", F. Ghisotti & G. Melone. Supplemento 1 [1977]-
13 [1978] "Guida illustrata all'identificazione delle Con-
chiglie del Mediterraneo", B. Sabelli & G. Spada. Con-
tinued as a supplement to Bollettino Malacologico. Ad-
ditional miscellaneous supplements, including reprints
of old papers. Indice Analitico Pluriennale 1965-1969
[1970]; 1970-1976 [1977],
The Connoisseur of Seashells. [Roma]. 1 [1985]-current.
{P.O. Box 561. 00187 Roma.ltaly}
Giornale di Malacologia. [Pavia], 1 [1853]-2 [1854].
Lavori della Societa Italiana di Malacologia. [Milano].
21 ["1984" 1985]-current. Continuation of Lavori della
Societa Malacologica Italiana. {address: see Bollettino
Malacologico}
Lavori della Societa Malacologica Italiana. [Milano]. 1
[1964]-20 [1982]. Continued as Lavori della Societa It-
aliana di Malacologia. [Early numbers are collected re-
prints from various journals].
Malacofauna Italica. (C. Alzona). [Genova]. 1(1) 1937-
1(7) [1940]. Posthumous revised edition [1971], Atti della
Societa Italiana di Scienze Naturali e del Museo Civico
di Storia Naturale di Milano 111:1-433.
Malacologia Mostra Mondiale. Associazione Picena di
Malacologia. [Cupra Marittima]. 1 [1988]-current. {Via
Adriatica Nord 240, 63012 Cupra Marittima, Italy}
* Mostra Malacologica Messinese. Associazione Mala-
cologica Messinese. [Messina]. 1 [1980?]-current?
Notiziario C.LS.M.A. [Centro italiano Studi Malacolo-
gici]. [Roma]. 1 [1979]-current. {Civico Museo de Zoolo-
gia, via V. Aldrovandi 18, 00197 Roma, Italy}
Notiziario S. L M. Pubblicazione mensile edita della
Societa Italiana di Malacologia. [Milano]. 1 [1983]-cur-
rent. {address: see Bollettino Malacologico}
Schede Malacologiche del Mediterraneo. Societa Ma-
lacologica Italiana. [Milano]. 1 [ 1 964 ]-60 [1978].
Snail Farming Research. Associazione Nazionale Elici-
coltori. [Cherasco]. 1 [1986]-current. {Via Vittorio
Emanuele 103, 12062 Cherasco, Italy}
R. Bieler and A. R. Kabat, 1991
Page 47
Netherlands
Achatina. A Newsletter of African non-marine malacol-
ogy. [Tervuren (Belgium); Leiden]. 1 [1970]-13 [1985].
Basteria. Tijdschrift van de Nederlands[cli]e Malacolo-
gische Vere[e]niging. [Leiden]. 1(1) [Feb. 1936]-current.
Supplement 1 [1959] "The Netherlands as an environ-
ment for moUuscan life", ed. W. S. S. van der Feen-van
Benthem Jutting & CO. van Regteren Altena. Supple-
ment 2 [198-1] "The marine Mollusca of the Bay of Al-
geciras, Spain . . .", J. van Aartsen et al. Index & dates
of publication, volumes 1-50 (1936-1986); Ende & Bank
(1989). {National Natuurhistorisch Museum, Postbus
9517, 2300 RA Leiden, The Netherlands!
Correspondentieblad van de !\ederlands[ch]e Malaco-
logische Ver[e]eniging. [Amsterdam]. 1 [1934]-current.
Inde.xes, published as "Bijlage van het . ..". Supple-
ments [1965] "Gegevens over Mariene Gastropoden van
Curacao"; [1968] "Gegevens over de Mollusken von Cu-
racao uitgezonderd de Mariene Gastropoden", K. M. de
Jong & I. Kristensen. See also Informatieblad. {address:
see Basteria!
Informatieblad van de Nederlandse Malacologische
Vereniging. [Amsterdam]. 1 [1973]-current. Published as
a supplement ["Bijlage bij ] to the Correspondentie-
blad . . . ; numbers 2-6 also simultaneously issued as sup-
plements to De Kruekel. {address; see Basteria!
Jaarverslagen, Schelpenwerkgroep Friesland. [Leeu-
warden]. 1 [1983]-current. { Schoenmakersperk 2, 8911
EM Leeuwarden, The Netherlands}
De Kreukel, uitgave van de Malacologische Contacl-
groep Amsterdam & Omstreken. [Amsterdam]. 1 [1964]-
current. Seven series of special publications and supple-
ments; (1) [1970-1983] "Proefschriftensupplementen",
1-5, lA-lG, II-IL, 4B, 5A-5G and 5L, A. F. J. Jansen.
(2) [1983; 1988] "Jubileumnummer". (3) [1988; 1989]
"Catalogus voor de Malacofilatelist", R. V. Eijkenduijn
& C. Touw. (4) [1976] "Malacologische Opstellen". ed.
W. Backhuys. (5) [1983] "Speciaal Polyplacophora-
Nummer", H. Huneker and A. van der Sman. (6) [1984]
"Bibliographia Malacofaunae Jugoslaviae", W. J. M.
Maasen. (7) [1984] "Bibliography of the Land- and Fresh-
water Molluscs of the Near East", Z. Bar. Index, R. Bank,
1983, De Kreukel Jubilieumnummer 1963-1983, p. 29-
44. {P.O. Box 6, 1100 Diemen, The Netherlands!
The Papustyla. Manus Land Snail Society. [Lorengau,
Manus Island & Woudenberg, The Netherlands]. 1 [1988]-
current. Continuation of Tree and Land Snail Collectors
of the World Newsletter. {P.O. Box 58, 3930 EB Wou-
denberg, The Netherlands!
Porphyria. Publicatie van de Werkgroep Olividae.
[Delft]' 1 [1985]-2 [1986],
Poland
Folia Malacologia — Scientific Bulletins of Stanslaw
Staszic Academy of Mining and Metallurgy. [Krakow].
1 [1987]-current. {Inst. Geol. Sur. Miner., al. Mickiewi-
cza 30, 30-050 Krak6w, Poland}
Portugal
Amphitrile — Publica^ao da Sec^ao de Malacologia do
Centre Porlugues de Actividades Subaquaticas. [Lisboa].
1 [1979].
Halia. Sociedade Portuguesa de Malacologia. [Lisboa]. 1
[1982]-current. {address: see Publica9oes Ocasionais!-
Publica^oes Ocasionais da Sociedade Portuguesa de
Malacologia. [Lisboa]. 1 [May 1982]-current. {Largo da
Princesa 24-l°E., 1400 Lisboa, Portugal}
Spain
*Hojas Malacologicas. Coleccion de Memorias, datos y
noticias sobre los rnoluscos terrestres de Espafia y Por-
tugal. [Madrid]. 1 [1870-1875?]-2 [1870].
Iberus. Revista de la Sociedad Espanola de Malacologia.
[Barcelona]. 1 [1981]-current. Suplemento 1 [1988] "Ca-
talogo actualizado y comentado de los opistobranquios
(Mollusca, Gastropoda) de la Peninsula Iberica, Baleares
y Canarias, con algunas referencias a Cueta y La Isla de
Alboran", J. L. Cervera et al. Suplemento 2 [1990], "Des-
cripcion de nuevas especies y subespecies del genero
Conus (Mollusca, Neogastropoda) para el Archipielago
de Cabo Verde", E. Rolan. {Museo Nacional de Ciencias
Naturales, Jose Gutierrez Abascal 2, 28006 Madrid, Spain}
Resenas Malacologicas. Sociedad Espafiola de Malaco-
logia. [Madrid]. 1 [1981]-current. {address; see Iberus}
Switzerland
Bulletin de la Societe Internationale de Conchyliologie
[S.I.C.j. [Lausanne]. 1 [1979]-current. {C.P. 1323, CH-
1001 Lausanne, Switzerland!
AFRICA
South Africa
Bulletinus Natalensis. Occasional newsletter of the Dur-
ban group of the Conchological Society of Southern Af-
rica. [Durban]. 1 [1983]-current. {address; see Strandlo-
per[
Circular, The Conchological Society of Southern Af-
rica. [Cape Town]. 1 [1958]-140 [1972]. Continued as
Strandloper. Supplement 1 [1960] "Eastern Province &
Natal Supplement [to the] Circular, The Conchological
Society of Southern .Africa". [Durban].
INewsletler of the Conchological Society of Southern
Africa. [Durban]. 1 [1989]-current. {address; see Strand-
loper !
Reefcomber. [Occasional newsletter of] the Natal Shell
Society. [Durban]. 1 [1979]-20 [1984].
Slug Newsletter. [Plumstead]. [198 ?]-current. [A par-
Page 48
THE NAUTILUS, Vol. 105, No. 2
ody, by Giis Ferguson {30 Firfield Road, 7800 Plumstead,
South Africa}].
Special Publication, Conchological Society of Southern
Africa. [Kenilworth], 1 [1960]-4 [1965].
The Strandloper. Bulletin of the Conchological Society
of Southern Africa. [Durban]. 141 [1972]-current. Con-
tinuation of Circular, the (Conchological Society of
Southern Africa, i Durban Museum, P.O. Box 4085, 4000
Durban, South Africa}
ASIA
China
Musee Heude, Notes de Malacologie Chinoise. Universi-
te I'Aurore. [Shanghai]. 1 [1934-1945]. [Issue 1(5) (1938)
as "Vol. V".]
Transactions of the Chinese Society of Malacology. [Bei-
jing/Qingdao]. 1 [1983]-current. {Institute of Oceanol-
ogy, Academia Sinica, 7 Nan-Hai Road, Qingdao, China}
Japan
Bulletin of the Institute of Malacology, Tokyo. [Tokyo].
1(1) [May 1979]-current. {6-36, Midoricho-3-chome,
Tanashi City, Tokyo 188, Japan!
Bulletin of the National Pearl Research Laboratory.
[Kashikojima]. 1 [1956]-22 [1978J
Bulletin of the Nippon Institute for Scientific Research
on Pearls. [Kyoto], 1 [1953]-4 [1963]. [Collected reprints
from various journals].
The Chiribotan. Newsletter of the Malacological Society
of Japan. [TokyoJ 1 [1960]-current. [Early numbers with-
out English title], {address: see Venus}
Conchologia Asiatica. [KyotoJ 1(1) [Oct. 1943]-1(4)
[1943],
The Conchological Magazine. [KyotoJ 1 [Nov 1907]-
4(1) [1915J [Subtitles varied; some of the articles had
English translations].
*Kzohelix. Sheller's Club Ezohelix. [Horai, Wakkanai].
?-current.
Hitachiobi. [TokyoJ 1 [1974]-current.
Illustrated Catalogue of Japanese Shells. [Kyoto]. 1(1)
[1949]-1(25) [1953]; 2(1, 2) [1955]; series B, 1 [1955]-24
[1955 J
Isokojiki. [KyotoJ 1 [1970]-current.
The Japanese Journal of Malacology. 13(1-4) [Dec.
1943]-15(l-4) [March 1948J Continuation of The Venus.
Continued as Venus, The Japanese Journal of Malacol-
ogy
JECOLN Annual Report for Japanese Expert Con-
sultation on Living Nautilus. [Kamagawa]. 1976-1977
[1977]-1978-1979[1980J [Collected reprints on Nauti/us
from various journalsj
Kainakama. Hanshin Shell Club. [Nishinomiya-shi, Hy-
ogo-kenj 1(1) [1967]-current.
Material of the National Pearl Research Laboratory.
[Ka.shikojimaJ I [1970]-6 [1978J
Occasional Publication of the Institute of Malacology:
North Pacific Shells. [Tokyo]. 1 [Nov. 1979]-current.
{address: see Bulletin of the Institute of Malacology, To-
kyo}
Ostrea. [Hiroshima} 1(1) [1969]-current.
Pacific Shell News. [TokyoJ 1 [Jan. 1970]-6[Dec. 1972J
See Emerson & Sage (1987).
Proceedings of the Malacozoological Meeting, "Ha-
gure-Garasu" of Tokyo. [Tokyo]. 1 [1987]-current.
Special Publication, Institute of Malacology of Tokyo.
[Tokyo]. 1 [1985]-current. [Number 2 was on corals].
The Venus, [subtitles varied]. The Malacological Society
of Japan. [KyotoJ l(l)[Nov. 1928]-12(3-4) [March 1943J
Continued as The Japanese Journal of Malacology. Ap-
pendix [1929-1935] "An illustrated catalogue of the Jap-
anese shells", T. Kuroda.
Venus. The Japanese Journal of Malacology. [Mukaishi-
ma; Fukuyama; TokyoJ 15(5-8) [Oct. 1949]-current.
Continuation of The Japanese Journal of Malacology.
Special Publication [1979] "Check list of new genera and
species; Author index; Date of publication" [volumes 1-
36, 1928-1977]. Supplement 1 [1989] "Taxonomy and
distribution of moUusks", ed. T. Okutani. {National Sci-
ence Museum, 3-23-1 Hyakunin-cho, Shinjuku-ku, To-
kyo 169, Japan}
South Korea
The Korean Journal of Malacology. [Seoul]. 1(1) [Dec.
1985]-current. {Institute of Tropical Medicine, Yonsei
Univ., G.P.O. Box 8044, Seoul, South Korea}
Philippines
Carfel. Philippine Shell News. A publication of Carfel
Seashell Museum. [ManilaJ 1 [1979]-8(5) [Sept.-Oct.
1986J [Issue 6(2) erroneously labelled as "Vol. VII No.
2"J
* Philippine Shell News. A Publication of the Association
of Philippine Shell Collectors. [Manila]. 7 (New Series
No. 1) [1972}-?. Continuation of The Philippine Shell
News.
*The Philippine Shell News. Association of Philippine
Shell Collectors, Inc. [Quezon CityJ 1 [1958]-6 [1964?J
Continued as Philippine Shell News.
Taiwan
Bulletin of Malacology, Republic of China. [TaipeiJ 5
[Nov. 1978]-current. Continuation of Bulletin of the Mal-
acological Society of China. {Taiwan Museum, 2 Siang-
Yang Road, Taipei, Taiwan}
Bulletin of the Malacological Society of China. [TaipeiJ
1 [Aug. 1974J-4 [Nov. 1977J Continued as Bulletin of
Malacology, Republic of China.
R. Bieler and A. R. Kabat, 1991
Page 49
OCEANIA
Australia
*Argonauta. Malacological Society of South Australia,
Inc. [Adelaide]. 1 [19 ?]-current.
Australian Newsletter. Malacological Society of Austra-
lia. [Sydney]. 6(20) [Jan. 1958]-15(60) [Jan. 1968]; new
series, 1 [= 16(61)] [1968]-18 [July-Oct. 1972]. Contin-
uation of The Malacological Society of Australia News-
letter. Continued as Australian Shell News.
Australian Shell News. Newsletter of the Malacological
Society of Australia. [Sydney]. 1 [Jan. 1973]-current. Con-
tinuation of Australian Newsletter. Supplement to No.
19[1977] "Family Mitridae Sub-family Mitrinae in West-
ern Australia", G. Hanson. Supplement to No. 33 [1981]
"Check Lists of the Muricidae of Queensland", O. H.
Rippingale. Additional miscellaneous supplements, pri-
marily membership lists, {address: see Journal of the
Malacological Society of Australia!
Brisbane Branch Newsletter, IVlalaeological Society of
Australia. [Brisbane, Queensland]. [Feb.-March 1969]-
[Dec. 1983]. [Issues not numbered sequentially]. Contin-
ued as the Moreton Bay Radula.
Cairns Shell News. Cairns Shell Club. [Cairns, Queens-
land]. 1 [Sept. 1977]-current.
Con's Conchology Newsletter of the Royal Zoological
Society of New South Wales Conchological Section.
[Sydney]. 1(1) [June-July 1979]-[1983]. Continued as The
Sydney Sheller.
The Cowry. [St. Austell (Great Britain); Port Macquarie,
New South Wales]. 2(1) [1968]. [Vol. 1 published in Great
Britain].
Field Naturalists' Section of the Royal Society of South
Australia Conchology Club (B. C. Cotton). [Adelaide]
4 [June 1947]-5 [Jan. 1948]. Continuation of Malacolog-
ical Society of South Australia Publications.
Journal de Conchyliologie. [Mosman, New South Wales].
108(1) [1970]-115(4) [December 1979]. [Previously pub-
lished in Paris; see entry under "France" for further
information].
Journal of the IVlalaeological Society of Australia. [Perth,
Western Australia]. 1(1) [Sept. 1957]-current. Index issue
(1-10), 1969. IDept. of Invertebrates, Museum of Vic-
toria, 285 Russell Street, Melbourne, Victoria 3000, Aus-
tralia}
Keppel Bay Tidings. Keppel Bay Shell Club. [Yeppoon,
Queensland], 1 [1962]-current.
Malacological Club of Victoria. [Letter-Bulletin; News-
letter]. [Melbourne]. 1 [1953]-4(14) [June 1956]. Contin-
ued as The Malacological Society of Australia Newsletter.
IVlalaeological Club of Victoria Publication (B. C. Cot-
ton). [Melbourne]. 1 [July 1956]-2[Oct. 1956]. Continued
as Malacological Society of Australia, Publication.
The Malacological Society of Australia Newsletter.
[Sydney]. 4(15) [Oct. 1956]-5(19) [1957]. Continuation of
Malacological Club of Victoria Newsletter. Continued as
Australian Newsletter.
The Malacological Society of Australia Publication (B.
C. Cotton). [Sydney]. 3 [March 1957]-4 [April 1957].
Continuation of Malacological Club of Victoria Publi-
cation.
Malacological Society of South Australia Publication
(B. C. Cotton). [Adelaide]. 1 [Dec. 1938J-3 [Jan. 1943].
[Note; No. 3 was not on Mollusca]. Continued as Field
Naturalists' Section of the Royal Society of South Aus-
tralia Conchology Club.
The Moreton Bay Radula. Brisbane Branch of the Mal-
acological Society of Australia. [South Brisbane, Queens-
land]. 1(1) [Feb. 1984]-current. Continuation of the Bris-
bane Branch Newsletter.
Pearl Oyster Information Bulletin. Pearl Oyster Special
Interest Group. South Pacific Commission/Pacific Islands
Marine Resources Information System [PIMRIS]. 1 [Feb.
1990]-current. {13 Toxteth Avenue, Austimmer, New
South Wales 2514, Australia!
Proceedings of the Miriam Vale Shell Club. [Miriam
Vale, Queensland]. [Oct. 1955]-[April 1959].
*Proserpine Shell Club Newsletter. [Proserpine,
Queensland]. ?-current.
Royal Society of South Australia, Malacological Sec-
tion, Publication. [Adelaide]. 1 [July 1952J-16 [1961].
Several numbers revised and reissued, not always dated.
*Sunshine Coast Shell Club News. [Golden Beach,
Queensland]. ?-current.
The Sydney Sheller. Royal Zoological Society of NSW
Conchology Section. [Sydney]. [1983]-current. Contin-
uation of Con's Conchology.
[Newsletter]. The Townsville Shell Club. [Townsville,
Queensland]. (1) [March 1983]-(14) [1987]. Continued as
Tide Watch.
Tide Watch. The Townsville Shell Club. [Townsville,
Queensland]. (15) [Aug. -Oct. 1987]-current. Continua-
tion of the newsletter of The Townsville Shell Club.
Victorian Branch Bulletin. Victorian Branch, Malaco-
logical Society of Australia. [Reservoir, Victoria]. 1 [Aug.
1968]-current.
[The] W. A. Shell Collector. Western Australian Shell
Club. [Perth]. [1966]-current. [Originally unnumbered;
Number 1 issued in December 1973]. Also a WA Branch
Newsletter for members.
Fiji
Fiji Shell Club Annual Magazine. [Suva]. [1979]-[1987].
Shell Gossip. A monthly newsletter issued to members
of the Fiji Shell Club. [Suva]. 1 [1960J-13 [1972]; new
series 1 [1972]-5 [1976],
Page 50
THE NAUTILUS, Vol. 105, No. 2
French Polynesia
Robusta. Bulletin de liaison de 1 '.Association Fran^aise
de Conchyliologie, section de Tahiti. [Papeete]. 1 [Jan.-
Feb. 1990}-current. {B.P. 12030, Papara, Tahiti}
La Sleeriae. Bulletin de liai.son du club des collection-
neurs de coquillages de Tahiti. [Papeete]. 1 [1976]-16
[1977].
New Caledonia
Rossiniana. Bulletin de I'Association Conchyliologique
de Nouvelle Caledonie [N.C.]. [Noumea]. 1 [1978]-cur-
rent. |B.P. 148, 18 rue Henri-Bonneaud, Noumea, New
Caledonia}
New Zealand
Bulletin, [Auckland Museum Conchology Club/Con-
chology Section of the Auckland (Institute and) Mu-
seum]. [Auckland]. 1 [Nov. 1940]-17 [1961]; new series
1 [1976]-2 [1977].
Cookia. Journal of the Wellington Shell Club, New Zea-
land. [Wellington]. 1 [1976]-current. Continuation of
Wellington Shell Club Newsletter.
News Letter [Conchology Section Auckland Institute
and Museum/Auckland Museum Conchology Section).
[Auckland]. [1962]-[1982]; new series, 1 [19831-current.
The new series is issued as a supplement to Poirieria.
Poirieria. [Conchology Section, Auckland Institute and
Museum/Auckland Museum Conchology Section].
[Auckland]. 1 [1962]-current. Special Issue 1 [1980].
Tidesong. Christchurch Shell Club. [Christchurch]. 1
[Dec. 1970]-current. Also a separate Newsletter on club
matters.
Wellington Shell Club [Newsletter/Bulletin]. [Welling-
ton]. 1 [Feb. 1967]-36 [1975]. Continued as Cookia.
Whangarei Shell Club Newsletter. [Whangarei]. [Oct.
1978]-current.
Journal of Cephalopod Biology. [St. John's, Newfound-
land]. 1 [1989 [-current. {Ocean Studies, Memorial Univ.
of Newfoundland, St. John's, Newfoundland, Canada AlC
5S7}
Molluscs in Archaeology & the Recent. [Vancouver,
British Columbia.] 1 [Aug. 1960]-7 [Aug. 1962].
Out of the Shell. IRDC [International Research and De-
velopment Centre] Mollusc Culture Network. [Halifa.\,
Nova Scotia]. 1 [Sept. 1989]-current. {Mollusc Culture
Network, IRDC, Dalhousie University, Halifa.x, Nova
Scotia, Canada B3H 4J II
*The Vancouver and District Shell Bulletin. [Vancou-
ver, British Columbia]. 1 [Jan. 1979]-current?
Cuba
Actas de la Sociedad Malacologica "Carlos de la Tor-
re", Durante el afio [HabanaJ 1942 [1943]-1944
[1945].
*Boletin del Grupo de Estudios Malacologicos "Rafael
Arango y Molina". 2 [1980]-current? Continuation of
the Contribuciones Malacologicas del Grupo de Estudios
Malacologicos "Rafael Arango y Molina ".
Circulares de la Sociedad Malacologica "Carlos de la
Torre". [Habana]. 1 [1947-1950J Continued as Circu-
lares del Museo y Biblioteca de Malacologia de La Ha-
bana.
Circulares del Museo y Biblioteca de Malacologia de
La Habana. [HabanaJ 2 [1950J-4 [1954]. Continuation
of Circulares de la Sociedad Malacologica "Carlos de la
Torre ".
Contribuciones Malacologicas del Grupo de Estudios
Malacologicos "Rafael Arango y Molina". [Habana]. 1
[1980]. Continued as the Boletin del Grupo de Estudios
Malacologicos "Rafael Arango y Molina ".
Revista de la Sociedad Malacologica "Carlos de la Tor-
re". [Habana]. 1(1) [May 1943]-9(2) [Jan. 1954]. Index:
Jacobson (1971).
Papua New Guinea
The Papustyla. Manus Land Snail Society. [Lorengau,
Manus Island & Woudenberg, The Netherlands]. 1 [1988]-
current. Continuation of Tree and Land Snail Collectors
of the World Newsletter. {P.O. Box 58, 3930 EB Wou-
denberg, The Netherlands}
Tree and Land Snail Collectors of the World Newslet-
ter. [LorengauJ 1 [1987-1988]. Continued as The Pa-
pustyla.
NORTH AMERICA
Canada
Conchologica Nonsensica. [Toronto, Ontario]. 1 [1939].
[A parody, by John Oughton].
United States of America
American Conchologist. Quarterly Bulletin of the Con-
chologists of America, Inc. [Louisville, Kentucky]. 15(2)
[1987]-current. Continuation of Conchologists of Amer-
ica Bulletin. {P.O. Box 8105, Saddle Brook, NJ 07026-
8105, USA}
American Journal of Conchology. [Philadelphia, Penn-
sylvania]. 1[1865]-7[1872J Indexes: Sykes (1901) Digesta
Malacologica, Lviii, 46; La Rocque (1961); see also Ab-
bott (1962). The dates of publication were printed on
the verso of the title page of each volume and compiled
by Sykes (1901).
American Malacological Bulletin. [Hattiesburg, Missis-
sippi; Indiana, Penns>lvania]. 1 [1983]-current. [Vol. 3(1),
"December 1984" published March 1985]. Special Edi-
tion 1 [1985] "Perspectives in Malacology". Special Edi-
R. Bieler and A. R. Kabat, 1991
Page 51
tioii 2 [1986] "Proceedings of the second international
Corbicula symposium, ed. J. C. Britten. Special Edition
3 [1986] "Proceedings of the symposium on the entrain-
ment of larval oysters". Continuation of Bulletin of the
American Malacological Union for Index, 198.'3-1988
(vols. 1-6), (1989) American Malacological Bulletin 6(2);
219-305, C. L. Counts. % Richard E. Petit, P.O. Bo.x 30,
North Mvrtle Beach, SC 29582, LISA.
The American Malacological Union, Annual Meeting
, Membership List .... [Philadelphia, Pennsylvania].
(5) [1936]-(10) [1942]. Continuation of Report, _ Annual
Meeting and Membership, The American Malacological
Union. Continued as The American Malacological Union,
News Bulletin and Annual Report,
The American Malacological Union [, Inc.] [and] Amer-
ican Malacological Union [, Inc.], Pacific Division, An-
nual Report[s] [for] _ (20) ["1953" 1954]-37 ["1970"
1971]. [Various places of publication]. Vol-
umes 22-37 officially assigned bulletin numbers. Contin-
uation of The American Malacological Union, News Bul-
letin and Annual Report Continued as The American
Malacological Union, Inc., Bulletin for
The American Malacological Union, Inc., Bulletin for
(38) ["1971" 1972]-(40) ["1973" 1974]. [Phil-
adelphia, Pennsylvania], Variant titles: inside front cover
as "The AMU Bulletin"; the header on articles as "Bul-
letin of the American Malacological Union, Inc., for ".
Continuation of The American Malacological Union . . . ,
Annual Reports for Continued as Bulletin of the
American Malacological Union, Inc.
The American Malacological Union, News Bulletin and
Annual Report, , Membership list. . . . [Philadelphia,
Pennsylvania]. (11) ["1943" 1944]-(19) ["1952" 1953].
Continuation of The American Malacological Union,
.\nnual Meeting Membership List .... Continued as
The American Malacological Union .. . , Annual Reports
for_.
American Malacological Union [, Inc.,] Newsletter.
[Various places of publication]. 1 [1968]-15(1) [1984].
[Also titled "Newsletter of the . . ." (1971)]. Continued
as AMU News.
Amici Castropodorum. Friends of the Gastropods News-
letter. [Various places; currently at Tampa, Florida]. 1
[1976]-current. [Variously subtitled; early issues not
numbered]. {Dept. of Geology, Univ. of South Florida,
Tampa, FL 33260, USA \
AMU News. [Publication of the American Malacological
Union]. [Ft. Pierce, Florida; Princess Anne, Maryland].
15(2) [1984]-current. Continuation of .\merican Mal-
acological Union [, Inc.,] Newsletter, {address: see Amer-
ican Malacological Bulletin r
*The Auger. The Austin Shell Club. [Pflugerville, Texas].
?-current?
The Beachcomber Report. Coastal Bend Shell Club.
[Corpus Christi, Texas]. [April 1965]-[Nov. 1980]. [Un-
numbered issues; those of Aug. 1969 to Sept. 1970 (or
Dec. 1970?) were published in "Star to Star", a magazine
of the Corpus Christi Junior Museum]. Continued as The
Mitchelli.
* Between the Tides. Gulf Coast Shell Club. [Beaumont,
Texas]. ?-? [Seen: 1963-1964, with some early issues
numbered {e.g., "Edition No. IX" = January 1964); 1965-
June 1970; issued monthly & not numbered]. Continued
(?) as the Shell and Tell.
Boston Malacological Club, Inc., Newsletter. [Various
places]. 1 [1969-1987]. Continued as The Epitonium.
Bulletin of Paleomalacology. An International Forum
for the Study of Systematics, Ecology, and Biogeography
of Worldwide Fossil Mollusks. [Charlottesville, Virginia].
1(1) [Feb. 1988]-1(4) [Dec. 1988].
Bulletin of the American Malacological Union [, Inc.]
[for]. (41) ["1974" 1975]-(4S) ["1981" 1982; 5 page er-
rata, 1983]. [Philadelphia, Pennsylvania]. Inside front
cover, title as "The AMU Bulletin". Continuation of The
American Malacological Union, Inc., Bulletin for
Continued as American Malacological Bulletin. Index to
years 1934-1974, (1975), 57 p. See Burch (1981) for dates
and contents of issues from 1931-1966; note that several
dates are incorrect therein.
Bulletin of the Brooklyn Conchological Club. [Brook-
lyn, New York]. 1 [1907].
The Busycon. Broward Shell Club. [Pompano Beach,
Florida]. 10[1975]-current. Continuation of Busycon Bu-
gle.
Busycon Bugle. Publication of the Broward Shell Club.
[Fort Lauderdale, FloridaJ [May, 1962]-9(8) [August
1974]. [Early issues not numbered]. Continued as The
Busycon.
The Capsule. The .Astronaut Trail Shell Club of Brevard,
Inc. [Melbourne, Florida]. 1 [Sept. 1967]-current.
Central Florida Shell News. Central Florida Shell Club.
[Orlando, Florida]. [1961]-current. [Issues numbered
starting with 1(1), Sept. 1980]. See also Clam Chatter.
The Chambered Nautilus Newsletter. [Wilmington, Del-
aware; Quincy, Massachusetts]. 1 [1974]-current.
Chicago Shell Club Newsletter. [Chicago, Illinois]. 1
[1966]-19 [1984]. Continued as Thatcheria.
Clam Chatter. [Orlando, Florida]. 1(2) [1980J A tem-
porary variant title of Central Florida Shell News.
The Collector. A Publication of the Sacramento Valley
Conchological Societv. [Sacramento, CaliforniaJ 1(1) [Jan.
1955]-8(4) [Dec. 1962J
*Conch Courier. North Texas Conchological Society.
[Piano, Texas]. ?-current. [Earliest issue seen: January
1985].
Las Conchas. Meeting Notice and Newsletter of Pacific
Shell Club, Inc. [Downey, California]. 1 [1969}-curreiit.
Continuation, in part, of News of the Western Association
of Shell Clubs.
♦Conchological Memoranda. [San Francisco?]. 1 [1866]-
4 [1868/9?]; 6 [1871]-13 [1875]; number 5 was not pub-
Page 52
THE NAUTILUS, Vol. 105, No. 2
lished. Reprinted, in part, by Smith (1976) [but note that
page 2 of number 7 was accidentally reprinted as page
2 of number 8].
Conchological INotes for [month, year]. Conchological
Club of Southern (California. [Los Angeles]. [April, 1939]-
[June 1941]. Issued monthly; not numbered. Continued
as Minutes of the Conchological Club of Southern Cal-
ifornia.
The Conchologist. Isaac Lea Chapter of Conchology
(No. 119) of the Agassiz Association. [Alameda, Califor-
nia]. 1(1-3) [January-March, 1901]. Continuation of the
Transactions of the Isaac Lea Conchological Chapter of
the Agassiz Association. See Roth & Carlton (1970).
Conchologist. [Houston, Texas]. 1(2) [Oct. 1964]-1(3)
[Nov. 1964]. Continuation of the Publication of the Con-
chology Group of the Outdoor Nature Club, Houston,
Texas. Continued as the Texas Conchologist.
The Conehologist's Exchange. [Philadelphia, Pennsyl-
vania]. 1(1) [1886]-2(9) [March-April 1988]. Continued
as The Nautilus [for index, see that entry].
Conchoiogists of America Bulletin. 1 [1974]-15(1) [1987]
[\ arious places of publication]. Continued as American
Conchologist. [Note: in volume 1, there were two dif-
ferent issues of "Number 7"; one "Number 8" and no
"Number 9"; fide American Conchologist (June 1989)
17(2): 11}
Cone Shell Alert (John K. Tucker). [Effingham, Illinois].
1(1) [Oct. 1980]-1(2) [Nov. 1980].
Contributions to Conchology (C. B. Adams; E. Chitty).
[New York; Kingston, Jamaica]. 1 [1849]-13 [1853].
Convention Papers, National Shellfisheries Association.
[Washington, D.C.]. (23) [1931 "1930"]-(44) [1953]. Con-
tinued as Proceedings of the National Shellfisheries As-
sociation. The numbers of 1931-1950 were printed on
"micro cards"; 1950-1953 were issued in mimeographed
form. "Volumes 1 (1909)-22 (1929)" do not exist, except
as meeting handouts. See Malacological Review (1970
"1969"), 2:153, footnote 4 regarding these complexities.
Corbicula Communications. [Cincinnati, Ohio]. 1 [1970]-
2 [1971].
Corbicula Newsletter. [Oak Ridge, Tennessee; Fort
Worth, Texas]. 1(1) [August 1975]-9(1) [May 1985].
Crown Crier. St. Petersburg Shell Club. [St. Petersburg,
Florida]. [1983]-[1988]. [Bimonthly report]. Combined
with the Tidelines in 1988.
The Double Helix (A Guide to the Konks of A.M.L.).
Unofficial Organ of the American Mal-ecological Union-
idae. Published under the auspices of the [New Orleans/
Deep South] Shell Club, New Orleans [La. /Louisiana].
[New Orleans, Louisiana]. 1(1) [July 1968]; 2(1) [July
1969]; 7(1) [July 1971]; possibly there were other issues.
[A parody, by D. Dundee].
Dredgings. Pacific Northwest Shell Club, Inc. [Seattle,
Washington]. 14(1) [Feb. 1974]-current. Continuation of
Pacific Northwest Shell News.
The Echo. Abstracts and Proceedings of the Annual
Meeting of the Western Society of Malacologists. [San
Francisco]. 1 [1968]-6 [1974]. Continued as Western So-
ciety of Malacologists Annual Report.
The Epitonium. Newsletter of the Boston Malacological
Club, Inc. [Newton, Massachusetts]. 1987-current. Con-
tinuation of Boston Malacological Club, Inc., Newsletter.
The Festivus. [A publication of the] San Diego Shell Club.
[San Diego, California]. 1(1) [Jan. 1969]-current. Sup-
plement [1984] "Illustration of the types named by S.
Stillman Berry in his 'Leaflets in Malacology' ", C. M.
Hertz. Supplement [1987] "A faunal study of the bivalves
of San Felipe and environs, Gulf of California . . .", J.
Gemmell et al. Supplement [1988] "An illustrated cat-
alogue of the family Typhidae Cossmann, 1903", A.
D'Attilio & C. M. Hertz. Author Index, volumes 1-18
(1987), 10 pp., by T. H. Fulsbv. Continuation, in part,
of News of the Western Association of Shell Clubs. {3883
Mt. Blackburn Avenue, San Diego, CA 92111, USA}
*Florida Keys Shell Club. [Key West, Florida]. 1(1)
[1980]-current?
*Fort Myers Shell Club Newsletter. [Fort Myers, Flor-
ida]. ?-? [Seen: 4, 1968; 12, 1977].
Friends of the Gastropods [Club] Newsletter. See Amici
Gastropodorum.
*GaIveston Shell Club [Newsletter/Quarterly]. [Galves-
ton, Texas]. ?-? [Issues seen: no. 58, May 1971; no. 87,
Dec. 1973; vol. 9 no. 1, Jan. 1974; vol. 9 no. 12, Dec.
1974; Winter 1976].
Gastropodia (Glenn Webb). [Everman. Texas; Kutz-
town, Pennsylvania; Fleetwood, Pennsylvania]. 1(1) [Oct.
1952]-current. [Page 21 issue 2(3) "reprinted" 22 May
1984; but the original not published until 12 March 1990].
{Route 1, Box 158, Fleetwood, PA 19522, USA}
Grand Strand Shell Club Newsletter. [Surfside Beach,
South Carolina]. 1(1) [March 1990]-2(1) [August 1990].
Continued as Grand Strand Shell News.
Grand Strand Shell News. Grand Strand Shell Club.
[Surfside Beach, South Carolina]. 2(2) [October, 1990J-
current. Continuation of Grand Strand Shell Club News-
letter.
*The Greater Tampa Shell Club, Newsletter. [Tampa,
Florida]. 1 [1976]-? [Last seen April 1979]. Continued,
in part, as Tampa Drill.
Guam Shell News. [Agana, Guam]. 1(1) [Jan.-Feb. 1979]-
1(4) [Dec. 1979].
Hawaiian Marine Mollusks. Recent [and New] Species
(C. S. Weaver). [Honolulu, Hawaii] 1 [1960]-2 [1966].
The Hawaiian Malacological Society [newsletter]. [Ho-
nolulu, Hawaii]. 1(1) [March 1952]-1(8) [Oct. 1952]. Con-
tinued as Hawaiian Shell News.
Hawaiian Shell News. Hawaiian Malacological Society.
[Honolulu, Hawaii]. 2(l)[Nov. 1953]-current. In addition
to volume/number, the issues are numbered consecu-
tively from number 1 of the "new series" [= 8(1)]. Sup-
R Bieler and A. R. Kabat, 1991
Page 53
plement 1 [1990] "Hawaiian Epitoniidae", H. DuShane.
Indexes, 1976-1981 (vols. 24-29), 1975-1984 (vols. 23-
32), 1985-1987 (vol.s 33-35), and 1985-1989 (vols. 33-
37), by Stuart Lillico, Hawaiian Malacological Society.
Continuation of The Hawaiian Malacological Society
[newsletter]. IP.O. Bo,\ 22130, Honolulu, HI 96823-2130,
USA}
Indo-Pacific IVIoIlusca. [Philadelphia, Pennsylvania;
Greenville, Delaware]. 1(1) [March 1959]-3(17) [Sept.
1976]. Dual pagination scheme utilized.
Irradians. [Baldwin, New York]. 7 [1980]-current. Con-
tinuation of Long Island Shell Club News.
The Jersey Sheller. Garden State Shell Club. [Newark,
New Jersey]. 1 [1966]-2 [1967].
Johnsonia. Monographs of the marine Mollusks of the
Western Atlantic. [Cambridge, Massachusetts]. 1(1) [Oct.
1941]-5(50) [July 1974]. The issues of volume 1 were
individually paginated. Volume 1 reprinted 1968; vol-
ume 2 reprinted 1975.
Journal for IVIedical and Applied Malacology/Revista
de Malacologia Medica y Aplicada. International Society
for Medical and Applied Malacology/Sociedad Inter-
nacional de Malacologia Medica y Aplicada. [Ann Arbor,
Michigan]. 1 [1989]-current. Supplement/Suplemento 1
[1989], "Proceedings of the First International Congress
on Medical and Applied Malacology/Actas del Primer
Congreso Internacional de Malacologia Medica y Apli-
cada", ed. J. B. Burch et al. Also a biannual newsletter.
{P.O. Box 2715, Ann Arbor, MI 48106, USA|
Journal of Shellfish Research. National Shellfisheries
Association. [Various places; now at West Boothbay Har-
bor, Maine]. 1 [1981]-current. Continuation of Proceed-
ings of the National Shellfisheries Association. 1 Dr. Tom
Soniat, Dept. of Biology, Univ. of New Orleans, New-
Orleans, LA 70148, USA}
*Junonia. Sanibel-Captiva Shell Club. [Sanibel, Florida].
?-current. Continuation of Sanibel-Captiva Shell Club
Newsletter.
The Laddergram. Palmetto Shell Club, Inc. [Columbia,
South Carolina]. 1 [Sept. 1987]-current.
Leaflets in Malacology. [Redlands, California]. 1(1) [Nov.
1946]-1(26) [Dec. 1969]. Illustrations of type specimens.
Hertz (1984); see Petit (1988) regarding two versions of
number 13 [printings of 7 and 9 July, 1956 of which the
former was suppressed].
The Littorina. The Louisville Conchological Society.
[Louisville, Kentucky]. 1 [1975]-ll(l)[Jan.-March 1986J
Long Island Shell Club News. [Flushing, New York;
Baldwin, New York]. 1 [1975]-6 [1980J Continued as
Irradians.
Louisiana Malacological Society Occasional Papers.
[Baton Rouge, Louisiana]. 1 [1988].
Low Country Shell Club Newsletter. [Charleston, South
Carolina! 1(1) [May, 1979]-2(7) [Sept. 1980} [Volume 1
comprised numbers 1-9 and 12; there were no numbers
10 or llj
Malacologia. [Ann Arbor, Michigan; Philadelphia, Penn-
sylvania]. 1(1) [Nov. 1962]-current. Dates of publication
given on the inside back cover of 11(2) and with the
index of later volumes. History: Burch & Huber (1968);
Burch (1973). I Malacology Department, The Academy
of Natural Sciences, 19th & The Parkway, Philadelphia,
PA 19103, USAl
Malacological Review. [Ann Arbor, Michigan; Boulder,
Colorado]. 1 [June 1969 "1968"]-current. Dates of pub-
lication are given on page ii of the latest volume. Sup-
plement 1 [1973] "The freshwater snails of Taiwan (For-
mosa)", G. L. Pace. 2 [1980] "The Meking Schistosome",
ed. J. I. Bruce et al. Supplement 3 [1982] "A dissection
guide, field and laboratory manual for the introduced
bivalve Corbicula fluminea" , J. C. Britton & B. Morton.
4 [1988] "Prosobranch Phylogeny", ed. W. F. Ponder et
al. History: Burch (1974 "1973"). {P.O. Box 637, Niwot,
CO 80544, USA}
Malacology Data Net [Ecosearch Series]. [Portland, Tex-
as]. 1(1) [April 1986]-current. [The cover of issue 2(1/2)
was erroneously labelled as "Vol. 2 No. 1"]. {325 E.
Bay view, Portland, TX 78374, USA}
Marginella Marginalia. [Dayton, Ohio]. 1(1) [March
1986}-current. {Dayton Museum of Natural History, 2629
Ridge Avenue, Dayton, OH 45414, USA!
Members of the American Malacological Union. [Phil-
adelphia, Pennsylvania]. (1) [1932]. Continued as Mrs.
Imogene C, Robertson's rambling notes. [An account of
the first meeting appeared in the Nautilus, 45(l):l-5
(1931).]
Miami Malacological Society Quarterly. [Miami, Flor-
ida]. 1(2) [1967]-5(4) [1972J Continuation of Quarterly
Report Miami Malacological Society.
Minutes [number, date] Conchological Club of Southern
California. [Los Angeles]. 1 [July, 1941]-200[June, 1960J
Continuation of Conchological Notes for ... . See Coan
(1976) regarding status of the new taxa proposed in this
serial.
The Mitchelli. Coastal Bend Shell Club. [Corpus Christi,
Texas]. 1(1) [Dec. 1980]-current. Continuation of the
Beachcomber Report.
Mollusca. [Tavares, Florida]. l(l)[Sept. 1944]-2(6) [Aug.
1949].
The Mollusca. Off the Beach Notes. Shell Club of the
Ryukyu Islands. [Ryukvu/APO San Francisco]. 1(1)
[1960]-2(8) [Dec. 1961]. [Although from Japan, this En-
glish-language newsletter was published by U.S. Armed
Services personnel and their families stationed in Ryu-
kyu].
Molluscan Digest. The International Publication for
Malacological Research. [Pismo Beach, California; Ox-
nard, California} 1(1) [Jan. 1971]-4(l-2) [Sept. 1976].
The Mollusk. Greater Miami Shell Club [previously as:
South Florida Shell Club]. [Miami, Florida]. 14(7) [1976]-
current. Continuation of South Florida Shell Club Mol-
lusk Chaser.
?a«;e 54
THE NAUTILUS, Vol. 105, No. 2
Mollusk-Chaser South Florida Shell Club. [Miami, Flor-
ida]. 5(4) [April 1967]-7(7) [July 1969]. Continuation of
The South Florida Shell Club's Newsletter Mollusk-
Chaser. Continued as the South Florida Shell Club Mol-
lusk Chaser.
IVIonographs of Marine Mollusca. [Greenville, Dela-
ware; Melbourne, Florida; Silver Springs, Maryland]. 1
[Dec. 1978]-current. [Early numbers (1-3) followed the
dual pagination scheme of the Indo-Pacific Mollusca and
the Standard Catalogue of Shells]. ! P.O. Box 7279, Silver
Spring, MD 20907-7279, USA}
Mrs. Imogene C. Robertson's Rambling Notes on the
Second Annual Meeting of the American Malacological
Union in Washington, D.C. May 26-28, 1932. [Buffalo,
New York]. (2) [1932]. Continuation of Members of the
American Malacological Union. Continued as Report,
Annual Meeting and Membership, The American Mal-
acological Union. [See also the Nautilus, 46(1): 1-3 (1932);
an account of the third meeting appeared in the Nautilus,
47(l):37-44 (1933)].
Naples Shell News. Naples Shell Club, Inc. [Naples, Flor-
ida]. 1 [1966]-current. [Sporadic numbering of volumes
and issues; "The Shell Case " a temporary variant title,
q.v.].
*I\ational Shellfisheries Association Quarterly News-
letter. [Port Norris, New Jersey]. ? [October 1989]-cur-
rent.
The Nautilus. [Philadelphia, Pennsylvania; Greenville,
Delaware; Melbourne, Florida; Silver Spring, Maryland].
3(1) [May 1889]-current. [Volumes 3-30 with 12 num-
bers each; subsequent volumes with four numbers each].
Supplement 1 [ 1990] "Catalogue of the Superfamily Can-
cellarioidea Forbes and Hanley, 1851 (Gastropoda: Pros-
obranchia", R. E. Petit & M. G. Harasewych. Continu-
ation of the Conchologist's E.xchange. Indexes: (1) "Index
to The Nautilus, vols. 3 to 34, 1889-1921 and to its
predecessor. The Conchologist's Exchange, vols. 1, 2,
1886-1888" by J. B. Hender.-on et ai, 1927, 434 p. (pub-
lished by G. M. Clapp & B. Walker). (2) "Index to The
Nautilus, volumes 35 to 60, 1921-1947" by A. LaRocque
et ai, 1951, 332 p. (published by Univ. of Michigan Press,
Ann Arbor). (3) "Author Index to The Nautilus, vols. 3-
75 ..." by A. LaRocque, 1963, 279 p. (Columbus, Ohio).
(4) "Author Index to The Nautilus, vols. 76-85" by A.
LaRocque, 1972, Sterkiana 47:21-38. (5) Inde.xes to The
Nautilus: Geographical (vols. 1-90) and scientific names
(vols. 61-90)" by R. T. Abbott et al.. 1979, iv + 238 p.
(American Malacologists, Inc., Melbourne, Florida). An
anthology: Abbott (1976). Dating of the first 75 volumes
best accomplished by reference to Clench & Turner
(1962). {P.O. Box 7279, Silver Spring, MD 20907-7279,
USA}
NCMC Newsletter. Northern California Malacozoologi-
cal Club. [Novato]. 1 [1988]-current. Continuation of the
Northern California Shell C:lub News.
[Newsletter]. National Capital Shell Club. [Washington,
D.C.]. 1 [1974)-current.
[Newsletter]. New Jersey Shell Club [May wood. New [
Jersey; Park Ridge, New Jersey]. 1973 [October]-1977
[May /June].
Newsletter, Marco Island Shell Club, Inc. [Marco Island,
Florida]. 1 [1982]-current.
The [month] Newsletter of the Jersey Cape Shell Club
[Stone Harbor, New Jersey]. [May, 1982]-current.
News of the Associated Shell Clubs of California.
[Northern California Malacozoological Club, Berkeley;
Pacific Shell Club, Los Angeles; Conchological Club of
Southern California, Los Angeles]. 1(1) [May 1960]-2(1)
[July 1961]. Continuation, in part, of the Proceedings
[Minutes] of the Pacific Shell Club. Continued as News
of the Western Association of Shell Clubs.
News of the Western Association of Shell Clubs. [As
above, with the San Diego Shell Club, San Diego]. 2(2)
[Oct. 1961]-10(4) [1969]. [Issue 7(1) (1966) erroneously
labelled as "Vol. 6 No. 7 "]. Continuation of News of the
Associated Shell Clubs of California. Continued, in part,
by The Festivus and Las Conchas.
New York Shell Club Notes. [New York]. 1 [1950]-cur-
rent. [Department of Invertebrates. American Museum
of Natural History, Central Park West at 79th Street,
New York, NY 10024, USA}
The Noble Pen. Newsletter of the Cen.[tral]
Penn.[sylvania] Beachcombers. [Elizabethtown, Pennsyl-
vania; Lancaster, Pennsylvania]. 1(1) [Nov. 1983]-cur-
rent. [The first two issues were untitled].
North Carolina Shell Club Bulletin. [Morehead City,
North Carolina]. 1 [1963J-9 [1978].
Northern California Shell Club News. Northern Cali-
fornia Malacozoological Club. [Novato]. 1 [1978]-10
[1987]. Continued as NCMC Newsletter. Note: The Ve-
liger, volumes 1-6 [1958-1963], was originally published
as the newsletter of this club.
Northwest Shell News. [Poulsbo, Washington]. 1(1) [Jan.
1961]-1(5) [Oct. 1961]. Continued as Pacific Northwest
Shell News.
Notes and Quotes. Rochester Shell and Shore Club.
[Rochester, New York]. 1 [.\ugust 1967]-66 [November
1980].
Occasional Papers on Mollusks. Department of Mol-
lusks, Museum of Comparative Zoology, Harvard Uni-
versity. [Cambridge, Massachusetts]. 1(1) [Feb. 1945]-
current. {Museum of Comparative Zoology, Harvard
Universit\. Cambridge, MA 02138, USA}
Of Sea and Shore. [Port Gamble. Washington]. 1 [1970]-
current. Supplements, [1971] 'Dictionary of English-
French French-English shell terms ", C. Geerts; [1971]
"Identification Guide: Cow ries of the Red Sea Area", J.
O'Malley. { P.O. Box 219. Port Gamble, WA 98364, USA}
Opisthobranch. .\ monthh publication on mollusks and
marine life. (Phoenix, Arizona]. 16(1-4) [Jan-April 1984].
Continuiition of Opisthobranch Newsletter. Continued
as Shells and Sea Life.
Opisthobranch Newsletter. [Pismo Beach, California]. 1
1969]-15 [1983]. Contiiuied as Opisthobranch.
R. Bieler and A. R. Kabat. 1991
Page 55
Oregon Shell News. Oregon Societ\ of Conchologists.
[Portland. Oregon]. 1(1) [1970]-current. [1(1) labelled as
"new series"].
The Original Tide-ings. Indiana's first shell club. [Crown
Point, Indiana]. 3(7) [July 1973]-6(1) [Jan.-March 1976].
See discussion under Tide-ings.
Pacific Northwest Shell News. [Poulsbo. Washington].
U6) [1961]-13(5) [Oct. 1973]. Continuation of Northwest
Shell News. Continued as Dredgings.
Palm Beach County Shell Club News. [Lake \\ orth.
Florida]. [1961]-[1961]. Continued as Seafari.
The Pariah. [Highstown. New Jersey]. 1[1977]-8[1980].
Pittsburgh Shell Club Bulletin. [Pittsburgh. Penns\l-
\ania]. (1) [1966]-(12) [1977] [most issues not numbered].
Proceedings of the .National Shellfisheries .\ssociation.
[Duxbury, Massachusetts} 45 [1955 "1954"]-70 [1980].
Continuation of Convention papers. National Shellfish-
eries -Association. Continued as Journal of Shellfish Re-
search. Inde.x, "Titles of papers presented at annual meet-
ings. 1930-1973". ii + 37 pages
Proceedings of the Pacific Shell Club. [Los -\ngeles]. 1
[1955]. .\lso issued as irregular "Minutes" and ".\nnual
Report" of the Pacific Shell Club; these publications were
only distributed at meetings and not mailed. Continued,
in part, b\ the News of the .Associated Shell Clubs of
California.
Proceedings of the Philadelphia Shell Club. [Philadel-
phia. Pennsylvania} IJ) [1955}-1(9! [1964'.
Proceedings of the Society for Experimental and De-
scriptive Malacology. Earlier reports published in Mal-
acological Review 2(1). 5(1), 6(1). 7(1) and 8(1-2). Later
reports published in Walkerana (q.v.).
Publication of the Conchology Group of the Outdoor
.Nature Club. Houston. Texas. [Houston. Texas]. 1(1)
[Sept. 1964]. Continued as the Conchologist.
Quarterly Report Miami Malacological Society. [Miami,
Florida]. 1(1) [1967]. Continued as Miami Malacological
Society Quarterly.
Report. .\nnual Meeting and Membership. The
.\merican Malacological I'nion. [Philadelphia, Penn-
sylvania]. (3) [1934 J-i4) [1935]. Continuation of Mrs. Im-
ogene C. Robertson s Rambling Notes ... Continued as
The American Malacological Union. .\nnual Meeting
, Membership List ....
*\ Review of the Broward Shell Club. [Pompano Beach,
Florida]. 1(1! [1963]-^ [last issue seen: 1(3). 1965].
Revista de Malacologia Medica y .\plicada: see entrv
under Journal for Medical and Applied Malacology .
*Sanibel-Captiva Shell Club Newsletter. [Sanibel Island,
Florida]. "^-^ [seen: 8 ^Nov. 1965); 9 (June 1966)]. Con-
tinued as the Junonia.
*San Marcos Shell Club Quarterly Newsletter. [San
Marcos, Texas]. 1 [1972]-^ [last issue' seen: 5(4). 1976].
Sarasota Shell Club Newsletter. [Sarasota, Florida].
[1982]-current.
The Sayana. South Carolina Shell Club Newsletter.
[Charleston, South Carolina]. [April 1989]-current. [1989
issues not numbered].
Seafari. Palm Beach County Shell Club. [Lake Worth;
\\est Palm Beach, Florida]. [i961]-current. Continuation
of Palm Beach Count\ Shell Club News. First numbered
issue is "6(12)" [Dec' 1964]; issues 9(3-8) [1967J erro-
neousK- labelled as volume 8.
The Searcher. New sletter of the Sea Shell Searchers of
Brazoria Count\. [Lake Jackson, Texas], [1977]-current.
*Shell and Tell. Newsletter of the Gulf Coast Shell Club.
[Beaumont. Texas]. ?-currenti^ [onl\ two issues seen: 4(1),
1984; 5(1) 1985]. Continuation of Between the Tides?
*The Shell Case. Naples Shell Club. [Naples, Florida].
[1971]-[1977:^]. [Seen: 1971, 2(1), 3(1), 4(1); a temporary
variant title of the Naples Shell News, q.v.].
Shell Collector. [Fort Lauderdale, Florida]. Premiere
[1978]; 1 [1978]-2 [1979].
The Shelletter of Shells and their Neighbors. [Redlands,
California] 1 [1960]-24 [1964],
Shell Museum & [Research/Educational] Foundation,
Inc.. Newsletter. [Sanibel, Florida} 1 [1988]-current,
Shell Notes (Frank L>Tnan). [Lantana, Florida]. 1(1)
[1942]-2( 13-151 [June 1951}
[The] Shell-O-Gram. Jacksonville Shell Club, Inc. [Jack-
sonville, Florida], 1 [1966]-current. [\'arious issues have
erroneous \olume and number labels].
Shells and Sea Life. [Phoenix, ,\rizona; Bayside, Cali-
fornia} 16(5) [May 1984]-current. Continuation of Opis-
thobranch. [\arious changes in format and subtitles; vol.
18(3), 1986, announced a change to three separate series:
(1) a monthly "Shells and Sea Life News"; (2) a "Shells
and Sea Life Quarterly"; and (3) a new technical journal
".\cta Mollusca". This plan was apparentK abandoned].
Supplement [1986] "Directorx of Clubs & Services". U701
Hyland Street, Bayside, Ca'95524, USA}
*The Shell Sheet. Reef Roamers Shell Club [formerlv.
The .\nderson Reef Roamers]. [Yigo, Guam} 1 [19 ?]-
current.
The Slit Shell. [Newsletter of the] Indianapolis Shell
Club. [Indianapolis, Indiana} [1977}-current. [Some is-
sues numbered].
The Slug. Perrine Muckulogical Society. [Miami, Flor-
ida} 1(1) [1 April 1989} [A parody}
South Florida Shell Club [incorporated/Inc] MoUusk
Chaser. [Miami, Florida} 7(8^ [August 1969]-14(6) [1976}
Continuation of the Mollusk-Chaser South Florida Shell
Club. Continued as The MoUusk. [Various changes in
format and style; some issues of volume 9 (1971) labelled
as "new series"; December issue of volume 10 (1972)
labelled as "Supplement"].
South Florida Shell Club Newsletter. [Miami, Florida}
1 [1963]-2(5) [1964} Continued as The South Florida
Shell Club's Newsletter Mollusk-Chaser.
The South Florida Shell Club's Newsletter Mollusk-
Page 56
THE NAUTILUS, Vol. 105, No. 2
Chaser. [Miami, Florida]. 2(6) [1964}-5(3) [March 1967].
Continuation of the South Florida Shell CAuh Newsletter.
Continued as Moilusk-C^haser South Florida Shell Club.
*South Padre Island Shell Club [Newsletter]. [South
Padre Island, Texas]. ?-? [seen; issues from 1962-Oct.
1980].
♦Southwest Florida Shell News. Journal of the Southwest
Florida Conchologist Society, Inc. [Fort Myers, Florida].
?-current.
Southwestern Malacological Society, Monthly Meeting
Notice and Newsletter. [Phoenix, Arizona]. [1966]-cur-
rent.
Special Occasional Publication. Department of Mol-
lusks. Museum of Comparative Zoology, Harvard Uni-
versity. [Cambridge, Massachusetts], 1 [1973]-current.
{Museum of Comparative Zoology, Harvard University,
Cambridge, MA 02138, USA}
Sterkiana. [Columbus, Ohio; Shawsviile, Virginia]. 1 [Nov.
1959]-71 [1981].
Suncoast Shorelines. Suncoast Conchologists. [Palm
Harbor, Florida]. 1 [1984]-current.
Tabulata. [Santa Barbara, California]. 1(1) [July 1967]-
7(4) [Oct. 1974]. Index (1974), 14 p.
*Tanipa Drill. The Greater Tampa Shell Club. [Tampa,
Florida]. ?-? [earliest issue seen: 1(3), 1972; latest issue
seen: May 1977 labelled as "volume II first issue"]. Con-
tinuation, in part, of The Greater Tampa Shell Club,
Newsletter.
Texas Conchologist. Houston Conchology Society.
[Houston, Texas]. 1(4-6) ["Dec. 1964-Jan./Feb. 1965"]-
current. [Also a newsletter for meeting announcements].
Continuation of Conchologist. {3706 Rice Boulevard,
Houston, TX 77005, USA 1
Texas Shell News of the San Antonio Shell Club. [San
Antonio, Texas]. 1 [1958]-15 [1972J ["Vol. VII" (1964)
erroneously also used for 1965 and 1966].
Thalcheria. [Chicago]. 19 [1984]-current. Continuation
of Chicago Shell Club Newsletter.
*Tibia Trumpet. Broward North Shell Club. [?, Florida].
1 [1973]-2 [I974J [No further issues?].
Tide-ings. The Newsletter of the Crown Point Shell Col-
lectors' Study Group, Inc. [Crown Point, Indiana]. 1
[ 197 1 ]-current. See also The Original Tide-ings; the latter
was started due to an editorial dispute within this shell
club.
Tide Lines. St. Petersburg Shell Club [, Inc.]. [St. Pe-
tersburg, Florida]. [1970]-current. Continuation of an
annual (uiuiamed) St. Petersburg Shell Club show sou-
venir booklet [1965-1969]. Combined with the Crown
Crier in 1988.
Tide Lines. Treasure Coast Shell Club, Inc. [Palm Citv,
Florida]. 1(3) [1987]-2(2) [1988]. Continued as Treasure
Coast Shell News. Note: this newsletter started in March
1986, but 1(1) and 1(2) were unnamed.
The Tidewater Triton. The Tidewater Shell and Fossil
Club. [Tidewater, Virginia]. 1 [1969J-4 (1973) [some is-
sues of volumes 3 (1972) and 4 (1973) erroneously la-
belled as "IV ' or "VI", respectively].
♦Transactions of the Isaac Lea [Conchological] Chapter
of the Agassiz Association. 1 [1889?]- 10? [1899?]. This
".serial" actually comprised members' reports, which were
annually compiled into a single (?) bound manuscript
volume, and circulated from one member to the next
during the following year. Extracts were published in
various journals, including The Nautilus, usually titled
as the "Isaac Lea Department ". We have not been able
to locate any extant set(s) or issues of the Transactions
(including the copy mentioned by Keen, 1954). See Coan
(1989) for further discussion. Continued as The Con-
chologist.
Transactions of the POETS Society. See Walkerana.
Treasure Coast Shell News. Treasure Coast Shell Club,
Inc. [Palm City, Florida} 2(3) [1988]-current. Continu-
ation of Tide Lines (q.v.) [the St. Petersburg Shell Club
complained of the theft of their title, so 2(3) was given
the current name].
Tryonia. Miscellaneous Publications of the Department
of Malacology of the Academy of Natural Sciences of
Philadelphia. [Philadelphia, Pennsylvania]. 1 [March
1979]-current. {address: see Malacologia]
♦[newsletter]. Utah Shell Society. [Salt Lake City, UtahJ
?-current.
The Veliger. [Berkeley; Davis, California]. 1(1) [June
1958]-current. Supplements (not numbered separately):
Supplement [Vol. 3, 1961 ]"Opisthobranchmollusks from
California", E. Marcus. Supplement [Vol. 3, 1968] "The
Order Anaspidea ", R. D. Beeman; and "Thecosomata
and Gymnosomata", J. A. McGowan. Supplement [Vol.
6, 1964] "The biology of Tegula funebralis (A. Adams,
1855)", ed. D. P. Abbott et al. Supplement [Vol. 7, 1965]
"A glossary of a thousand-and-one terms used in con-
chology", W. H. Arnold. Supplement [Vol. 11, 1968]
"The biology of Acmaea", ed. D. P. Abbott et al. Sup-
plement [Vol. 14, 1971] "The Northwest American Tel-
linidae", E. V. Coan. Supplement [Vol. 15, 1973] "A
systematic revision of the Recent cypraeid [sic] family
Ovulidae (Mollusca: Gastropoda)", C. N. Gate. Supple-
ment [Vol. 16, 1974] "The Panamic-Galapagan Epito-
niidae ", H. DuShane. Supplement [Vol. 17, 1974] "Growth
rates, depth preference and ecological succession of some
sessile marine invertebrates in Monterey Harbor ", E. C.
Haderlie. Supplement [Vol. 18, 1975] "The biology of
chitons", ed. R. Burnett et al. Supplement (new series)
1 [1990] "Catalogue of Recent and fossil molluscan types
in the Santa Barbara Museum of Natural History. I . . . ",
by P. H. Scott et al. History, Keen (1983) and Gate (1989);
index to opisthobranch taxa, Carlton & Behrens (1981).
{Museum of Paleontology, 3 Earth Sciences Building,
Univ. of California, Berkeley, CA 94720, USA)
Walkerana. Transactions of the POETS Society. [Whit-
more Lake, Michigan; Niwot, Colorado; Ann Arbor,
Michigan]. 1 []980]-current. [Volume 1(1), undated (ap-
R. Bieler and A. R. Kabat, 1991
Page 57
parently 1980?), was also designated as the "Transactions
of the Physiological, Oecological, Experimental Taxo-
nomic and Systematic Society . . . from the Proceedings
of the Society for Experimental and Descriptive Mala-
cology ]. There are discrepancies in the dates and order
of several issues: 1(5) ["1982" 30 December 1983]; 2(6)
["1988" 6 June 1989]; 2(7) [31 December 1984]; 2(8)
["1987" 29 April 1988]; 3(10) ["1989" 15 January 1990].
Issues are numbered consecutively independent of the
volume numbers. {P.O. Box 2701, Ann Arbor, MI 48106,
USA}
West American MoIIusca. [San Diego, California]. 1
[1900-1902], See Coan (1966) for dates of publication.
Western Society of IVlalacologists, Annual Report. [Los
Angeles, California]. 7 [1974]-current. Continuation of
Echo. {1633 Posilipo Lane, Santa Barbara, CA 93018,
USA}
Western Society of IVlalacoIogists, Occasional Paper.
[Los Angeles, California]. 1 [1975]-2 [1976].
The Whelk Wavelength. The Georgia Shell Club, Inc.
[Marietta, Georgia]. 1 [1985]-current.
The Xenophora Newsletter. [Van Nuys, California]. 1
[Jan. 1976]-7 [Dec. 1976].
The Young Shell Collectors Quaterly. International
League of Young Shell Collectors. [Lima, Peru; Arling-
ton, Massachusetts]. 1 [1967]-2 [1969].
CENTRAL AND SOUTH AMERICA
Chile
Sociedad Chilena de Malacologia Comunicaciones. [Vifia
del Mar], 1 [1979].
Brasil
Informativo S.B.lVl. [Sociedade Brasileira de Malaco-
logia]. [Porto Alegre; Sao Paulo]. 1 [1969]-current. {Inst,
de Biociencuas, Univ. Sao Paulo, Caixo Postale 20520,
Sao Paulo, Brasil}
Siratus. Periodico Bimensal dos Conquiliologistas do
Brasil/Bimonthly Journal of [the] Conquiliologistas do
Brasil. [Sao Paulo]. 1 [Nov.-Dec. 1989]-current. {C. P.
15011, Sao Paulo/SP, Brasil 01599}
Panama
Donax Panamensis. Sociedad Panamena de Malacolo-
gia. [Panama]. 1 [Feb. 1981]-58/59 [Nov.-Dec. 1985].
Thais. Publicaciones diversas de la Sociedad Panamefia
de Malacologia. [Panama]. 1 [29 June 1981]-4 [17 January
1983].
Peru
The Young Shell Collectors Quarterly. International
League of Young Shell Collectors. [Lima; Arlington, Mas-
sachusetts (USA)]. 1 [1967]-2 [1969].
Uruguay
Comunicaciones de la Sociedad Malacologica del Uru-
guay. [Montevideo]. 1 [Sept. 1961]-current. {Casilla de
Correo 1401, Montevideo, Uruguay}
Venezuela
*Bollettino informativo. Sociedad Venezolana de
Malacologia. [Caracas]. 1 [198 ?]-current.
INDEX
The purpose of this index is to guide the reader to the
appropriate country for the malacological journals and
newsletters listed herein, especially those that do not have
(well known) geographical indication(s) in their title.
Long titles have been abbreviated or truncated to save
space.
Abhandlungen des Archiv
Achatina
Acta Conchyliorum
Actas de la Sociedad
Amenites Malacologiques
American Conchologist
American Journal of Conchology
American Malacological Bulletin
American Malacological Union
Amici Gastropodorum
Amphitrite
AMU News
Annales de Malacologie
Annales de la Societe Malacologique
Annales de la Societe Royale
Apex
Archiv fiir Molluskenkunde
Archives Malacologiques
Argamanit
.\rgamon
.Argonauta
■Argonauta
.Arion
Auger, The
Australian Newsletter
Australian Shell News
Basteria
Beachcomber Report, The
Beilage zum Nachrichtsblatt
Between the Tides
Bibliotheque Conchyliologique
Boletin del Grupo de Estudios
Bollettino Informativo
Bollettino Malacologico
Boston Malacological Club
Brisbane Branch Newsletter
British Shell Collectors' Club
Bulletin, Auckland Museum
Bulletin de la Societe Inter.
Bulletin de liaison de Malac. [Elona]
Bulletin Mensuel d Information
Bulletin of Malacology
Bulletin of Paleomalacology
Bulletin of the American Malac Union
Germany
Belgium/Netherlands
Germany
Cuba
France
USA
USA
USA
USA
USA
Portugal
USA
France
Belgium
Belgium
Belgium
Germany
France
Israel
Israel
Australia
Italy
Belgium
USA
Australia
Australia
Netherlands
USA
Germanv
USA
France
Cuba
Venezuela
Italy
USA
Australia
Great Britain
New Zealand
Switzerland
France
Belgium
Taiwan
USA
USA
Page 58
THE NAUTILUS. Vol. 105. No. 2
Bulletin of the Brooklyn Conch. Club
Bulletin of the Inst, of Malacology
Bulletin of the Malac. Soc. of China
Bull, of the Malac. Soc. of London
Bull, of the Natl. Pearl Res. Lab.
Bulletin of the Nippon Institute
Bulletin Periodique, Fondation Conch.
Bulletino della Societa Malac Italiana
Bulletino Malacologico Italiano
Bulletins de la Sociele Malac. de France
Bulletinus Natalensis
Busycon Bugle
Busycon, The
Cairns Shell News
Capsule, The
Carfel
Central Florida Shell News
Cephalopod Newsletter, The
Chambered Nautilus Newsletter, The
Chicago Shell Club Newsletter
Chiribotan, The
ClAC Newsletter
Circular, The Conchological Society
Circulares de la Sociedad Malacologica
Circulares del Museo y Biblioteca
Clam Chatter
Club Conchylia Informationen
Collector, The
Comunicaciones de la Sociedad Malac.
Conch Courier
Concha
Conchas, Las
Conchiglia, La
Conchiglie
Conchioline, La
Conchologia Asiatica
Conchologica Nonsensica [parody]
Conchological Leaflets
Conchological Magazine, The
Conchological Memoranda
Conchological Notes
Conchological Society
Conchologi.sche Mittheilungen
Conchologist, The
Conchologist, The
C'onchologist
C-"onchologist"s Exchange, The
Conchologists" Newsletter, The
Conchologists of America Bulletin
Cone Shell Alert
Connoisseur of Seashells, The
Con's (^onchology
Contribuciones Malacologicas
Contributions a la faune malacologique
Contributions to Conchology
Convention Papers, Natl. Shellfish. Assoc.
Cookia
Corbicula Communications
Corbicula Newsletter
Correspondentieblad
Cowry, The Great
Crown Oier
Dige.sta Malacologica
Dona.x Panamensis
Double Helix, The [parody]
USA
Dredgings
USA
Japan
Echo, The
USA
Taiwan
Fiona
France
Great Britain
Epitonium, The
USA
Japan
Ezohelix
Japan
Japan
Festi\ us. The
USA
Belgium
Field Naturalists' Section
Australia
Italy
Fiji Shell Club .Annual Magazine
Fiji
Italy
Florida Keys Shell C:lub
USA
France
Folia Malacologia
Poland
South Africa
Fort Myers Shell Club Newsletter
USA
USA
Friends of the Gastropods
USA
USA
Fiir die Litteratur und KenntniB
Germany
Australia
Galveston Shell Club
USA
USA
Gastropodia
USA
Philippines
Giornale di Malacologia
Italy
USA
Gloria Maris
Belgium
Great Britain
Grand Strand Shell Club Newsletter
USA
USA
Grand Strand Shell News
USA
USA
Greater Tampa Shell Club
USA
Japan
Guam Shell News
USA
Great Britain
Halia
Portugal
South Africa
Haliotis
France
Cuba
Handlists of the Molluscan Coll.
Great Britain
Cuba
Hawaiian Malacological Society
USA
USA
Hawaiian Marine Mollusks
USA
Germany
Hawaiian Shell News
USA
USA
Heldia
Germany
Uruguay
Helix
Great Britain
USA
Hitachiobi
Japan
Germany
Hojas Malacologicas
Spain
USA
Iberus
Spain
Italy
Illustrated Catalogue of Japanese Shells
Japan
Italy
Indo-Pacific Mollusca
USA
Belgium
Informatieblad
Netherlands
Japan
Informationen des Club Conchylia [Concha]
Germany
Canada
Informations de la Societe Beige
Belgium
Great Britain
Informative S.B.M.
Brasil
Japan
Irradians
USA
USA
Isokojiki
Japan
USA
Jaarverslagen, Schelpen. Friesland
Netherlands
Great Britain
Jahrbiicher der Deutschen Malak. Ges.
Germany
Germany
Japanese Journal of Malacology, The
Japan
Great Britain
JECOLN Annual Report
Japan
USA
Jersey Sheller, The
USA
USA
Johnsonia
USA
USA
Journal de Conchyliologie
France/Australia
Great Britain
Journal for Medical and .Applied Malacology
USA
USA
Journal fiir die Liebhaber des Steinreichs
Germany
USA
Journal of Cephalopod Biology
Canada
Italy
Journal of Conchology, The
Great Britain
Australia
Journal of Malacology, A [Levantina]
Israel
Cuba
Journal of Malacology, The
Great Britain
France
Journal of Moll. Behav, [parody]
Great Britain
USA
Journal of Molluscan Studies, The
Great Britain
USA
Journal of Shellfish Research
USA
New Zealand
Journal of the Malacological Society
Australia
USA
Junonia
USA
USA
Kainakama
Japan
Netherlands
Keppel Ba\ Tidings
Australia
itain/Australia
Korean Journal of Malacology, The
South Korea
USA
Kreukel, De
Netherlands
Great Britain
Laddergram, The
USA
USA
Lavori della Societa Italiana di Malac.
Italy
USA
Lavori della Societa Malac. Italiana
Italy
R. Bieler and A. R. Kabat. 1991
Page 59
Leaflets in Malacology
Levantina
Littorina, The
Littorinid Tidings, The
Long Island Shell Chib News
Louisiana Malacologicai Society
Low Country Shell Club Newsletter
Magasin de Conchyliologie
Maiacofauna Italica
Malacologia
Malacologia Mostra Mondiale
Malacologicai and Conch. Magazine
Malacologicai Club of Victoria
Malacologicai Newsletter [Levantina]
Malacologicai Review
Malacologicai Society of Australia
Malacologicai Society of London
Malacologicai Society of South Aust.
Malacology Data Net
Malakologiai Tajekoztato
Malakologische Abhandlungen
Malakozoologische Blatter
Mappa
Marginella Marginalia
Material of the National Pearl Res. Lab.
Members of the American Malac. Union
Miami Malacologicai Society Quarterly
Minutes, Conch. Club of Southern Calif.
Miscellanees Malacologiques
Mitchelli, The
Mitteilungen der Berliner Malakologen
Mitteilungen der Deutschen Malak. Ges.
Mollusca
Mollusca, The Off the Beach Notes
Molluscan Digest
Molluscs in Archaeology & the Recent
Mollusk Chaser
Mollusk, The
Monographs of Marine Mollusca
Moreton Bay Radula, The
Mostra Malacologica Messinese
Mrs. Imogene C. Robertson's Rambling Notes
Musee Heude, Notes de Malacologie
Nachrichtsblatt der Deutschen Malak. Ges.
Naples Shell News
National Capital Shell Club
National Shellfish. Assoc. Quarterly News
Nautilus, The
NCMC Newsletter
Neue Litteratur and Beytrage zur KenntniB
New Jersey Shell Club
New York Shell Club Notes
News Letter, Auckland Museum
Newsletter, Marco Island Shell Club, Inc.
Newsletter of African [Achatina]
Newsletter of the Conch. Society
Newsletter of the Jersey Cape Shell Club
News of the Assoc. Shell Clubs of Calif.
News of the Western Assoc, of Shell Clubs
Noble Pen, The
North Carolina Shell Club Bulletin
Northampton Nat. Hist. Soc
North Texas Conchological Soc. Quarterly
Northern California Shell Club News
Northwest Shell News
USA
Notes and Quotes, Rochester Shell
USA
Israel
Notiziario C.I.S.M.A.
Italv
USA
Notiziario S.I.M.
Italy
Great Britain
Nouveau Pes-Pelicani, Le
France
USA
Nouvelles Miscellanees Malacologiques
France
USA
Novitates C-onchologicae
Germanv
USA
Occasional Papers on Mollusks
USA
France
Occasional Publ. of the Inst of Malac.
Japan
Italy
Of Sea and Shore
USA
USA
Opisthobranch
USA
Italy
Opisthobranch Newsletter
USA
Great Britain
Oregon Shell News
USA
Australia
Original Tide-ings, The
USA
Israel
Ostrea
Japan
USA
Out of the Shell
Canada
Australia
Pacific Northwest Shell News
USA
Great Britain
Pacific Shell News
Japan
Australia
Pallidula
Great Britain
USA
Palm Beach County Shell Club News
USA
Hungary
Papustyla, The
Papua New Guinea
Germany
Pariah, The
USA
Germany
Pearl Oyster Information Bulletin
Australia
France
Pes-Pelicani
France
USA
Philippine Shell News
Philippines
Japan
Pittsburgh Shell Club Bulletin
USA
USA
Puirieria
New Zealand
USA
Porphyria
Netherlands
USA
Proi^eedings of the Malac. Society
Great Britain
France
Proceedings of the Malacozool. Meeting
Japan
USA
Proceedings of the Miriam Vale
Australia
Germany
Proceedings of the Natl. Shellfish. Assoc.
USA
Germany
Proceedings of the Pacific Shell Club
USA
USA
Proceedings of the Philadelphia Shell Clu
lb USA
USA
Proceedings of the Soc. for E.\p. [Walkerana] USA
USA
Proces-Verbau.x des Seances de la Societe
Belgium
Canada
Proserpine Shell Club News
Australia
USA
Publicafoes Ocasionais da Sociedade
Portugal
USA
Publication of the Conchology Group
USA
USA
Pyrum
France
Australia
Quarterly Journal of Conchology
Great Britain
Italv
Quarterly Report Miami Malac. Soc.
USA
USA
Reefcomber
South Africa
China
Resenas Malacologicas
Spain
Germany
Review of the Broward Shell Club, A
USA
USA
Revista de la Sociedad Malacologica
Cuba
USA
Revue Biographique de la Societe Malac.
France
USA
Robusta
French Polynesia
USA
Rossiniana
New Caledonia
USA
Royal Society of South Australia
Australia
Germany
Sanibel-Captiva Shell Club News Letter
USA
USA
San Marcos Shell Club Quarterly Newsletter USA
USA
Sarasota Shell Club Newsletter
USA
New Zealand
Sayana, The
USA
USA
Schede Malacologiche del Mediterraneo
Italy
Netherlands
Schriften zur Malakozoologie
Germany
South Africa
Seafari
USA
USA
Searcher, The
USA
USA
Series Conchyliologiques
France
USA
Shell and Tell
USA
USA
Shell Case, The
USA
USA
Shell Collector
USA
Great Britain
Shell Gossip
Fiji
USA
Shell Museum . Newsletter
USA
USA
Shell Notes
USA
USA
Shell, The [La Conchiglia]
Italy
Page 60
THE NAUTILUS, Vol. 105, No. 2
Shell Sheet, The
Shell-0-Gram
Shelletter of Shells and their Neighbors
Shells and Sea Life
Si rat us
Slit Shell, The
Slug Newsletter [parody]
Slug, The [parody]
Snail Farming Researili
Sociedad (^hilena de Malacologia
Soosiana
South Florida Shell Cluh
.South Padre Island Shell Chih Newsletter
Southwest Florida Shell News
Southwestern Malaeologieal Society
Special Occasional Publication
Special Publication, Inst, of Malac.
Species Conchyliorum
Spicileges Malacologiques, Les
Steeriae, La
Sterkiana
Strandloper, The
Subscriber's Bulletin
Suncoast Shorelines
Sunshine Coast Shell Club News
Sydney Sheller, The
Tabulata
Tampa Drill
Tentacle
Texas Conchologist
Texas Shell News
Thais
Thatcheria
Tibia Trumpet
Tide-ings
Tide Lines
Tidesong
Tide Watch
Tidewater Triton, The
Townsville Shell Club
Transactions of the Chinese Society
USA
USA
USA
USA
Brasil
USA
South Africa
USA
Italy
Chile
Hungary
USA
USA
USA
USA
USA
Japan
Great Britain
France
French Polynesia
' USA
South Africa
Great Britain
USA
Australia
Australia
USA
USA
Great Britain
USA
USA
Panama
USA
USA
USA
USA
New Zealand
Australia
USA
Australia
China
Zeitsehrift fiir Malakozoologie
Zoological Becord (Mollusca Sect.)
ACKNOWLEDGEMENTS
Germany
Great Britain
Transactions of the Isaac Lea Conch. Chapter USA
Transactions of the POETS Soc [Walkerana] USA
Treasure Coast Shell News LTSA
Tree and Land Snail Collectors Papua New Guinea
Tryonia USA
UN IT AS Malacologica Newsletter Germany
Unterhaltungen fiir Conchylienfreunde Germany
Utah Shell Society USA
Vancouver & District Shell Bulletin, The Canada
Veliger, The USA
Venus, The Japan
Vertigo France
Victorian Branch Bulletin Australia
Walkerana USA
W. A. Shell (Collector Australia
Wellington Shell Clhib New Zealand
West American Mollusca USA
Western Society of Malacologists. Ann. Rep. USA
Western Society of Malacologists, Occ. Paper USA
Whangarei Shell Club New Zealand
Whelk Wavelength, The LISA
.Xenophora France
Xenophora Newsletter, The USA
Young Shell Collectors Quarterly, The Peru/ USA
We thank the following colleagues who have checked
the entries for various countries or states on our behalf:
R. Tucker Abbott, Philippe Bouchet, Walter Cernohor-
sky, Eugene Coan, Richard Kilburn, Ian Loch, Bruce
Marshall, Akihiko Matsukuma, Paula Mikkelsen, Robert
Moolenbeek, Richard Petit and Walter Sage; their assis-
tance was indispensable.
In addition, a large number of individuals provided
information on certain publications (usually of shell clubs).
These include Ruby Barry, Alan Bebbington, Connie
Boone, Carol Boswell, Jean Cate, Barb Collins, Phil Col-
man, Phyllis Diegel, Laurie Ford, Peggy Fox, Levente
Fiikoh, Dorothy Germer, Raye Germon, Alison Haynes,
Russell Jensen, Rudolf Kilias, L. A. Koestel, Florence
Kuczynski, Betty Lawson, Charlotte Lloyd, Jordan
Marche, George Metz, Henk Mienis, Walter Morgan,
Olive Peel, Frances Perry, Charles Pettitt, William Pitt,
Berneice Plummer, Thomas Rice, Gary Rosenberg, Rich-
ard Salisbury, J. B. Sessoms, Carol Skoglund, Shirley Slack-
Smith, John Stanisic, Mike Sweeney, Jose Templado, Jane
Topping, F. White and Peggy Williams; we appreciate
their help.
For helpful discussion concerning the content and the
introductory material of this paper, we thank Kenneth
Boss, Eugene Coan, William Emerson, M. G. Harase-
wych, Richard Johnson, Richard Petit and Gary Rosen-
berg.
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London 22:357.
1988/89 Zoological Record Serial Sources (covering Zoological
Record, X'olumes 124-125). 1990. Biosis, Philadelphia,
X -f 291 pp.
THE NAUTILUS 105(2):62-80, 1991
Page 62
First Records For Cymathim mundum (Gould) in the
Eastern Pacific Ocean, with Comments on the
Zoogeography of the Tropical Trans-Pacific
Tonnacean and Non-Tonnacean Prosobranch
Gastropods with Indo-Pacific Faunal
Affinities in West American Waters
William K. Emerson
Dt-parlment of Invertebrates
American Museum of Natural History
Central Park West at 79th Street
New York, NY 10024, USA
ABSTRACT
Cymalium (Monoplex) mundum (Gould, 1849), a wide-rang-
ing Indo-West Pacific species, is recorded from the Galapagos
Islands. This taxon has yet to be recognized elsewhere in the
eastern Pacific Ocean, but is here reported from the western
Atlantic for the first time. A lectotype is selected for Cymatium
gemmatum (Reeve, 1844), which was previously confused with
C. mundum. The occurrences are documented in west Amer-
ican waters for other trans-Pacific Tonnacean species of the
following genera: Tonna in the Tonnidae, Charonia. Cyma-
tium. Linatclla in the Ranellidae, Distorsiu in the Personidae,
and Bursa in the Bursidae.
The modern Tonnacean faunal element in the eastern Pacific
apparently reflects survivors of a previously widespread, post-
Tethyan biota modified by factors resulting from the closure
of the Central American Seaways about three million years ago,
together vvitli post-Pliocene supplementation from the central
Pacific, The present non-Tonriacean prosobranchs in the eastern
Pacific with Indo-Pacific faunal affinities appear to represent
largely post-Pliocene additions to the eastern Pacific that were
derived from western Pacific communities.
Key words: Tonnacea; systematics; new records; Ranellidae;
Personidae; Bursidae; zoogeography; Indo-Pacific Prosobran-
chia in eastern Pacific Ocean; marine snails.
INTRODUCTION
This paper records the presence in the eastern Pacific
Ocean of the Indo-West Pacific ranellid gastropod, Cy-
malitim mundum (Gould, 1849). The occurrences of this
wide-ranging species in the New World are based on
specimens collected by Jacqueline DeRoy and Carmen
Angermeyer in the Galapagos Islands, some 15 to 35 years
ago. This taxon is differentiated from Cymatium gem-
matum (Reeve, 1844), with which it was commonly con-
fused. The first records of C. mundum in the western
Atlantic are here reported, based on specimens collected
by Frank and Vera Lyman off Florida some 50 years
ago. Additional specimens of Cymatium muricinum
(Roding, 1798), a tropical cosmopolitan species known
elsewhere in the eastern Pacific from Panama (Emerson,
1983:119, figs. 15, 16) are reported from the Galapagos
Islands confirming a previous record which was based
on a specimen in the DeRov Collection (Emerson, in
Radwin, 1969:235).
A zoogeographical survey was undertaken of the trop-
ical trans-Pacific Tonnacean species and non-Tonnacean
prosobranchs with Indo-Pacific faunal affinities known
to occur in the western and eastern Pacific. On the basis
of the available data, the origins of these faunal elements
in the eastern Pacific Ocean are discussed in terms of
their distribution in time and space.
ABBREVIATIONS
AHF = Allen Hancock Foundation Collection; see Eraser
(1943) and McCulloch (1977) for station data (depos-
ited in the LACMNH)
AMNH = American Museum of Natural History, New
York
BM(NH) = The Natural History Museum, London [for-
merly the British Museum (Natural History)]
CASIZ = California Academy of Natural Sciences, San
Francisco, CA
LACMNH = Los Angeles County Museum of Natural
History, CA
MCZ = Museum of Comparative Zoology, Harvard Uni-
versity, Cambridge, MA
MNHN-Paris = Museum National d'Histoire Naturelle,
Paris
W. K. Emerson, 1991
Page 63
NMNH = National Museum of Natural History, U.S.
National Museum Collection (USNM), Smithsonian In-
stitution, Washington, D.C.
NZGS = New Zealand Department of Scientific and
Industrial Research, Geology and Geophysics Division
[formerly the New Zealand Geological Survev], Lower
Hutt
SBMNH = Santa Barbara Museum of Natural History,
Santa Barbara, CA
article 49b, in place of Neptunellininae Gray, 1854 as
used by Waren and Bouchet, 1990:58.
Genus Cyrnatitan Boding, 1798
Type species: Miirex jemorale Linne, 1758, by subse-
quent designation of Dall (1904:133). Synonyms: Loto-
rium Montfort, 1810; Tritociirris Lesson, 1842; Nyctilo-
chus Gistel, 1848.
SYSTEM.ATIC TREATMENT
For man\ \ears. the identit>' of Cymatium mundum
(Gould, 1849) has been misunderstood by most workers,
who erroneously considered Gould's ta.xon to be a junior
subjective synonym of C. gemmatum (Reeve, 1844). This
confusion dates from Reeve's (1844a) original description
and illustrations in which specimens of what are now
recognized as C. mundum (Gould, 1849) were consid-
ered by him to be a variety of C. gemmatum (A. Beu,
in litt.. November 13, 1989). As indicated by Springsteen
and Leobrera (1986:116), C. mundum differs from C.
gemmatum in the shell being ". . . shorter and wider
and more coarsely sculptured; [with] the varices . . . thick-
er and [with] the intervarical cords . . . more pronounced
especially on the spire whorls. ..." The slender, more
delicate Cymatium gemmatum is rarely white and is
commonly orange-brown in basal color, with darker
brown and white varical bands as shown in color by
Springsteen and Leobrera (1986:115, pi. 31, figs. 12a,
12b). The more massive C. mundum is yellowish to white
in basal color and specimens commonly exceed 30 mm
in length (Abbott and Dance, 1982:120, us C. gemmatum
Reeve). In contrast, specimens of C. gemmatum rarely
attain 30 mm in length and they lack, in maturity, the
heavy outer lip characteristic of C. mundum on which
the apertural lip projects above the labral surface (cf.
figure 8 with figure 24).
Cymatium gemmatum appears to be largely confined
in distribution to the western Pacific, where in the central
Philippine Islands specimens are found in depths to 100
meters. Verified records of this taxon from localities on
the Pacific Plate, however, are scarce. In the case of C.
mundum, the range is spread throughout the Indo-Pa-
cific faunal province. Moreover, specimens of this taxon
are commonly found at localities on the Pacific Plate,
from the intertidal zone to moderate depths. See range
data given below.
In the following section, the taxonomic status of these
two taxa is reevaluated and the two nominal species are
recognized as separate and distinct taxa.
Family Ranellidae Gray, 1854
(= Cymatiidae Iredale, 1913; see Beu and Cernohorskv,
1986:242)
Subfamily Cymatiinae Iredale, 1913; see Jansson and
Beu, 1990a, proposal to conserve Cymatiinae under ICZN
Subgenus Monoplex Perry, 1811
Type species: Monoplex australasiae Perry, 1811 [=
Cymatium parthenopeum (von Salis, 1793)], by subse-
quent designation of Dall (1904:138), fide Beu and Kay
(1988:197).
Synonyms: Lampusia Schumacher, 1817; Cymatriton
Clench and Turner, 1957.
Cymatium {Monoplex) gemmatum
(Reeve, 1844)
(figures 1-8)
Triton gcmniatns Reeve, 1844a, Triton text, pi. 15, figs. 60a, b;
figs. 1, 2 herein.
Type locality: Island of Ticao, Philippines (12°30'N,
123°42'E), "Found on stones at low water); Cuming".
Reeve, 1844b: 117.
Cymatium {Monoplex) gemmatum, Beu, 1985:58, "West Pa-
cific".
Cymatium (Septa) gemmatum, Springsteen and Leobrera, 1986:
114. 116. pi 31. figs, 12a, b, "Camotes Sea and Sulu Sea",
Philippines,
Triton (Simpulum) gcmmatus, Tryon, 1880:13. in part, pi, 7,
fig. 41 only [copy of Reeve, 1844a, pi, 15, fig. 60a].
Range: western Pacific: Philippine Islands (Bohol Straits,
AMNH 232143; Punta Engano, Mactan Island, Cebu,
ex.— F. J. Springsteen, AMNH 232142); Palau Islands
(Kror Island, AMNH 92774), Indonesia (Ambon Island,
USNM 746407); Solomon Islands (AMNH 93092; NZGS);
Fiji Islands (Vanua Levu, USNM 695079); New Cale-
donia (MNHN-Paris; teste Beu); Coral Sea (Chesterfield-
Bellona Plateau, MNHN-Paris; teste Beu), Austral Is-
lands (Tubuai Island, USNM 705501 ).
Remarks: The probable syntypes of Triton gemmatus
Reeve in the BM(NH) [lot #198055] consist of 4 speci-
mens, which range from 16.2 mm to 28.9 mm in height.
One of these specimens appears to be the specimen used
to illustrate Triton gemmatus (Reeve. 1844, pi. 15, figs.
60a, b) and is here selected as the lectot\pe (cf. figures
1, 2 with 3, 4 herein). The lectotype (28.9 mm in height)
is slightly larger than Reeve's figure (28. 1 mm). The early
whorls are missing owing to breakage, as is the case in
Reeve's figure, but the lectotype possesses a varix on the
third postnuclear whorl which is not shown on Reeve's
illustration Otherwise, the lectotype compares favorably
with the figure. Three paralectotypes of C. gemmatum
Page 64
THE NAUTILUS, Vol. 105, No. 2
Figures 1-8. Cymatium gemmatum (Reeve); x 2. 1, 2. Copy of Triton gemmatus Reeve, 1844a, pi. 15, figs. 60a, b. 3, 4. Lectotype
BM(NH) 198055. 5, 6. Paralectotype BM(NH) 198055a. 7. 8. .\MNH 232142, off Mactan Island. Cebu, Philippines, ex-F. J.
Springstein. Figures 9, 10. Cymatium muricinum (Roding) Crabbed specimen, Santa Cruz Island, Galapago.s Ids.. Oct. 1969,
AMNH 232144, ex-J DeRoy; xl Figures 11-15. Cymatium mundum (Gould); x 2. 11. Copy of Triton mumium Gould, 1849,
after Gould, 1856, pi. 17, fig. 297. 12, 13. Lectotype USNM 5695. 14, 15. Copies of Triton mundum Gould, 1849, after Gould,
1856, pi. figs. 297a,b.
W K. Emerson, 1991
Page 65
are here recognized in the type lot [BM(NH) 198055].
The largest of the three paralectotypes is illustrated here-
in (figures 5, 6).
The old locality label accompaning the type lot reads,
"T. gemmatus, var., I. Masbate [12°11'N, 123°30'E] on
the reefs, under stones. H. [ugh] C. [uming]". Whereas
the "Hab. [itat]" given by Reeve (1844a); for T. gem-
matus is "Island of Ticao, Philippines" [12°13'N,
123°42'E], (found under stones at low water; Cuming)."
These islands are in close pro.ximity, being separated by
the Masbate Pass. Inasmuch as the existing label accom-
panying this lot refers to a variety of T. gemmatus, the
original locality stated by Reeve (1844a); for this taxon
is here retained as the type locality.
At the time Triton gemmatus was proposed, Reeve
(1844a, pi. 15, fig. 60c) briefly described and illustrated
a specimen of what is now recognized as Cijmatium
mundum under the appellation of "Triton gemmatus
variety B. ' There are three specimens in the type lot
[BM(NH) #196736]. The largest specimen most closely
resembles Reeve's figure (see figure 20, herein), but this
specimen measures 40.9 mm in height vs. the figure
which is 38 mm in height. The other two specimens are,
respectively, 38.9 and 33.5 mm in height. Although these
specimens are typical of examples of Ctjmatium mun-
dum (Gould, 1849), they retain the status of paralecto-
types of Ctjmatium gemmatum (Reeve). The old label
accompanying the type lot cites only the "Philippines"
as the locality.
Cymatium (Monoplex) mundum
(Gould, 1849)
(figures 11-24)
Triton gemmatus variety B, Reeve, 1844a, Triton text,
pi. 15, fig. 60c [figure 20 herein], "Island of Annaa (Chain
island). South Pacific Ocean, and island of Burias, Phil-
ippines (found under stones in both localities at low wa-
ter); Cuming". Reeve, 1844b: 117. Not T. gemmatus
Reeve.
Triton mundum Gould, 1849:143; Gould, 1856:506,
pi. 17, figs. 297, 297a-b [figures 11, 14, 15 herein]; Gould,
1862:66; Johnson, 1964:112. Type locality: "Tutuilla [sir;
Tutuila] Samoa Islands" (American Samoa, 14°18'S,
170°42'W).
Cymatium {Morwplex) mundum, Beu 1985:58, "Indo-
West Pacific".
Cymatium (Septa) mundum, Springsteen and Leo-
brera, 1986:116, pi. 31, fig. 13, "Camotes Sea and Sulu
Sea , Philippines.
Triton (Simpulum) gemmatus, Tryon, 1880:13, in part,
pi. 7, figs. 43, 44 [copy of Gould, 1856, figs. 279, 279a]
only. Not T. gemmatus Reeve.
Cymatium (Septa) gemmatum. Clench and Turner,
1957:222, in part, reference in synonymy to Triton mun-
dum Gould only. Not T. gemmatus Reeve. [Clench and
Turner mistakenly referred their western Atlantic rec-
ords of the Indo-west Pacific species, C. vespaceum (La-
marck, 1822), to C. gemmatum. Beu (1985:60) now con-
siders these east American populations to be a distinct
species, namely: C. comptum (A. Adams, 1854)]. Finet,
1985:19, "Galapagos Islands", based on AM.\H 1.39535.
Not T. gemmatus Reeve.
Cymatium gemmatum, Salvat and Rives, 1980:306,
fig. 173, "lies de la Societe aux Tuamotu a Raevavae
(Australes) et aux Marquises"; Abbott and Dance, 1982:
120, "Indo-Pacific; West Indies, under coral, on sand,
shallow water; uncommon". Not C. gemmatum Reeve.
Range (Selected localities from 50 AMNH lots): West-
ern Indian Ocean (Durban Bay, South Africa, AMNH
196938); (off Gedi, Kenya AMNH 96528); (off Kizimkazi,
Zanzibar, AMNH 101032); off Somalia (Mogadiscio,
AMNH 142680); (Mauritus, AMNH 113606); eastward
into the Pacific Ocean, northward to southern Japan
(Okinawa, AMNH 169800) and southward to the Fiji
Islands (Suva Reef, AMNH 214467), Solomon Islands
(Malaita Island, AMNH 118330) and New Caledonia
(Touho, AMNH 104266), northeastward to the Hawaiian
Islands (Oahu Island, AMNH 152284; Maui Island,
AMNH 147510) and southeastward to the Societv Islands
(Huahine Island, AMNH 231972; 239056); French Aus-
tral Islands (Tubuai, AMNH 84641) and Marquesas Is-
lands (Salvat and Rives, 1980:305). NEW RECORDS For
The Eastern Pacific Ocean: Galapagos Islands, Academy
Bay, Isla Santa Cruz, 1 fresh specimen, found on beach,
June, 1966, ex-J. DeRoy (AMNH 139535); 1 crabbed
specimen, found at low tide, December, 1965, ex-J. DeRoy
(AMNH 232145), see figures 21, 22. Galapagos Islands,
1 crabbed specimen, South Channel (Isla Santa Cruz-
Isla Baltra) collected about 1975, C. Angermeyer collec-
tion. NEW RECORDS For The Western Atlantic: 1 spec-
imen, south end of Lake Worth, Palm Beach County,
Florida, Vera Lyman!, ex-A. D'Attilio collection (NZGS
WM 15228, teste A. Beu). 1 specimen dredged from Gulf
Stream, off Palm Beach County, Florida, Frank Lyman!
1940, ex-AHF (LACM 115537, teste A. Beu).
Remarks: In a review of the taxa described by Augustus
A. Gould, Johnson (1964:112) selected for Triton mun-
dum Gould (1849:506, pi. 17, fig. 297, a-b) the lectotype
(USNM 5695, cf. figures 12, 13 with Figure 11, herein)
and a "paratype" [= paralectotype] (USNM 612311, cf.
figures 14, 15 with figures 16, 17 herein). These speci-
mens, which measure 29.9 and 21.9 mm in height, re-
spectively, compare favorably with Gould's somewhat
stylized drawings. As Johnson (1964:112) noted, an ad-
ditional specimen, which is labeled a paratype [= para-
lectotype] from the A. A. Gould Type Collection, No.
A427, is present in the Museum of Comparative Zoology
(#169249). The MCZ specimen (figures 18, 19), which
had been obtained from the New York State Museum,
measures 28.8 mm in height.
The two type specimens (lectotype and paralectotype)
in the USNM collection are presumably from the type
locality, "Tutuilla, Samoa Islands " (Gould, 1849), but the
existing labels only identify the source of the two lots,
namely: the U.S. Exploring Expedition (1838-42). There
Page 66
THE NAUTILUS, Vol. 105, No. 2
Figures 16-24. Cyinatium mtindum (Gould); x2. 16, 17. Paralectotype USNM 612311. 18. 19. Paralectotype MCZ 169249.
20. Copy of Triton gemmatus var. B. Reeve, 1844a, pi. 15, fig. 60c [= C. mundum]. 21, 22. Crabbed specimen, Santa Cruz
Island, Galapagos Ids., ,\MNH 232145, e.\-J. DeRoy 23, 24. Huahine Island, Society Ids., AMNH 239056, note fully developed
outer lip
are no additional data in the USNM catalogue for either
lot (teste M. G. Harasewych).
ZOOGEOGRAPHIC IMPLICATIONS
Tables 1 and 4 enumerate and document the trans-
Pacific Tonnacean species known to occur in the tropical
eastern Pacific, including records for the oceanic islands
(Clipperton Island, Revillagigedo Islands, Cocos Island,
and Galapagos Islands) as well as those from the conti-
nental borderland. Some of the records are based on
limited samples of one to three individuals that are known
only from dead-collected specimens. These include Ton-
na perdix perdix, Cyrnatium aquatile. C. mundum. and
W. K. Emerson, 1991
Page 67
Linatella succincta. The rarity of these and other records
may reflect, in part, the lack of extensive collecting by
means of SCUBA diving and dredging operations. The
marine moUusks of Cocos Island (about 9 km in circum-
ference) are now reasonably well known owing largely
to the intensive collecting activities during the past de-
cade of Drs. Donald R. Shasky and Michel Montoya with
the cooperation of their numerous field associates (Shas-
ky, 1989c; Montoya, 1983, 1988). Whereas the moUuscan
faunas of most of the other oceanic islands may have
received attention for a longer period of time, much of
this collecting predates the use of SCUBA diving, and
the insular faunas of these widely isolated and not easily
accessible islands still remain poorly known. This is also
largely true of the Galapagos Islands because of the geo-
graphical extent of this archipelago and the long-stand-
ing restrictive regulations pertaining to the collection of
specimens (cf. Wellington, 1984). On the mainland, for
most of the tropical zone (Mexico to Ecuador), much of
the subtidal coastal waters and those of the near-shore
islands have not been extensively explored by collectors.
This is especially true of the communities associated with
the coral-reef biotope (Emerson, 1967:91). A possible
exception are the coral reefs of Panama where James
Ernest, Royce Hubert and others have concentrated their
collecting efforts for mollusks in recent years (Emerson,
1983). Nevertheless, the majority of the Tonnacean rec-
ords are based on live-collected or well-preserved dead
specimens from numerous sources over a considerable
period of time, commencing with explorations early in
the nineteenth century by Alexander von Humboldt and
Aime Bonpland (Valenciennes, 1833).
Students of zoogeography have long noted the pres-
ence of marine mollusks with Indo-Pacific faunal affin-
ities in the subtropical and tropical waters of the eastern
Pacific (Carpenter, 1857:346; Darwin, 1860:391 and oth-
ers). Hertlein (1937) was the first, however, to discuss
specifically this subject and detail the shallow-water mol-
luscan species then known to be common to Polynesian
and western American waters. He concluded that these
Indo-Pacific species had likely dispersed the 5,000-km-
wide body of open ocean (Ekman's East Pacific Barrier)
separating the most eastern situated Polynesian Islands
and the New World by means of long-lasting larval stages
transported by ocean currents, and/or by attachment to
drifting objects, to pelagic fishes, to mammals, or on the
feet of wide-ranging oceanic birds. He also noted that
nearly all of these mollusks were gastropods, many in
the Conidae, Cymatiidae [= Ranellidae and Personidae]
and Cypraeidae, and most of them were known in the
eastern Pacific only from the oceanic islands, far from
the West American Continental Borderland.
Our present knowledge largely serves to support Hert-
lein s conclusions. In the tropical Tonnacea, several spe-
cies of Cymatium including C. pileare martinianum,
and C. miiricinum, have been shown to have teleplanic
larvae (Scheltema, 1989:439). Additionally, a larval stage
lasting nearly a year in the water column is known for
the cosmopolitan species Cymatium parthenopeum and
C. nicobaricum (Scheltema, 1971: table 1). Furthermore,
C. parthenopeum and others are known to feed in the
plankton and to be able to delay metamorphosis until a
shallow-water substrate is reached (Pechenik et al., 1984).
Oceanic currents appear to be the primary vehicle for
the passive dispersal of mollusks with hemipelagic larvae
(Scheltema, 1986, 1989; Zinsmeister & Emerson, 1979;
Emerson, 1990) and for the distribution of species that
attach to, or burrow into floating objects. Rafting may
be the means of dispersal for some epifaunal bivalves
(Turner, 1966:52, 1971; Scheltema, 1977;93; Emerson,
1978:92) and gastropods (Jokiel, 1990:71). The major cur-
rent circulation in the tropical Pacific apparently has
favored east to west dispersal pathways since the late
Neogene, or perhaps much earlier (cf. Grigg, 1988; New-
ton, 1988; Scheltema, 1988; Skelton, 1988). On the other
hand, the paucity of mollusks with Panamic faunal af-
finities in Polynesian waters suggests that recruitment of
tropical Panamic species into the central Pacific via dis-
persal by the westward flowing current patterns has been
largely unsuccessful (cf. Briggs, 1970:134; Zinsmeister &
Emerson, 1979:36; Scheltema, 1988:149). This mostly one-
way trans-Pacific distribution exists even though the
westward transport of organisms across the East Pacific
Barrier would presumably be facilitated by the accel-
eration of westward flowing currents during pre-El Nifio/
Southern Oscillation events (Jokiel, 1990:69).
As noted previously, Hertlein (1937:309) pointed out
that many of the Indo-Pacific species recorded by him
occurred only on the oceanic islands in the eastern Pacific
and were not known to have spread eastward to the
continental borderland of the Americas. This observation
remains largely true for the non-Tonnacean prosobranch
species, of which only 20 of the 48 taxa (41%) compiled
herein are known from the mainland (tables 2, 3, Ap-
pendix). In contrast, 42 (87%) of these 48 taxa are re-
ported from the oceanic islands in the tropical eastern
Pacific, with 33 (68%) at Clipperton Island, 16 (33%) at
Cocos Island, 10 (20%) in the Galapagos Islands, 7 (14%)
in the Revillagigedo Islands, and 1 (2%) at Guadalupe
Island (table 4). These species represent an attenuated
Indo-Pacific element that has reached the oceanic islands
in the eastern Pacific, but the majority of these ta.xa (59%)
have not been reported from the west American main-
land (table 4). Furthermore, none of the 48 species is
known from the western Atlantic, although one survives
in the eastern Atlantic and the Mediterranean, presum-
ably as a post-Tethyan faunal relict (table 4).
The only fossil evidence for the temporal existence of
the non-Tonnacean Indo-Pacific faunal element in west
America is the presence of Cypraea cernica Sowerby in
the Pleistocene of Guadalupe Island (Lindberg et al.,
1980:52) and Hastula albula (Menke) in the Miocene of
California, a species living at the present time in the
Revillagigedo Islands (Bratcher & Burch, 1971; table 3,
herein). Nor have any representatives of this faunal el-
ement been recovered from Mesoamerican archaeolog-
ical sites (Emerson, 1983:120). If large and colorful gas-
tropods such as Mitra mitra (Linne) had been commonly
Page 68
THE NAUTILUS, Vol. 105, No. 2
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W. K. Emerson, 1991
Page 69
Table 2. Shallow-water Indo-Pacific prosobranch gastnipods {excluding the Toiiiiacea) recorded from the West American Con-
tinental Borderland (Appendix) and their distribution on the eastern Pacific islands (records from F.merson, 1978. 1983; Everson,
1988; Robertson, 1979: Shasky, 1983-1989, and herein). None of these species is known from the .'\tlantic Ocean, except Quoyula
madreporaruiu uhich was reported from the Cape Verde Islands (Emerson, 1983:122)
Re\il-
C'lipperton
lagigedo
txicos
Galapagos
CiiadaluiJc
Island
Islands
Island
Islands
Island
Neritacea
Titiscaniidae
1. THificania liiiuicina (Rergh)
.■\rchitectonicacea'
.'Krchitectonicidae
2. Hcliactis Irochoidcs (Desha\es)-l-#
3. Philippic! radiiita (Roding) + #
Littorinidae
4. Littorina pintado (\Vood)#
Cypraeacea
Cypraeidea
5. Cijpraea captitserpcntis Linne*-l-#
6. Cypraea talpa Linne* + #
7. Cypraea teres CJmelin [= ■'alisunae Burgess]*#
Ovulidae
8. Pseudoeypraea adamsonii (8owerby) +
Muricacea
Muricidae
9. Qitoyula madreporarum (Sowerby)* + #
10. Reliquiaeeava rohiUardi (Lienard)*
11. Rhizochilus antipathieus (Steenstrup)#
Mitracea
Mitridae
12. Mitra mitra (Linne)* + #
Conacea
Conidae
13. Conus chaldcus (R6ding)*-l-#
14. Conus ebraeus Linne*-l-#
15. Conus tessulatus Born* +
Terebridae
16. Terebra a ffinis Gray* + tt
17. Terebra laevigata Gra\ +#
18. Terehra niaeulota (Linne)* + #
Turridae
19. Kermia felina (Hinds)-I-
20. Microdaphnc Inelwdes (Dalll*-i-
Total \ taxa on mainland = 20
X
X
X
X
X
X
X
9(45"
3(15?
X
X
X
X
X
X
10 (50''
X
X
X
X
X
6(315
Explanation of symbols: asterisk (*) = taxa also known from Line Islands (Jewell, 1962; Kay. 1971; Ka\ and Switzer, 1974); plus
sign ( + ) = known from French Polynesia (Dautzenberg and Bouge, 1933; Rehder, 1968; Salvat and Rives, 1980; Richard, 1985);
and number sign (#) = known from Hawaiian .Archipelago (Gage, 1962; Schmeltz, 1978, 1979; Kay, 1979)
' For the purposes of this tabulation, .\rchitectonicacea is retained in the subclass Prosobranchia and not transferred to the subclass
Heterobraiichia (see classification of Ponder and Waren, 1988:308),
available in west American waters, one would expect
specimens to have been reported in the kitchen middens
and burial sites of the Native Americans. On the other
hand, the tropical shallow-water Tertiary record is poorly
preserved on the northwest American borderland. Until
recently, no fossiliferous marine Tertiary onshore de-
posits were recognized regionally, from southern Nica-
ragua northward (Woodring, 1966, 1978; Perrilliat, 1978),
except for the long-known exposures on the Tres Marias
Islands, off San Bias, Mexico, onshore and on the islands
in the Gulf of California and in southeastern California
(Durham & Allison, 1960; Powell, 1988; Smith, 1989).
In the mid-1970's however, reconnaissance exploration
along the Pacific coast of Mexico, northwest of Tehuan-
tepec (Durham et ai. 1981) determined the presence of
seven onshore Tertiary sedimentary basins on the North
America Plate, negating the belief pre\iousK held that
most of these sediments had been lost to subduction or
continental-margin truncation into the Middle American
Trench system (Karige^ ai. 1978; Beck & Plumley, 1979),
Four of these onshore west Mexican basins have yielded
fossiliferous marine sediments of Miocene and younger
Page 70 THE NAUTILUS, Vol. 105, No. 2
Table 3. Shallow-water Iiuio-Pafific pr().sobranch gastropods (excludiiii; the Tonnacea) recorded from the eastern Pacific Oceanic
IslaiKJs that are not known from the West .Vmericaii Continental Borderland (records from Bakus, 1975; Cate, 1969; Kmerson,
1978, 198.3; Everson, 1988; Finel, 1987a,b; Kay, 1979; Shasky. 198:5-89, antl herein). None of these species is known from the
.Mlanlic Ocean.
C:lipperton Revillagigedo Cocos Galapagos Guadalupe
Island Islands Island Islands Island
Fissurellacea
Fissurellidae
1. Diodora granifera ( Pease)* + # X — — —
Trochacea
Phasianellidae
2. Tricolia variabilis ( Pease)* + # — — X — —
Neritacea
Neritidae
3. Serita plicata Liimi:* + U X — — — —
Cerithiacea
Cerithiidae
4. Cerithitini ncsioticiitn Pilsbry & Vanatta* + # X — — — —
Cypraeacea
Cypraeidae
5. Cypraea depressa Gray*+ X — — — —
6. Cypraea helvola Linne* + # X — — — —
7. Cypraea maculijera (Schilder)* + # X — — — —
8. Cypraea moneta Linne* + # X — XX —
9. Cypraea rashleighana Melvill + * X — ? — —
10. Cypraea schilderorum (Ireddle)* + tt X — — — —
11. Cypraea .srurra Gmelin* + # X — — — —
12. Cypraea vitellus Linne* + # X — — — —
Muricacea
Muricidae
13. Drupa nioruni Roding* + # X — — — —
14. Drupa ricinus (Linne)* + # X — — X —
15. Moru/a uya (Roding)* + # X — X — X
16. Na,v.va .serfa (Bruguiere)* + # X — — — —
17. Coralliohia fimlmata (A. Adams)*# X — — — —
18. Corallioplula violacea (Kiener)* + # X X X X —
Buccinacea
Buccinidae
19. Clivipollia costata (Pease) + tt X — — — —
20. Clivipollia fragaria [WoocD + tt — — X — —
21. Cantharus fumostts {Di]]\\\n} + ff — — — X —
Mitracea
Mitridae
22. Mitra edentula Swainson + # X — — — —
23. Mi(ra /crn/guico Lamarck* + # X — X — —
24. Mitra papalis (Linne) + # X — X — —
25. Strigatella litleraia (Lamarck)* + # X — — — —
Volutacea
Harpidae
26. Harpa gracilis Ikoderip & .Sowerby+ X — — — —
Conacea
27. Terefora crenu/a*a (Linne)*-l-# X X X — —
28. /7a«<«/a a//w/a (Menke) + # — X — — —
Total N of insular taxa = 28' 24(85%) 3(10%) 8(32%) 4(14%) 1(3%)
Explanation of ssnibols: asterisk (*) = taxon also known from Line Islanils (Jewell. 1962; Kay, 1971; Ka\ and Switzer, 1974); plus
sign ( + ) = known from French I'oKnesia (Dautzenberg and Bouge, 1933; Rehder, 1968; Salvat and Rives. 1980; Richard, 1985);
and number sign (#) = known from Hawaiian Archipelago (Schmeltz, 1978, 1979; Kay, 1979).
' Additional unverified records: 1, Vanihuro acuta (Rechiz) from Cocos Island (Shasky, 1987:49) and Triphora trilicea Pease from
the Galapagos Islands (Shasky, 1989a:8).
W. K. Emerson, 1991
Page 71
ages (Durham et al., 1981; Perrilliat, 1981). Indo-Pacific
molluscan elements have yet to be reported from these
basins (PerriUiat, 1987). Furthermore, no mollusks of the
non-Tonnacean Indo-Pacific faunal element have been
recorded from the numerous fossiliferous terrace deposits
of Pleistocene age along the west American mainland,
from Mexico to Peru. Thus, the paucity of tropical Ter-
tiary fossils from much of the middle West American
Continental Borderland has largely limited paleogeo-
graphic interpretations of the faunistic history of the
modern molluscan Panamic Province to the regional
Pleistocene fossil record and to comparisons with the
Tertiary faunal records of the adjoining Californian and
Caribbean regions.
The non-Tonnacean element (tables 2, 3) is present
only at the genus-group level in the Tertiary faunas of
the Caribbean region. A small Indo-Pacific relict element
(conid subgenera, etc.) is recognized by Petuch (1988:
94, 96) as regionally extinct species in the West Indian
Gurabo Formation (Pliocene). However, none of the non-
Tonnacean Indo-Pacific species now living in the eastern
Pacific is known to inhabit the western Atlantic (table 4)
and none of these species has been recognized in the
fossil record of the east American tropics. The modern
and fossil occurrences of the species of the non-Tonna-
cean Indo-Pacific faunal element in the eastern Pacific,
therefore suggest that these taxa largely represent post-
Pliocene colonizers derived by trans-Pacific dispersal from
populations in the central Pacific.
This distributional pattern is in sharp contrast to that
for the 13 trans-Pacific Tonnacean species cited in table
1 . Eleven of the 13 species (84% ) also occur in the western
Atlantic, 7 (53%) of which are recognized as subspecif-
ically divergent taxa. The vast majority (8 of the 13 taxa)
are represented in the eastern Atlantic and several (5 of
the 10 taxa) also have populations in the Mediterranean
Sea (see table 1). Only eight (61%) of these trans- Amer-
ican taxa, however, can be termed truly circumtropical
in distribution as the others are not known from the
eastern Atlantic (table 4).
The fossil record of the Tonnacean species in the New
World tropics, although sparse, is somewhat better doc-
umented at the species-group level than that of the non-
Tonnacean trans-Pacific prosobranch species cited herein
(tables 2, 3, Appendix). In the Tertiary, species of Cy-
matium (sensn lato) are recognized in Oligocene de-
posits of Panama and Mississippi (MacNeil and Dockery,
1984; Schmelz, 1989). Other cymatiid taxa are known
from the Neogene period and Pleistocene epoch of the
eastern Pacific and western Atlantic tropics. Beu and Kay
(1988:196) recorded the presence of ". . . both C. pileare
and C. aqitatile . . in the Pacific [Basin] since Miocene
time", and they noted ". . . that C. aquatile lived in
northwestern Ecuador at some time [Pliocene] during
the Neogene [Olsson (1964; pi. 30, fig. 3, as C. cf. pi-
leare)], and that unnamed taxa close to, but different
from, both C. aquatile and C. martiniantim lived in the
Caribbean during Miocene to mid-Pliocene time." The
genus Distorsio is also known from the Oligocene of
Mississippi (MacNeil, 1984) and elsewhere in Neogene
and Pleistocene deposits of the east Americas, from Flor-
ida to Brazil (Pilsbry, 1922; Emerson & Puffer, 1953;
Woodring, 1959). In west America, Distorsio is reported
from the Mio-Pliocene of southeastern California (Pow-
ell, 1988:17, as D. constricta), of Costa Rica (Aguilar &
Fischer (1986:223) and of northwestern Ecuador (Pilsbry
and Olsson, 1941:40; Olsson, 1964:174). Distorsio first
appears in the Cretaceous of Madagascar (Beu, 1988:89),
Cijmatium (sensu lato) is known from the Paleogene
and Neogene of the Old World [e.g., Oligocene onwards
in the Aquitaine Basin), and Charonia occurs in the Oli-
gocene of Europe and the Neogene of both southern
Europe and the Caribbean area (Beu, 1970; Gibson-Smith,
1971) suggesting that the New World representatives of
these taxa form part of a post-Tethyan distributional
pattern.
All 13 of the Tonnacean species listed in table 1 are
known to occur on the oceanic islands within the eastern
Pacific. Ten (76%) of the species are recorded from the
Galapagos Islands, with 7 (53%) at Cocos Island, 4 (30%)
at Clipperton Island and 3 (23%) in the Revillagigedo
Islands. Seven species (58%) are reported from the main-
land, including Cymatium corrtigatum amictuni, a non-
trans-Pacific species with divergent subspecies in the
western and eastern Atlantic Ocean (west Africa) and
the Mediterranean Sea (table 1). Only 4 of the 13 taxa
are commonly found on the west American mainland
(Cymatium corrugatum amictum, C. pileare macrodon,
C. parthenopeum keenae, and Distorsio c. constricta).
One of these taxa (C. p. macrodon) is recognized by Beu
and Kay (1988:211) as a full species on the basis of shell
morphology, although they note the close resemblance
of the eastern Pacific specimens to specimens of "typical"
C. pileare in the western Pacific. Moreover, a faunal
antiquity for the east American Tonnacea is further sup-
ported by the recognition in the western Atlantic of 7 of
the 1 1 Pan-Panamian species divergent at the subspecies
level.
The presence of 10 of the 13 eastern Pacific Tonnacean
species living in the Galapagos Islands, with only 7 species
on the west American mainland, may identify this ar-
chipelago as a post-Pliocene refuge for some of these
Tonnaceans. This hypothesis is supported by the low num-
bers of Tonnacean species known from the other oceanic
islands, especially Clipperton Island (4 species) and Cocos
Island (7 species). Adverse post-Pliocene conditions in
the tropical eastern Pacific may have caused the extinc-
tion of some elements of the insular and mainland Ton-
nacean populations. If this is the case, some of the present
trans- Pacific Tonnacean fauna in the eastern Pacific must
represent elements derived by dispersal from central Pa-
cific populations commencing with the emergence of the
Panamanian isthmus. Dana (1975) made a similar case
for the decline of the Indo-Pacific hermatypic corals in
the west American tropics during the Pliocene and their
subsequent trans-Pacific replacement in the Pleistocene-
Holocene. These factors (fossil record and the modern
distributional pattern in the New World tropics) suggest
Page 72 THE NAUTILUS, Vol. 105, No. 2
Table 4. A compilation comparing the distribution of the Incio-Pacific non-Tonnacean and Tonnacean prosobranchs in the eastern
Pacific Ocean, with their occurrences in the Atlantic Ocean and Mediterranean Sea (based on tables 1-3, .•\ppendix)
Non-Tonnacean species
1 Diodora granifera
2. Tricolia varialrilis
3. Nerita plica ta
4. Titiscania limacina
5. Littorina pintado
6. Cerithimii nesiolicuni
7. Helianis trochoides
8. Philippiu radiata
9. Cijpraea capiitserpcntis
10. Cijpraea depressa
11. Cijpraea helvola
12. Cijpraea niaculifera
13. Cypraea moneta
14. Cypraea rashleighana
15. Cypraea schilderorum
16. Cypraea scurra
17. Cypraea lalpa
18. Cypraea teres
19. Cypraea vitellus
20. Pseudocypraea adamstmii
21. Drupa rnorum
22. Drupa ricinus
23. Morum twa
24. Nassa serta
25. Quoyula madreporanim
26. Reliqniaecava rohillardi
27. Rhizochilus antipathiciis
28. Coralliohia finihriata
29. Coralliophila violacea
30. Clivipollia eostata
31. Clivipollia jragaria
32. Canthanis fumosus
33. Mitra edentula
34. Mitra ferruginea
35. Mitra mitra
36. Mitra papalis
37. Strigatella litterata
38. Harpa gracilis
39. Conus chaldeus
40. Conus elnaeus
41. Conus tessulatus
42. Terehra ajfijis
43. Terelna crenulata
44. Terelna laevigata
45. Terelna maculata
46. Hastula allmla
47. Kerrnia felina
48. Microdaphne trichodes
Total N of taxa = 48
Tonnacean species
1. Tonna p. perdix
2. Charonia t. tritonis
3. Cymatium aquatile
4. Cymatium corrugatutn amictum
5. Cymatium mundum
Eastern Pacific Ocean
.Atlantic
Ocean
Oceanic
Western
Eastern
Mediter-
Islands
Mainland
Atlantic
Atlantic
ranean Sea
X'
X^
X'
—
—
—
—
X'
X'
X
X
"
=
=
X''
X
X
—
—
—
X1.3
X'
X'
X'
Xl.3.4
Xl.'3
X'
X'
x^
X
—
—
—
—
—
—
—
X
—
—
—
\1.3.-l
X'
X3.J
X'
X
—
—
—
X
—
—
—
X1.3..';
X'
Xl,2
—
—
—
—
X
I
X
X"
X'
X
X
—
—
—
X'
X 1.2.3.4
X'
x^
x^
—
—
—
—
—
—
—
—
X'
Yl,3
—
—
—
—
X3.4
X
—
—
—
XI. 3
X'
—
—
—
—
X'
Xl.3.4
X
—
—
—
Xl.3.4
X
—
—
—
Xl.2.3
X
X
X
X
—
—
—
XI. 2.3
—
—
—
X..2.3
I
x^
X
X
—
—
—
x^
—
—
—
42 (87%)
20(41%)
0
1 (1%)
1 (1%)
x^
_
X''
_
X3.4
—
X"
X'
X'
X'
X
X
X
—
XM.4
X
X"
X9
X»
X'
—
X
—
—
W. K. Emerson, 1991
Table 4. (^mtinued
Page 73
Eastern Pacifi(
lOi
cean
.•\tlai
itic Ocean
Oceanic
Western
Eastern
Mediter-
Islands
M;
ainlainl
Atlantic
Atlantic
ranean Sea
6. Cumatium muricinum
X^
x
X
X
X
7. Cymatium pileare macrodon
X 1.2.3.4
X
X'"
X'"
—
8. Ci/matium nicobaricum
XI.3
—
X
X
X
9. Ciimatium parthenopeum kecnac
X^
X
X"
X"
X"
10. Linalella succincta
x^
—
—
X
—
11. Distorsio c. constricta
XM
X
X12
—
—
12. Bursa asperrima
Xi.-i
—
—
—
—
13. Bursa g. granularis
X1.M
X
X13
—
—
Total N of taxa = 13
13(100%)
7
(53%)
11 (84%)
8(61%)
5 (.38%)
Non-Tonnat'ean and Tonnacean species
Total N of taxa = 61
.5,5(90<7)
27
(44%)
11 (18%)
9(14%)
6(9%)
Known from: 1 = Clipperton Island, 2 = Revillagigedo Islands, 3 = Cocos Island, 4 = Galapagos Islands, and 5 = Guadalupe
Island. Recognized as subspecies: 6 as T p. maculosa. 7 as C. /. variegata. 8 as C. c. krebsii. 9 as C. c. corrugatum. 10 as C. p.
martinianum, 11 as C. p. parthenopeum. 12 as D. c. macgintyi. 13 as B. g. cubaniana. 14 described as Q. m. var, mediterranea
Parenzan (1970).
a dual origin for elements of the Tonnacean species now-
living in the eastern Pacific: 1, Survivors of a previously
widespread Pan-Panamian post-Tethyan biota and 2, New
arrivals of trans-Pacific species from the Central Pacific
after the closure of the Central American Seaways.
The apparently rare occurrences of many of the taxa
of both the Tonnacean and non-Tonnacean constituents
found on the islands and/or the mainland may reflect in
part records of species that cannot maintain viable pop-
ulations for any significant duration without frequent
replacement by larval recruitment or by other means of
dispersal from established populations in the mid-Pacific.
One of the major causes of temporal instability of these
populations may result from the harsh environmental
changes caused by major El Nifio events in the eastern
Pacific. Shasky (1985:4) reported a definite decline in
numbers of some of the more common non-Tonnacean
Indo-Pacific species and in the condition of the reef -coral
at Cocos Island following the warming of the local waters
as a result of the severe 1982-83 El Nino episode. He
noted in 1984 the mortality of Pocillopora and found no
living Coralliophila violacea until 1989, when a single
living specimen was taken (Shasky, in litt., August 19,
1990). Unfortunately the collection database for the mol-
lusks provided herein (tables 1-3, Appendix) is not suf-
ficient to determine the long-term influence of the El
Nifio induced changes on the survival of the Indo-Pacific
faunal elements in the tropical eastern Pacific. The El
Nifio disturbances may temporarily benefit some faunal
elements and harm others, e.g.. cause adverse effects on
the communities associated with the coral-reef biotope
(Glynn, 1988; Richmond, 1990; Robinson, 1985, 1987).
For an excellent review of the many factors affecting
the biotic composition of eastern Pacific stony corals and
other tropical marine organisms in time and space, see
Glynn and Wellington (1983:167-207).
The establishment at Clipperton and Cocos Islands of
reproductively viable populations of the Indo-Pacific
faunal element during "normal" conditions (non El Nino
periods) would appear to be the key to the post-Pliocene
dispersal of these faunal consituents to other areas in the
tropical eastern Pacific. The large number of the non-
Tonnacean Indo-Pacific taxa known from Clipperton and
Cocos Islands serves to support this thesis (tables 2, 3).
Twenty-four (85%) of the 28 taxa that are not known
from the mainland have been reported from Clipperton
Island, whereas 8 (28%) occur on Cocos Island. Of the
20 species known from the mainland, 9 (45%) have been
recorded from Clipperton Island, and 10 (50%) occur on
Cocos Island. In contrast, 10 of the 13 (76%) Tonnacean
species occur in the Galapagos Islands, 7 (53%) on Cocos
Island, and 4 (30%) on Clipperton Island. If all the Indo-
Pacific prosobranch gastropods (Tonnacean and non-
Tonnacean) in the eastern Pacific are considered, 55 (90%)
of the 61 taxa are known from the oceanic islands and
27 (44%) are reported from the mainland (table 4). Thir-
ty-seven of the 61 taxa (60%) are recorded from Clip-
perton Island, 25 (40%) from Cocos Island, 18 (29%) from
the Galapagos Islands, 9 (14%) from the Revillagigedo
Islands, and 1 (<1%) from Guadalupe Island (table 4).
The presence at these stepping-stone islands of several
species of hermatypic corals, mostly with western Pacific
faunal affinities (Glynn & Wellington, 1983:176), affords
a habitat for Indo-Pacific molluscan species associated
with the coral reef biotope.
If the established populations of the Indo-Pacific ele-
ment at Cocos and Clipperton Islands serve as a major
source for larval dispersal to the Galapagos Islands and
the west American mainland via the North Pacific Equa-
torial Countercurrent (Abbott, 1966:109; Emerson, 1967:
89; Wellington, 1984:256; Glynn & Wellington, 1983:
167), any prolonged disruption of the reproductive vi-
Page 74
THE NAUTILUS, Vol. 105, No. 2
ability of the populations at these insular sites would
affect the gene replenishment required to reestablish de-
clining or defunct populations on the mainland and in
the Galapagos Islands. The subsurface Equatorial Un-
dercurrent (Cromwell Current), however, is a possible
source for larval transport directly to the Galapagos Is-
lands from the central Pacific. This subsurface current
passes eastward through the Line Islands and could pre-
sumably disperse to the Galapagos Islands tropical or-
ganisms tolerant of temperate subtropical temperatures
(20-25°C at its core of ma.ximum velocity, between 50
and 100 m depth, teste R. S. Scheltema; cf. Glynn &
Wellington, 1983:168). Furthermore, with the com-
mencement of El Nino conditions, the surface flow east-
ward from the Line Islands extends to the Galapagos
Islands and onward toward the mainland (Richmond,
1990). Glynn (1985) estimated the period of uninter-
rupted growth of living Pocillopora in the Gulf of Pan-
ama at about 190 years, based on the death of the pro-
tective coral barriers as a result of warming during the
El Nino event of 1982-83. In terms of geological time,
if only one such El Nifio-related dispersal event occurred
every 200 years, this would result in 5,000 disruptive
episodes per million years and have a significant influ-
ence on the composition of the biota, past and present,
in the eastern Pacific (Richmond, 1990).
ACKNOWLEDGMENTS
In addition to Mesdames Jacqueline DeRoy and Car-
men Angermeyer, I am pleased to acknowledge the co-
operation of a number of people for their aid in the
completion of this paper. Types and other specimens, as
well as data, were kindly provided by Kathie M. Way
of The Natural History Museum, London, Silvard Kool
of the Museum of Comparative Zoology, Harvard Uni-
versity, Terrence M. Gosliner and Elizabeth Kools of the
California Academy of Sciences, M. G. Harasewych and
Raye N. Germon of the National Museum of Natural
History, Smithsonian Institution, James H. McLean, Gale
Sphon and Helen DuShane of the Los Angeles County
Museum of Natural History, Henry W. Chaney of the
Santa Barbara Museum of Natural History, Twila Bratch-
er of Hollywood, California, Donald R. Shasky of Red-
lands, California and Jody Woolsey of Los Angeles, Cal-
ifornia. Specimens used for comparative study were
generously donated by Robert Foster of Abbey Specimen
Shells, Santa Barbara, California and F. Jim Springsteen
of Victoria, Australia. I am indebted to Alan Beu of the
New Zealand Department of Scientific and Industrial
Research, Geology and Geophysics Division, Rudolf S.
Scheltema of the Woods Hole Oceanographic Institution,
Donald R. Shasky and two anonymous reviewers for
kindly reading drafts of the manuscript and offering
many helpful suggestions. I am grateful to my AMNH
colleagues Walter E. Sage, III for technical assistance,
Andrew S. Modell for photographic services, and Ste-
phanie Grooms for word-processing the manuscript.
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Schmeltz, R. L. 1978. A check list of Midwav s molluscs.
Hawaiian Shell News 26(8): 13.
Schmeltz, R. L 1979. Shells from Midway: a further checklist.
Hawaiian Shell News 27(12):11.
Schmelz, G. W. 1989. Notes on the fauna of the Chipola
Formation — .\XXII. A new species of the genus Cyma-
tiiim (Mollusca: Gastropoda). Tulane Studies in Geology
and Paleontology 22(4): 137-139.
Shasky, D. R. 1983a. New records of Indo-Pacific Mollusca
from Cocos Island, Costa Rica. The Nautilus 97(4): 144-
145.
Shasky, D. R 1983b Update on mollusks with Indo-Pacific
faunal affinities in the tropical eastern Pacific. The F"e,stivus
15(2):27. 28.
Shasky, D. R. 1984a. Update on mollusks with Indo-Pacifit-
faunal affinities in the tropical eastern Pacific III. The
Festivus 16(11):I23-124.
Shasky, D. R. 1984b. A preliminary checklist of marine mol-
lusks from Manabi Province, Ecuador. The Western So-
ciety of Malacologists, Annual Report for 1983, 16:25-32
[May 31, 1984].
Shasky, D. R. 1985. Further notes on Cocos Island species.
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Shasky, D R 1987. Update on mollusks with Indo-Pacific
affinities in the tropical eastern Pacific — Part V. The Fes-
tivus 19(6):48-50.
Shasky, D. R. 1989a. Update of molluscs with Indo-Pacific
affinities in the tropical eastern Pacific VIII. The Western
Society of Malacologists, Annual Report, for 1988 21:8
[Marcii 7, 1989].
Shasky, D R. 1989b. .An update on the molluscs from the
Galapagos Islands as listed in the preliminary faunal list
of marine mollusks of the Galapagos Islands by Yves Finet.
Ibid. 21:8-10.
Shasky, D. R. 1989c. Mv last seven years — at Cocos Island.
The Festivus 21(8):72-75.
Skelton, P. W. 1988. The trans-Pacific spread of equatorial
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Charles, M. G. and A. Hallam (eds.) Gondwana and Te-
th\s. Oxford University Press, Geological Society Special
Publication no. 37:247-253.
Smith, J. T 1989. Contrasting niegafaunal and sedimentary
records from opposite ends of the Gulf of California: im-
plications for interpreting its Tertiary history. In: Abbott,
P. L. (ed.). Geologic studies in Baja California. Pacific
Division, Society of Economic Paleontologists and Min-
eralogists, Los Angeles, Book 63:27-36.
Sphon, G. D. 1976. The Mitridae of the Galapagos Islands.
The Nautilus 90(2):63-64.
Springsteen, F. J. and F. M. Leobrera. 1986 Shells of the
Philippines. Carfel Seashell Museum, Manila, 337 p.
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Teredinidae (Mollusca: Bivalvia). Museum of Compara-
tive Zoology, Harvard University, Cambridge, MA, 265 p.
Turner, R. D. 1971. Australian shipworms. Australian Natural
History 17(4):139-145.
Tryon, G W., Jr 188^-1881]. Family Tritonidae, manual of
conchology. Philadelphia, Series 1, 3:(5-6):l-97 [p. 1-64,
1880:65-128, 1881].
Valenciennes, A. 1833. Oiquilles univalves de 1 Amerique
Eqinoxiale, recueillies pendant le voyage de MM. de Hum-
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Bonpland (eds). Voyage aux regions equinoxialesdu Nou-
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von Cosel, R. 1977. First record of Mitra mitra (Linnaeus,
1758) on the Pacific coast of Colombia, South America.
The X'eliger 19(4):422-424.
Waren, A. and P Bouchet. 1990. Laubierinidae and Pisania-
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noidea (Gastropoda: Prosobranchia). The Veliger 33(1):
56-102.
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THE NAUTILUS, Vol. 105, No. 2
Wellington, G. M. 1984. Marine environment and protection
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I7:2-47-26.'3, International Union For (Conservation of Na-
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Geological Survey Professional Paper 306-B, i-iii + 147-
239.
Woodring, W. P. 1966. The Panama land bridge as a sea
barrier. Proceedings of the American Philosophical Society
110(6):425-433.
Woodring, W. J. 1978. Distribution of Tertiary marine mol-
luscan faunas in southern Central America and northern
South America. Universidad Nacional Autonoma de Mex-
ico, Instituto de Geologia, Boletin 101:153-165.
Zinsmeister, W. J. and W. K. Emerson. 1979. The role of
passive dispersal in the distribution of hemipelagic inver-
tebrates, with examples from the tropical Pacific Ocean.
The Veliger 22(l):32-40.
APPENDIX
Geographic records of the 20 shallow-water Indo-Pacific
prosobranch gastropods (other than Tonnacea) known
from the West American Continental Borderland, from
Mexico to Peru including the Near-shore Islands. None
of these taxa is known from the Atlantic Ocean except
Qitoyula madreporarum reported from the Cape Verde
Islands (Saunders, 1976:14).
Not cited in this list are unverified records of west
American taxa that are poorly understood taxonomically,
but may have populations in Indo-Pacific waters. These
records include 1. Phenacovolva ? brevirostris (Schu-
macher) Isla Los Zurrones, Panama, 1 specimen, AMNH
198612, R. E. Hubert, 1979 (see Bertsch and Bibbey,
1982:430 [= ?P. lenoreae Cardin and Walls (1980:1), IsJa
Gobernadora, Panama, 32 specimens]. 2. Bizetiella shas-
kiji Radwin and D'Attilio (1972:347), known from the
Gulf of California, Islas las Tres Marias, Jalisco, Mexico,
and the Galapagos Islands, with a single specimen cited
from Moorea, French Polynesia (Shasky, 1983b). This
may be a west American species that has extended into
the central Pacific. 3. Tripterotyphis lowei (Pilsbry),
known from the Gulf of California to Panama (Keen,
1971), is recognized at the subspecies level from Austra-
lia, Norfolk Island, and the Solomon Islands by D'Attilio
and Hertz (1984), who also report the nominate form
from the Galapagos Islands. 3. Amphithalamus inclusns
Carpenter, known from California to the (iulf of Cali-
fornia (Keen, 1971) [= ?A. trosti Strong and Hertlein,
type locality: Panama], was reported on the basis of 2
specimens from Midway Island by Shasky (1987:49).
I. MEXICO
Conus tessulatus Born, "west Mexican coast, especially
on the offshore islands. " (Keen, 1971:667). Los Frailes,
Baja California Sur, living specimen dredged in 23 m,
IX-1975, ex-S. Bennett coll., AMNH 232136; Small Lsland
off La Paz, 1 fresh specimen, 19.50, ex-A. Barlow coll.,
AMNH 239982.
Littorina pintado (Wood). "Southern Baja California
to Panama" (Keen, 1971:366, as L. puUata Carpenter;
see Reid, 1989:96).
Microdaphne trichodes (Dall). "Puertecitos, head of
the Gulf of California, to Gorgona Island, Colombia."
(Keen, 1971:762, fig. 1848). Cedros Island, west coast of
Baja California, to Puertecitos, Baja California Norte in
the Gulf of California, southward to Punta Ancon, Santa
Elena Peninsula, Peru, intertidal to 146 meters (83 lots
in the LACMNH, teste J. H. McLean).
Quoyula madreporarum (Sowerby). "The southern
part of the Gulf of California and Tres Marias Islands,
Mexico, to Panama." (Keen, 1971:546). Bahia Balandra,
Baja California Sur, 1 dead specimen, IX-1960, Ariel
Exped., ex-H. DuShane coll., AMNH 242842; Cabo Pul-
mo, 8 living specimens on coral (Pocillopora), IV- 1966,
H. DuShane leg., AMNH 242843; Pulmo Reef, in 2-3
meters, 1965, D. R. Shasky, leg. and coll. Bahia Jaltemba,
Nayarit, 7 living specimens, ex-C. Skoglund coll., AMNH
200776. Isla Maria Madre, Nayarit, in 1-5 meters, XII-
1961, D. R. Shasky, leg. and coll. Barra de Navidad,
Jalisco, in 2-3 meters, VII- 1965, D. R. Shasky leg. and
coll.
Rhizochilus aff. R. antipathus Steenstrup, off Sonoran
coast, on antipatharian coral, in 20-60 meters (Foorman,
1981:165).
Terebra affinis Gray. "Gulf of California, Mexico"
(Bratcher and Cernohorsky, 1986:80). Bahia San Luis
Gonzaga, Baja California Norte, 1 living specimen
(Bratcher, 1970:6; Keen, 1971:672, fig. 1519).
Titiscania limacina (Bergh). west of Puerto Pefiasco,
Sonora, sandy beach, C. Skoglund coll. Puerto Lobos,
Sonora (Marcus and Marcus, 1967b:145).
II GUATEMALA
Conus ebraeus Linne. San Jose, Escuintla Department,
1 specimen, tidal flats, 1947, G. Farris leg. (Emerson,
1968:33; AMNH 114575, ex-J. Zager coll.).
III. COSTA RICA
Conus chaldeus (Roding). Isla de Cafio, Puntarenas Prov-
ince, 1 specimen, IV-1977 (Anders, 1978:17, fig. 1); 1
living specimen, III-18, 1972, intertidal, J. H. McLean
leg., LACMNH loc. 72-68. Playas del Coco, Guanacaste
Province, 1 living specimen, IV-24, 1972, intertidal, R.
Koch leg. & coll.
Conus ebraeus Linne. Playas de Jaco, 1 living speci-
men, IV-25, 1975 (Sutton, 1975:79; AMNH 187658, ex-
B. Sutton coll.). Guanacaste Province, 1 living specimen,
T. Dranga leg. (Hertlein and Emerson, 1953:351). Playas
del Coco, Guanacaste Province, 2 living specimens, III-
18, 1965 (Houbrick, 1968:292, MCZ 256447, ex-Hou-
brick coW, fide Emerson, 1968:33). Isla de Cario, Puntare-
nas Province, 1 fragment, intertidal, III-18, 1972, J. H.
McLean leg., LACMNH loc. 72-68. 1 hermit crab spec-
W. K. Emerson, 1991
Page 79
imen, on rock pinnacle, near Isla de Cano, Puntarenas
Province, 21 to 41 meters, III-18, 1972, J. H. McLean
leg., LACMNH loc. 72-65. Isla de Cano, Puntarenas
Province, 9 specimens, IV-8-1977 (Anders, 1978:17, fig.
2).
Microdaphne trichodes (Dall). 10 localities, from Ba-
liia Salinas to Isla de Cano, in 2.4 to 73 meters, LACMNH
coll., 1935 to 1972, teste]. McLean. Bahia Bullena, Golfo
de Nicoya, in 14 meters. III- 1984, D. R. Shasky leg. and
coll.
Mitra mitra (Linne). Isla de Cano, Puntarenas Prov-
ince, 1 living specimen, on sand near coral patches, 7 to
12 meters, III-1972, McLean and Wheeler leg., LACMNH
loc. 72-63 (teste J. H. McLean; Sphon, 1976:63). Playas
del Coco, Guanacaste Province, Golfo de Papagayo, 1
living specimen, intertidal on sand, 1968, Houbrick leg.
IV. PANAMA
Conns chaldeus (Roding). Isla Gobernadora, Golfo de
Montijo, 2 living specimens on rocks, extreme low tide,
V-1979, ex-R. E. Hubert, AMNH 203815 (Emerson, 1983:
122, figs. 7, 8). Isla Canal de Afuera, Golfo de Veraguas,
1 living specimen, on dead coral, snorkling, V-79, 1982,
ex-J. Ernest, AMNH 206683.
Conns ebraeus Linne. Isla Gobernadora, Golfo de
Montijo, 1 fresh specimen, low tide under rock, VII- 1979,
J. Ernest leg., AMNH 242839, ex-H. DuShane coll.
Conus tessulatus Born. Isla Pedro Gonzales, Golfo de
Panama, 1 fresh specimen, 11-1981, ex-R. E. Hubert;
AMNH 203266 (Emerson, 1983:122, figs. 1, 2). Isla Mem-
brillos, Archipielago de las Perlas, Golfo de Panama, 1
fresh specimen, III-1975, e.x-R. E. Hubert, AMNH 183218
(Emerson, 1983:122, figs. 3, 4). Isla Boyarena, Archipie-
lago de las Perlas, 1 living specimen, 1982, ex-J. Ernest,
AMNH 206080 (Emerson, 1983:122, figs. 5, 6); Bahia
Anton Viejo, Isla Los Pajaros, 2 living specimens, IV-6,
1981, H. DuShane leg., AMNH 242837, specimens,
crawling at night on sand exposed at low tide (several
other specimens collected by field party). Isla Canal de
Afuera, Golfo de Veraguas, 1 living juvenile, dredged
15-24 m, 1983, ex-J. Ernest, AMNH 242133.
Cypraea teres Gmelin [= ?alisonae Burgess]. Islas Se-
cas, Golfo de Chiriqui, 1 specimen, II-4, 1935 (Bakus,
1968:94; Emerson and Old, 1968:99, pi. 12, figs. 1-3).
Bahia Honda, Golfo de Chiriqui, 1 specimen 11-21, 1934
(Bakus, 1968:94). Isla Pedro Gonzales, Archipielago de
las Perlas, 2 lots, ex-R. E. Hubert, 11-75, 1 living specimen
under a rock near sand, AMNH 183217; 11-1981, 2 living
specimens, shallow water in coral, AMNH 203809 (Em-
erson, 1983:122, figs. 9, 10); 1 living specimen, low tide,
exposed in coral, 1984, ex-H. DuShane coll., AMNH
242836. Isla Taboga, Bahia de Panama, VI-1979, 1 living
specimen in coral, ex-R. E. Hubert, AMNH 203812. Isla
Los Zurrones, Golfo de Montijo, 2 living specimens, shal-
low water, in coral, VIII- 1979, ex-R. E. Hubert, AMNH
208750. Isla Gobernadora, Golfo de Montijo, 2 living
specimens under rocks around coral, IV-1980, ex-R. E.
Hubert, AMNH 203810; 1 living specimen in 2-3 m, 11-
1985, J. Ernest leg., AMNH 219982; 1 living specimen,
coral heads, 1984, J. E. Ernest leg., ex-H. DuShane coll.,
AMNH 242835. Isla Canal de Afuera, Golfo de Veraguas,
3 specimens, under coral in 1.8 to 3.6 meters, 1982, J.
Ernest leg., AMNH 206081, and elsewhere in the Golfo
de Panama and off Isla Cebaco (teste J. Ernest, in litt.
1983).
Cypraea talpa Linne. Isla Canal de Afuera, Golfo de
Veraguas, 1 living specimen, on dead coral, 1981, ex-J.
Ernest, AMNH 206760 (Emerson, 1983:122, figs. 13, 14);
1 fresh specimen, under coral, 1-1984, ex-H. DuShane
coll., AMNH 242832; Isla Gobernadora, Golfo de Mon-
tijo, 1 living specimen from coral head, V-1984, J. Ernest
leg., AMNH 242831, ex-H. DuShane coll.
Kermia felina (Hinds). Isla Taboga, in 5 meters (Shas-
k\, 1983b:28 as K. maculosa Pease); see Richard (1985:
432).
Microdaphne trichodes (Dall). 9 localities from off Isla
Secas to off^ Bahia Honda, in 9 to 91 meters, LACMNH
coll., 1934 to 1965, teste J. H. McLean.
Mitra mitra (Linne). Los Zurrones, off Isla Cebaco, 1
living specimen, on sandy bottom, 1979, J. Ernest leg.,
AMNH 198611 (Emerson, 1983:122, figs. 11, 12). Isla
Canal de Afuera, Golfo de Veraquas, 1 living specimen
(129 mm by 36 mm) on white sand, near coral, X-1981,
J. Ernest leg., AMNH 206082. Isla Contadora, Golfo de
Panama, 2 living specimens, coral-sand VIII- 1982, P.
Piantino leg. (Anonymous, 1983:15). Isla Mogo Mogo,
Archipielago de las Perlas, 1 living specimen (120 mm
by 34.5 mm) on sand in 1.8 to 3 meters, 1982, snorkling,
J. Ernest leg., AMNH 206083; 1 dead specimen, on beach,
IV-1981, H. DuShane leg., AMNH 242841.
Quoyula madreporarum (Sowerby). "Panama" (Keen,
1971:546). Isla de Boyarena, Archipielago de las Perlas,
intertidally. III- 1977, D. R. Shasky, leg. and coll.
Terebra laevigata Gray. "Panama Bay" (Keen, 1971:
680). Based on type specimen of T. stylus Dall from
"Panama", but west American record questioned by
Bratcher and Cernohorsky (1986:54).
Terebra maculata (Linne). Isla Gobernadora, 1 spec-
imen dredged, 1987, J. Ernest leg., T. Bratcher coll.
Titiscania limacina (Bergh). "Pacific Coast of Pana-
ma" (Marcus and Marcus, 1967a: 124).
V. COLOMBIA
Conus chaldeus Roding. Isla de Gorgona, 1 beach shell.
". . . gathered by the prisoners for shellcraft" (von Cosel,
1977:423).
Cypraea caputserpentis (Linne) Isla de Gorgona, 1
dead specimen, XII-1988 (J. R. Cantera K., 1991).
Cypraea teres Gmelin [= ?alisonae Burgess]. Isla de
Gorgona, "present" (von Cosel, 1977:424; 3 specimens
III-30, 1980, 2 specimens XI-4, 1984, near coral (Cantera,
1986:23). Isla del Malpelo, III-1972 (Birkeland et al,
1975:67).
Microdaphne trichodes (Dall). Isla de Gorgona (Keen,
1971:762; 3 localities, AHF-LACMNH, 1-1935.
Page 80
THE NAUTILUS, Vol. 105. No. 2
Mitra mitra (Linne). Isla de Gorgona, 1 living speci-
men, 11-1972 (von Cosel, 1977:422, figs. 5a, 5b).
Philippic radiata (Roding). Isla de Gorgona, 1 dead-
collected specimen, IX- 1979 (Robertson, 1979:191, figs.
1-3; ANSP 348879).
Quoytda madreporarum (Sowerby). Isla de Gorgona,
"present" (von Cosel, 1977:424).
Reliquiaecava robillardi (Lienard). Isla de Gorgona, 1
specimen on coral (Pavona), LACMNH loc. 35-51 (AHF
412-35), 1-19S5, fide Massin (1987:81). "Gorgona Island",
as Coralliobia cumingii (H. and A. Adams), based on
above specimen (Keen, 1971:546, fig. 1070). Earlier names
for this taxon may be Campulotus cumingii H. and A.
Adams, 1864, and Concholepas (Coralliobia) fimhriata
A. Adams, which has priority and dates from 1854.
Microdaphne trichodes (Dall). Punta Ancon, Santa
Elena Peninsula, intertidal, J. H. McLean, LACMNH
loc. 70-11, III-1970. Manabi Province (Isla de la Plata;
Isla de Salango; and Punta Mala); 1978-1980 (Shaskv
1984b:30).
Psetidocypraea adamsonii (Sowerby). Isla de la Plata,
VI-21, 1979, 1 living specimen, under a rock in 17 meters,
D. Shasky leg. and coll. (Emerson, 1982:13; Shaskv, 1983b:
28).
Quoyula madreporarum (Sowerby). Isla de la Plata,
in 1-3 meters, on coral, V-1979, D. R. Shaskv, leg. and
coll.
Rhizochilus aff. R. antipathus Steenstrup. Isla de la
Plata, Manabi Province, on antipatharian coral, in 30
meters, V-1979, D. R. Shasky, leg. and coll.
VI. ECUADOR
Heliacus trochoides (Deshayes). Ayangue, 30 km north-
east of Punta Santa Elena, 1 living specimen, 11-27, 1971
(Robertson, 1976:13; figs. 1-3; ANSP A6572).
VII. PERU
Microdaphne trichodes (Dall). Isla Lobos de Afuera, 1.5
to 9 meters, J. H. McLean et ai, LACMNH loc. 74-6,
XII-1976.
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Miami, FL 33149
Mr. Richard E. Petit
P.O. Box 30
North Myrtle Beach, SC 29582
Dr. Edward J. Petuch
Department of Geology
Florida Atlantic University
Boca Raton, FL 33431
Dr. David H. Stansbery
Museum of Zoology
The Ohio State University
Columbus, OH 43210
Dr. Ruth D. Turner
Department of Mollusks
Museum of Comparative Zoology
Harvard University
Cambridge, MA 02138
Dr. Geerat J. Vermeij
Department of Geology
Universitv of California at Davis
Davis, CA 95616
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THEC?NAUTILUS
CONTENTS
Volume 105, Number 3
__ Jiugust 27. 1991
'^'^""6 BiolooffS?^^ 0028-1344
LIBRARY
James F. Quinn, Jr.
Lamellitrochus. a nev\' genus of Solariellinae (Gastropoda:
Trochidae), with descriptions of six new species from the
western Atlantic Ocean
81
J. S. Lucas
E. Ledua
R. D. Braley
Tridacna tevoroa Lucas, Ledua and Braley: A recently-
described species of Giant Clam (Bivalvia; Tridacnidae)
from Fiji and Tonga
92
B. A. Marshall Dates of publication and supraspecific taxa of Bellardi and
Sacco's (1873-1904) 'T molluschi dei terreni terziarii del
Piedmonte e della Liguria" and Sacco's (1890) "Catalogo
paleontologico del bacino terziario del Piedmonte " 104
Ronald B. Toll A note on supposed homonyms of Octopus australis Hoyle,
1885, with comments on Octopus campbelli Smith, 1902
(Cephalopoda: Octopodinae) 116
News and Notices 118
THE NAUTILUS 105(3):81-91, 1991
Page 81
Lamellitrochiis, a New Genus of Solariellinae
(Gastropoda: Trochidae), with Descriptions of
Six New Species from the Western Atlantic Ocean
Janie§ F. Quinii, Jr.
Florida Mariiif Research Institute
Department of Natural Resources
100 Eighth Avenue. S E
St. Petersburg, PL 33701-5095, L'SA
ABSTRACT
Lamellitrochiis new genus is erected for eight western Atlantic
species of Solariellinae. The genus is distinguished by having
conicoturbinate shells with angular whorls; strong, usually la-
mellif arm axial riblets that usualK become obscure or are absent
on last v\horls; strong subsutural angulation bearing rounded,
conical, or lamellate tubercles; strong, smooth to tuberculate
peripheral carina; strong, circumbasal carina; intritacalx-like
outer shell layer; usualK niicropustules on earl\ whorls; and
radula lacking a lateroniarginal plate Included in Lamellitru-
chus are the type species, Margarita laniellosa Verrill & Smith,
1880; Solariclla pourtalesi Clench & Aguayo, 19.39; and six new
species: L. inceratus, L. carinatiis. L. sitavis. L. filosiis. L.
fenestratus, and L. bicoronatus.
Key words: Gastropoda; Trochidae; Solariellinae, Lamellitru-
chits ; systematics; new species.
INTRODUCTION
The Solariellinae Powell, 1951, was erected for trochid
genera having short radulae with reduced numbers of
marginal teeth (usually 10 or fewer pairs per row). More
recent work has shown that features of the external anat-
omy (a broad, fringed snout; propodium with lateral
horns; reduced epipodium) are also characteristic of the
subfamily (Fretter & Graham, 1977; Herbert, 1987;
Hickman & McLean, 1990). Although the subfamily is
well defined, Herbert (1987) has shown that animals with
similar shell characters can have different radulae, and
that assignment of these species to genera based solely
on shell characters is sometimes inadvisable. Indeed, this
is reflected in the low number of extant genera (10)
recognized at present (Hickman and McLean, 1990).
Despite the obvious convergence of shell characters of
many species, some species defy ready assignment to
existing genera. Eight such species occur in the western
Atlantic Ocean, and the genus Lamellitrochiis is pro-
posed to include those taxa.
Institutional abbreviations used in this paper are as
follows; DMNH (Delaware Museum of Natural History,
Wilmington. Delaware); FSBC I (Florida Marine Re-
search Institute, St. Petersburg, Florida); MNHN (Mu-
seum National d'Histoire Naturelle, Paris); MORG (Mu-
seu Oceanografico da Fundacao Universidade do Rio
Grande, Rio Grande, RS, Brazil); UF (Florida Museum
of Natural History, L'niversit\ of Florida, Gainesville,
Florida); UMML (Rosenstiel School of Marine and At-
mospheric Science, University of Miami, Miami, Flori-
da); and L'SNM (National Museum of Natural History,
Smithsonian Institution, Washington, DC).
SYSTEMATICS
Family Trochidae Rafinesque, 1815
Subfamily Solariellinae Powell, 1951
Genus Lamellitrochiis Quinn new genus
Margarita: Verrill & Smith in N'errill, 1S80;.392 (partim), Ver-
rill, 1880:.378 (in 1880-1881); 1881:406; 1882:530, .531
ipartim): 1885:527; Dall, 1881:40 (partim).
Margarita (Solariclla): Dall, 1889a:378-382 (partim); 1889b:
164 (partim): 1903:164 (partim). Pilsbry. 1890:307-3.30
(partim ).
Solariclla: DaW. 1927:128-130 (par(im); Johnson, 193471 (par-
tim): Clench & Aguayo, 1939:190, 191; Rice & Kornicker,
1965:117 (partim). Porter, 1974:21 [partim): .Abbott, 1974:
41 (partim): Treece, 1980:5.59,
Solariclla (Machacroplax): .-Abbott, 1974:40 (partim).
Solariclla (Solariclla): Quinn, 1979:37-42 (partim).
Diagnosis: Shell small (usualK <10 mm); subsutural
angulation tuberculate; peripheral carina prominent,
smooth to tuberculate; umbilicus broad; sharp, usually
lamellate axial riblets on early teleoconch whorls; mi-
crosculpture usually of irregular pustules; intritacalx-like
outer shell layer; and oblique, circular to ovate aperture.
Description: Shell small, largest attaining height of about
10 mm but usualK less than 5 mm, conicoturbinate,
umbilicate, nacreous under thin intritacalx and white
porcelaneous layers. Whorls tubular, shouldered, periph-
Page 82
THE NAUTILUS, Vol. 105, No. 3
eralK- carinate, with promiiioiU axial and spiral sculpture.
Axial sculpture of strong, sharp lamellate riblets on first
2-4 teleoconch whorls; later whorls (when present) with
discontinuous renuiants of riblets, most prominent on
spiral carinae, and weak to strong axial rugae; generally
axially oriented micropustules on adapical surface of te-
leoconch whorls. Spiral sculpture above suture of strong
cords or angulations, often tuberculate, and usualK fine
threads between angulations; peripheral carina strongest,
tuberculate, undulate, or smooth. Base weakly convex to
flat, bounded by strong, smooth carina, and with weak
to strong, smooth or finely beaded spiral cords. LImbilicus
broadly open, funnel-shaped, bounded by one (rarely
two) strong, tuberculate spiral cord; walls usually with
spiral cords and axial rugae. Aperture oblique, circular
to ovate.
External anatomy typically solarielline, with broad
snout having numerous, finger-like projections at tip;
long, micropapillate cephalic tentacles; lateral extensions
of the propodium; cephalic lappets lacking. Eyestalks
small, slender, with terminal eye; right postoptic tentacle
lacking. Epipodium reduced; left neck lobe represented
by pair of short tentacles, anterior one near base of left
cephalic tentacle, posterior one near base of anterior
epipodial tentacle; right neck lobe simple or with small
triangular extension, and may be partially fused to basal
half of right e>estalk. Epipodium with 3 pairs of micro-
papillate tentacles; anterior pair long, about midway be-
tween cephalic tentacle and second pair; second pair long
or short, near anterior edge of opercular lobe; posterior
pair long, near posterior edge of opercular lobe; small,
thin, triangular flap may be present between anterior 2
pairs.
Radula short, broad, with about 20-30 transverse rows
of teeth; formula 6-8.4.1.4.6-8. Rachidian tooth cusp
triangular, denticulate, excavated dorsally to accom-
modate next anterior rachidian tooth cusp; base trian-
gular, articulating with inner lateral tooth base. Lateral
teeth 4 pairs, similar to those of llanga (Herbert, 1987).
Marginal teeth 6-8 pairs, denticulate on outer edge near
tip. Lateromarginal plate lacking.
Type species (here designated): Margarita lameUosa
Verrill & Smith, 1880.
Etymology: From the Latin lamella, a little blade, and
trochus, a child s hoop; gender masculine.
Included species: Margarita lamellosa Verrill & Smith,
1880; Solariella pourtalesi ('lench & Aguayo, 1939; La-
mellitrochns cariiiatus Quimi new species; L. inceratus
Quinn new species; L. suatns Quinn new species; L. fi-
losus Quinn new species; L. jenestratus Quinn new spe-
cies; and L. Incoronatus Quirm new species.
Remarks: The shells of species assigned to Lamellitro-
chus are distinguished from all other solarielline genera
by their strongly angular whorl profiles and distinctive
macro- and microsculpture. The angular whorl profile is
caused by the development of a strong subsutural an-
gulation set with rounded, conical, or lamellate tubercles;
a strong peripheral carina that is smooth, undulate, or
tuberculate; a strong basal carina that is usually smooth,
but ma\ be fineK rugose; and a strong, tuberculate cir-
cumumbilical cord. The whorl surfaces between these
angulations are almost flat, and usually have a varying
number of fine spiral lirae. Several species currently as-
signed to Solariella Wood, 1842, also have shells with
angulate whorl profiles: S. triplostephaniis Dall. 1910; S.
patriae Carcelles, 1953; and some forms of i>. cinctus
(Philippi, 1836) and S. intermissa Thiele, 1925. I have
not examined specimens of the first two species, and
illustrations and descriptions (Carcelles, 1953; McLean,
1971) are not detailed enough for me to determine the
species generic assignment; however, I doubt that these
two species are congeneric with the Lamellitrochus spe-
cies. Scanning electron micrographs of shells of S. cinctus
(Fretter & Graham, 1977; fig. 32) and S. intermissa (Her-
bert, 1987: figs. Ill, 112) reveal that neither species has
micropustules on the early whorls.
An extremely thin, calcified shell layer, which ma\' be
chalk} or polished, overlies the outer porcelaneous shell
layer of the last one or two teleoconch w horls. This thin
outer layer may be the calcified periostracum, or intri-
tacalx, described by D'Attilio and Radwin (1971). Nei-
ther an intritacalx nor a periostracum have been reported
for species of other solarielline genera, but they may
have been overlooked; the systematic significance of the
presence or absence of intritacalx in the Solariellinae is
unknown at present. The surface of the first 2-3 whorls
usually has a covering of micropustules. although the
pustules may be restricted to the sutural area. In addition
to those for Solariella cinctus and S. intermissa. SEM
micrographs of the apical whorls of shells of llanga Her-
bert, 1987, Spectamen Iredale, 1924, Minolops Iredale,
1929, and leminolia Finlay, 1927 (Herbert, 1987), and
Minolia Adams, 1860 (Hickman & McLean, 1990) have
been published. Of the species illustrated, only the South
African species Spectamen adarticulatum (Barnard,
1963) (see Herbert, 1987: fig. 124) has discernible mi-
cropustules. Conversely, Lamellitrochus pourtalesi
seemingly lacks micropustules (figure 6). Micropustules
also do not occur on shells of western Atlantic Solariella.
Microgaza Dall, 1881, or Suavotrochus Dall, 1924 (per-
sonal observations), but are present in Pagodatrochus
Herbert, 1989 (Herbert, 1989: fig. 4a,b).
Six of the eight species discussed in this paper have
well-developed, usually lamellate axial riblets that ap-
pear on the first half-whorl, usually immediately follow-
ing the terminal rim of the protoconch. Shells of a ninth
species from Argentina (off Rio de la Plata, USNM 330860)
also have this characteristic, but, because the two spec-
imens are juveniles, this species is not described here.
The illustrations of the first whorls of the genera cited
above (Fretter & Graham, 1977; Herbert, 1987; Hickman
& McLean, 1990) show that shells of other .solarielline
genera lack strong axial riblets on the first whorl, and
that the riblets, present on later whorls are generalK
weaker, more rounded, and more closely spaced than
those of Lanu'llitrochus.
J, F. Quinn. Jr., 1991
Page 83
,f"^
\
Figures 1-6. 1-3. Lamellitrochus lamellosus (Verrill & Smith, 1880) from Eolis Station 351, SE of P'owey Light, Florida, 165 m
(USNM 438440). 1. .\pertural view, 11.6 x. 2. Basal view (oblique), 12.7 x, 3. Protoconch, 118 x, 4-6. Lamellitrochus pourtalesi
(Clench & Aguayo, 1939). 4, 5. Shell from John Elliott Pillsbury Station P-747, 11°46'N, 67°05.7'W, 1,175-1,098 m (UMML
30.6832). 4. Apertural view, shell height 9.1 mm. 5. Basal view, shell diameter 7.6 mm. 6. Protoconch of shell from Alaminos
Station 69A11-86, 21°41'N, 96°51'W, 969-1,079 m, FSBC I 40300, 53 x.
The radiila of Lamellitrochus lacks any trace of a
lateromarginal plate (figures 28-36) and resembles Ilan-
ga Herbert, 1987, "some North Atlantic taxa" (Herbert,
1987:287), and possibly Zetela Finlay, 1927 (Herbert,
1987) in lacking this structure. Herbert (1987) used the
presence or absence of a lateromarginal plate to assign
species with similar shell morphologies to different gen-
era, but he did not speculate on the broader systematic
implications of the character because of the lack of de-
tailed knowledge of many genus-level groups.
The presence of an intritacal.x is the only character
that seems to be unique to this species group, but in
combination with presence of strong axial riblets and
distinct micropustules on the early whorls of most species;
presence of strong, usually tuberculate, subsutural and
peripheral carinae; and lack of a lateromarginal plate of
the radula, it supports establishment of a new genus-
group taxon. Until more detailed analyses of shell and
radular characters of more solarielline species groups are
made, I prefer to treat Lamellitrochus as a genus rather
than a subgenus of Solariella.
Lamellitrochus pourtalesi is illustrated here for com-
parison with the other species, but a complete species
account is found in Quinn (1979).
Key to species of Lamellitrochus.
1. Protoconch diameter greater than 350 fj.m 2
1. Protoconch diameter less than 350 ^ni 3
2. Protoconch diameter 350-375 /urn; axial riblets
present on first half-whorl . . inceratus new species
2. Protoconch diameter 525-550 /im; axial riblets
absent on first half-whorl
pourtalesi (Clench & Aguayo, 1939)
3. Tubercles on subsutural angulation lamellate .... 4
3. Tubercles on subsutural angulation conical or
rounded 5
4. Axial riblets continuous between suture and pe-
riphery; shell wider than high; height less than 3
mm carinatus new species
4. ,\xial riblets not continuous from suture to pe-
riphery; shell higher than wide; height greater
than 3 mm 5
5. Peripheral carina smooth to weakly undulate . .
lamellosus (Verrill & Smith, 1880)
5. Peripheral carina with strong, conical tubercles
filosus new species
6. Subsutural angulation and circumumbilical cord
formed by double rows of rounded tubercles . .
bicoronatus new species
Page 84
THE NAUTILUS, Vol. 105, No. 3
6. Subsiitural angulation and circuiniimbilical cord
formed by single rows of conical tubercles 7
7. Axial riblets present on first half-whorl; peripheral
carina tuberculate suavis new species
7. Axial riblets absent on first half-whorl; peripheral
carina undulate fenestratus new species
Lamellitrochus lamellosus (Verrill & Smith, 1880)
(figures 1-3, 28-30)
Margarita lamellosa Wrriil & Sinitli in \crrill. 1880;.392, 397,
398.
Solariella (Solariella) lamellosa: Quiiiii, 197940, 42 (partini).
Description: Shell small, attaining 4.2 mm height, 3.55
mm width, conicoturbinate, peripherally carinate,
broadly umbilicate, white, nacreous under thin outer
porcelaneous layer. Protoconch 280-310 nm maximum
diameter. Teleoconch whorls about 5, tubular, shoul-
dered, carinate; first two whorls with strong, sharp, axial
lamellae extending from suture to suture; axials becom-
ing discontinuous or obsolete on subsequent whorls; mi-
cropustules rather strong, covering entire surface of first
1.5-2 whorls; subsutural angulation and peripheral ca-
rina appear on second whorl; subsutural angulation be-
coming strong, tuberculate, tubercles lamellate and closely
set; peripheral carina becoming strong, tuberculate, but
usually becoming smooth or irregularly undulate on last
1-2 whorls; as many as 13 weak spiral threads present
between subsutural angulation and peripheral carina; as
many as 12 weak spiral threads present between periph-
eral and basal carinae. Base flat, circumscribed by strong,
smooth spiral carina; as many as 20 weak, smooth spiral
cords present between basal carina and strong, tuber-
culate circumumbilical cord; umbilicus broadly open,
35-45% maximum shell diameter, funnel-shaped, walls
with about 10 fine, weak spiral threads and weak axial
rugae. Aperture oblique, circular, lips thin; peristome
complete. Animal and radula as for genus.
Holotype: USNM 44738, height 3.0 mm, width 3.0 mm.
Type locality: Off Martha's Vineyard, Massachusetts,
Fish Hawk Station 871, 40°02'54"N, 70°23'40"W, 210 m.
Other material examined: More than 100 lots in USNM,
DMNH, FSBC I; see Quinn (in press) for listing.
Remarks: Shells of Lamellitrochus lamellosus are most
similar to those of L. filosus in shape and in having
lamellate tubercules on the subsutural angulation; how-
ever, the former species is absolutely and relatively larger
(4.2 mm vs. 3.15 mm with about 5 whorls) but has a
smaller protoconch (280-310 nm ds. 315-325 jum). The
tubercles on the peripheral carina of shells of L. filosus
are stronger, conical, and more widely spaced than those
of L. lamellosus. and the spiral threads in the intercarinal
spaces are weaker and the axial threads are stronger in
L. filosus than in L. lamellosus. The shell of L. suavis is
also very similar in shape and general sculpture, but the
subt hiral tubercles are rounded, the periphery has rath-
er strung tubercles, the micropustules are not as promi-
nent and are concentrated near the suture, and the in-
tercarinal spiral sculpture is weaker than in L. lamellosus.
Lamellitrochus lamellosus is distributed along the Unit-
ed States east coast from Massachusetts southward through
the Florida Keys, and in the Gulf of Mexico off western
Florida, Texas, and the Campeche Bank in depths of 50-
250 m (usually in 100-200 m).
Lamellitrochus carinatus new species
(figures 7-12, 34, 35)
Description: Shell very small, attaining 2.75 mm height,
2.90 mm width, depressed conicoturbinate, peripherally
carinate, broadly umbilicate, white, nacreous under thin
outer porcelaneous layer. Protoconch 275-300 nm max-
imum diameter. Teleoconch whorls about 4, tubular,
shouldered, carinate; first two whorls with strong, sharp,
axial lamellae extending from suture to suture, but end-
ing at or just above peripheral carina on subsequent
whorls; micropustules rather strong, covering entire sur-
face of first 2 whorls; subsutural angulation and periph-
eral carina appear on second whorl, peripheral carina
becoming strong, smooth to weakly undulate; 1-7 weak
spiral threads present between subsutural angulation and
peripheral carina; 0-4 weak to strong spiral threads may
be present between peripheral and basal carinae; whorl
surface with microscopic pustules. Base weakly convex,
circumscribed by strong, smooth spiral carina; 1-8 (usu-
ally 3-6) weak to strong, smooth to weakly undulate
spiral cords present between basal carina and strong,
tuberculate circumumbilical cord; umbilicus broadly
open, 35-45% maximum shell diameter, funnel-shaped;
walls with 1-7 weak spiral cords and strong axial rugae
that radiate onto inner part of base, forming strong,
radially elongate tubercles on circumumbilical cord. .Ap-
erture oblique, circular, lips thin, peristome complete.
Holotype: DMNH 179393, height 2.50 mm, width 2.70
mm.
Type locality: SW of Egmont Key, Florida, 366-421
m; J. Moore, collector.
Paratypes: DMNH 186768, 22 specimens; USNM
859408, 1 specimen; FSBC I 37633, 1 specimen; UF
161754, 1 specimen; all from same lot as holotype. —
USNM 859431, 13 specimens; Eolis Station 360, off Fow-
ey Light, Miami, Florida, 146 m.— -USNM 438401, 3
specimens; Eolis Station 321, off Western Dry Rocks,
Florida, 119 m.— 31, DMNH 179392; 100 mi SW of
Egmont Kev, Florida; D. Steger, collector. — FSBC I
24259, 5 specimens; MAFLA Station 2746-41, 27°07'N,
84°13'W, 122 m; 23 August 1977; box corer.— USNM
500245, 37 specimens; State Universitv of Iowa Barbados
Station 29, off Lazaretto, 165-183 m.— USNM 500266,
State University of Iowa Barbados Station 77, off Cable
Station, 73-137 m.— MORG 26529, 1 specimen; Marion-
Dufresne Cruise MD-55, Station DC-61, 20°29'S,
29°18'W, 63 m; May 1987; dredge.— MNHN uncata-
logued, 1 specimen; Marion-Dufresne Cruise MD-55,
J. F. Quiiin, Jr., 1991
Page 85
8
,«:'V
H^
11
'^^^^
.12f^
Figures 7-12. LamellUwchtis carinatus new species. 7-9. Paratypes from SW of Egmont Key, Florida, 366-421 m (DMNH
186768). 7. Apertural view, 15 x 8. Basal view, 17 x . 9. Protoconch, 134 x 10-12. Paratvpe from Marion-Dujresne Cruise
MD-55, Station DC-61, 20°29'S, 29°18'W, 63 m (MORG 26529) 10. Apertural view, 15.3 x . 1 1. Basal view, 14.9 x 12. Protoconch,
82 X.
Station DC-59, 20°.30'S, 29°19'W, 52-60 m; May 1987;
dredge.
Other material examined: More than 50 lots in USNM
and FSBC I; see Quinn (in press) for listing.
Remarks: Shells of Lamellitrochus carinatus are dis-
tinguished from those of all other Lamellitrochus species
principally in being distinct!) wider than high and usu-
ally having strong and continuous a.xial riblets on all
whorls. This species also has the smallest shells, with
heights of 2.50-2.75 mm and widths of 2.75-2.90 mm;
adult shells of all other species exceed 3 mm in height.
Variation in shell morphology occurs between northern
and southern populations, but this is a reflection of the
frequency of individual character states because the same
variations also occur within individual populations. Shells
of the northern (Florida) populations tend to have smooth
to weakly undulate peripheral carinae; 2-4 strong, often
unequally sized basal spiral cords; fewer, usually weaker
spiral threads in the spaces between the subsutural an-
gulation and peripheral carina (usually 1-3), and the
peripheral and basal carinae (0-1); and narrower spaces
between the axial riblets (about 0.15-0.20 mm apart on
the third whorl). Shells from Barbados and Brazil usually
have distinctly undulate peripheral carinae, the crests of
the undulations often sharp; the basal spiral cords tend
to be more numerous (usually 3-6, sometimes 7 or 8)
and often equally sized and spaced; spiral threads in the
intercarinal spaces are relatively numerous (usuallv 3-4
in each) and distinct; and the axial riblets are more widely
spaced (about 0.25 mm apart on third whorl). This spe-
cies is the most widely distributed of all Lamellitrochus
species, ranging from off North Carolina southward to
both sides of Florida; off Cuba, Puerto Rico, Antigua,
and Barbados; and Trinidade Island off eastern Brazil,
usually in depths of 100-200 m.
Lamellitrochus fenestratus new species
(figures 13-15)
Description: Shell small, attaining 3.35 mm height, 2.95
mm width, conicoturbinate, peripherally carinate,
broadly umbilicate, white, nacreous under thin outer
porcelaneous layer. Protoconch 275-300 ^m maximum
diameter. Teleoconch whorls 4.6, tubular, shouldered,
carinate; first 0.5 whorl w ith 4 spiral cords, increasing to
6 on second whorl; whorls 0.5-2 with initially weak,
progressively stronger, rounded, axial riblets extending
from suture to suture; axials becoming discontinuous or
obsolete on subsequent whorls; micropustules weak, re-
stricted to sutural area of first 2 whorls; subsutural an-
gulation and peripheral carina appear on third whorl,
peripheral carina becoming strong, undulate; 5-7 spiral
threads present between subsutural angulation and pe-
ripheral carina; 2-5 weak spiral threads and distinct axial
rugae present between peripheral and basal carinae. Base
flat, circumscribed by strong, smooth spiral carina; 6-9
weak, smooth spiral cords present between basal carina
Page 86
THE NAUTILUS, Vol, 105. No. 3
14
20
Figures 13-21. 13-15. Lamellitrochns fenestrates new species, paratype from off Barbados, 18.3 m (USNM 859429). 13. Apertural
view, 14 X, 14. Basal view, 14 x, 15. Protoconch, 84 x, 16-18. Lamellitroclms filosus new species, holotype from off English
Harbor, Antigua (USNM 5002.30). 16. Apertural view, 14.3 x. 17. Basal view, 14.2 x^ 18. Protoconch, 91 x, 19-21. Lamcllitwchus
suacis new species, holotvpe from off Barbados (USNM 500224). 19. Apertural view, 13.7 x . 20. Basal view, 13.1 x , 21. Protoconch,
92 X. '
and strong, tuberculate circumiimhilical cord; umbilicus
broadly open, 30-35% ma.ximum shell diameter, funnel-
shaped, walls with about 4 spiral cords and strong axial
rugae. Aperture oblique, circular, lips thin; peristome
complete.
Holotype: USNM 859428, height 3.15 nmi, width 3.00
mm.
Type locality: Off Barbados, Blake Station (data im-
recorded), 183 m.
Paratypes: USNM 859429, 4 specimens; from same lot
as holotype. — USNM 500202, 1 specimen; State Univer-
sity of Iowa i: rbados Station 62, off Pelican Island, 229
m— IISNM 859430, 1 specimen; State University of Iowa
Barbados Station (data unrecorded), "deep".
Remarks: Shells of L. fenestratus are most similar to
those of L. bicoronaius but differ in having only a single
row of conical tubercles forming the subsutural angu-
lation rather than a double row of rounded tubercles, in
having a stronger peripheral carina w ith stronger tuber-
cles, and in ha\ iiig a single rather than double circumum-
bilical cord. Shells of L. fenestratus are also similar to
tho.se of L. filosus and L. suavis in size, protoconch width,
and general shape, but differ from both in lacking axial
ribletson the first half-whorl; in having conical tubercles
in the subsutural angulation as in L. filosus, but a more
J. F. Quinn, Jr., 1991
Page 87
undulate peripheral carina rather than that of L. suavis;
and the spiral threads between the subsutural and pe-
ripheral carinae are stronger in L. fencstratus than in
either L. filostis or L. suavis.
Lamellitrochus filosus new species
(figures 16-18)
Description: Shell small, attaining 3.15 mm height, 2.85
mm width, conicoturbinate, peripherally carinate,
broadK umbilicate, white, nacreous under thin outer
porcelaneous la\er. Protoconch 315-325 jum maximum
diameter. Teleoconch whorls about 5, tubular, shoul-
dered, carinate; first two whorls with strong, sharp, la-
mellate axial riblets extending from suture to suture;
axials becoming discontinuous or obsolete on subsequent
v\ horls; micropustules weak, restricted to sutural area on
first 2 whorls; subsutural angulation and peripheral ca-
rina appear on second whorl; subsutural angulation be-
coming strong, tuberculate, tubercles lamellate; periph-
eral carina becoming strong, tuberculate; 13-15 very
weak spiral threads present between subsutural angula-
tion and peripheral carina; as many as 11 very weak
spiral threads present between peripheral and basal ca-
rinae; spiral threads intersecting equally sized axial
threads forming file-like surface sculpture. Base flat, cir-
cumscribed by strong, smooth spiral carina; as many as
20 weak, smooth spiral cords present between basal ca-
rina and strong, tuberculate circumumbilical cord; um-
bilicus broadly open, 40-45% maximum shell diameter,
funnel-shaped, walls with 6-8 or more very weak spiral
cords and weak, lamellate axial rugae. Aperture oblique,
circular, lips thin; peristome complete.
Holotype: USNM 500230, height 2.75 mm, width 2.50
mm.
Type locality: State University of Iowa Antigua Station
116, off English Harbor, "deep".
Paratypes: USNM 859425, 4 specimens; from same lot
as holot\pe.
Remarks: Shells of L. filosus are most similar to those
of L. lamellosus. particularly in having lamellate tuber-
cles on the subsutural angulation, but differ in being
smaller (3.15 mm vs. 4.20 mm, respectively, at whorl
4.9), in having strong, conical tubercles on the peripheral
carina, and in having much weaker spiral threads be-
tween the subsutural carination and peripheral carina.
Shells of L. filosus are also similar to the holotype of L.
suavis, but have lamellate rather than conical tubercles
on the subsutural angulation, and a fine, file-like sculp-
ture on the surface of the last whorl.
Lamellitrochus .mavis new species
(figures 19-21)
Description: Shell small, attaining 3.35 mm height, 2.90
mm width, conicoturbinate, peripherally carinate,
broadly umbilicate, white, nacreous under thin outer
porcelaneous layer. Protoconch 325 fxm maximum di-
ameter. Teleoconch whorls 4.6, tubular, shouldered, car-
inate; first two whorls with strong, sharp, lamellate, axial
riblets extending from suture to suture; axials becoming
obsolete on subsequent whorls; micropustules weak, scat-
tered over entire surface of first 2.5 whorls; subsutural
angulation and peripheral carina appear on second whorl;
subsutural angulation becoming strong, tuberculate, tu-
bercles bluntly conical; peripheral carina becoming
strong, tuberculate; 5 weak spiral threads present be-
tween subsutural angulation and peripheral carina; 3 or
4 extremely weak spiral threads present between pe-
ripheral and basal carinae on last whorl. Base flat, cir-
cumscribed by strong, smooth spiral carina; 10 weak,
smooth spiral cords present between basal carina and
strong, tuberculate circumumbilical cord; umbilicus
broadly open, 35% maximum shell diameter, funnel-
shaped, walls with weak axial rugae. Aperture oblique,
circular, lips thin; peristome complete.
Holotype: USNM 500224, height 3.35 mm, width 2.90
mm.
Type locality: State University of Iowa Barbados Station
(data unrecorded), "deep".
Remarks: The shell of L. suavis resembles a miniature
L. inceratus in shell shape and sculpture, but is smaller
and has a smaller protoconch (325 ixm vs. 350-375 ^m,
respectiveh ). Shells of L. filosus are similar to that of L.
suavis in size, protoconch width, and tuberculate pe-
ripheral carina, but differ in having lamellate rather than
conical tubercles on the subsutural angulation, and in
having fine, crowded axial threads that interact with
equally fine spiral threads to produce a fine, file-like
sculpture on the vn horl surface.
Lamellitrochus bicoronatus new species
(figures 22-24)
Description: Shell small, attaining 3.55 mm height, 3.20
mm width, conicoturbinate, peripherally carinate,
broadK umbilicate, white, nacreous under thin outer
porcelaneous la\er. Protoconch 290-300 ^m maximum
diameter. Teleoconch whorls 4.3, tubular, shouldered,
carinate; first whorl with 5 strong spiral cords with ad-
ditional threads appearing on second whorl and increas-
ing in number on subsequent whorls; 3 cords, 1 just below
suture, 1 above midvvhorl, and 1 below midwhorl
strengthening on second whorl, forming tuberculate sub-
sutural and undulate shoulder angulations, and weakly
undulate peripheral carina, respectively; low, rounded
axial riblets appear at end of first whorl, becoming stron-
ger and more lamellate on whorls 2-3, and becoming
low, broad folds between suture and shoulder angulation
on last whorl; micropustules weak, scattered over entire
surface of first 2 whorls, but most concentrated near
suture; last whorl with 3-4 spiral threads between sub-
sutural and shoulder angulations, 5-7 spiral threads be-
tween shoulder angulation and peripheral carina, and 4-
5 spiral threads and distinct axial rugae between pe-
bS
THE NAUTILUS, Vol. 105, No. 3
23
Figures 22-27. 22-24. LameUitrochus bicoronatus new species, paratype from off Barbados, 183 m (USNM 859436). 22. .\pertural
view, 13.1 X. 23. Basal view, 14.3 x. 24. Protoconch, 119 x. 25-27. LameUitrochus inceratus new species. 25, 26. Paratype
from Cerda Station G-967, 24°15'N, 82°26'W, 499-503 m (UMML 30.8062). 25. Apertural view, shell height 8.3 mm. 26. Ba.sal
view, shell diameter 7.0 mm 27. Protoconch of paratype from Albatross Station 2644, 25°40'N, 80°00'W, 353 m (USNM 859432),
81 X.
ripheral and basal carinae. Base flat, circumscribed by
strong, smooth spiral carina; 6-8 weak to strong, smooth
spiral cords present between basal carina and 2 strong,
tuberculate circumumbilical cords; umbilicus broadly
open, about 40% ma.ximum shell diameter, funnel-shaped,
walls with 2-4 weak spiral cords and weak axial rugae.
Aperture oblique, circular, lips thin; peristome complete.
Holotype: USNM 859435, height 3.55 mm, 3.20 mm.
Type locality: Off Barbados, Blake Station (data un-
recorded), 183 m.
Paratypes: 4 specimens, USNM 859436; from same lot
as holotype.
Hcmarks: Shells of L. bicoronatus differ from those of
all other Larnetlitrochiis species in having most of the
first teleoconch whorl devoid of axial riblets, and having
a double spiral row of strong, rounded tubercles forming
the subsutural angulation, and a similar double spiral
row of tubercles circumscribing the umbilicus.
LameUitrochus inceratus new species
(figures 25-27, 36)
Calliostoma tiara: Dall, 1889a;.365 (partim).
Solariella amalrilis: Dall, 1889a;378, 379 (partim).
Solariella (Solariella) lamellosa: Quinn, 197940-42, figs. 61.
62 (partim).
Description: Shell small, attaining 8.2 mm height, 7.3
mm width, conicoturbinate, peripherally carinate,
broadly umbilicate, white, nacreous under thin outer
porcelaneous layer. Protoconch 350-375 /um maximum
diameter. Teleoconch whorls 6.0, tubular, shouldered,
carinate; first vvhorl with strong, sharp, axial lamellae
extending from suture to suture, rapidh becoming re-
stricted to subsutural angulation and peripheral carina
on subsequent whorls; micropustules rather weak, cov-
ering entire surface of first 2 whorls; first whorl with 2-
4 weak spiral threads, two of which later interact with
axial lamellae to form tuberculate subsutural angulation
and peripheral carina; 0-5 very weak spiral threads pres-
ent between subsutural angulation and peripheral carina;
spiral threads absent between peripheral and basal ca-
rinae. Base flat, circumscribed b\ strong, smooth spiral
carina; 5-11 weak, flat, smooth spiral cords present be-
tween basal carina and strong, tuberculate circumum-
bilical cord; center part of base often smooth; umbilicus
broadly open, 45-50% maximum shell diameter, funnel-
shaped, walls with 0-4 weak spiral cords and w eak axial
rugae. Aperture oblique, circular, lips thin; peristome
complete.
J. F. Quinii, Jr., 1991
Page 89
^
'^fid^SS^ '^^^^^^1
1
1
^^f
^p^
^
H
3^
^i^s^
g
3
i^f^^i
^
u
Figures 28-36. Radulae of Lamellitrochus species. 28-30. Lamellitrochus lamellosus (Verill & Smith, 1880) from specimen in
Figure 1. 28. .Anterior segment, 339 x, 29. Middle segment, 339 x. 30. Anterolateral area, 690 x, 31-33. Lamellitrochus
pourtalesi (Clench & .Aguayo, 1939) from specimen in Figure 6. 31. Middle segment, 240 x , 32. Rhachidian and laterals, 600 x .
33. Anterolateral area, 600 x. 34, 35, Lamellitrochus carinatus new species, from paratype from Eolis Station 360, off Fowey
Rocks, Florida, 183 m (USNM 859431). 34. Anterior segment, 600 x .^ 35. Same, 1,200 x. 36. Lamellitrochus inceratus new
species, from paratype in Figure 27, right anterior fragment, 169 x.
Hololype: USNM 94946, height 7.05 mm, width 6.90
mm
Type locality: Off Cape Florida, Key Biscavne, Florida,
Albatross Station 2644, 25°40'N, 80°bo"W, 353 m.
Paratypes: USNM 108141, 1 specimen; USNM 754272,
1 specimen; Albatross Station 2668, 30°58'30"N,
79°38'30"W, 538 m.— UMML 30.8051, 2 specimens;
Gerda Station G-300, 26°16'N, 79°30'W, 640 m.— UMML
30.8025, 1 specimen; Gerda Station G-4, 25°49'N,
79°59.5'W, 256 m.— UMML 30.8338, 2 specimens; John
Elliott Pillsbitry Station P-1309, 25'"44.5'N, 79°50.0'W,
311 m.— UMML 30.7565, 1 specimen; Gerda Station
G-830, 25°40'N, 79°59'W, 342 m.— USNM 859432, 6
specimens; USNM 330559, 4 specimens; same locality as
holotype.— UMML 30.8099, 2 specimens; Gerda Station
G-23, 25°32'N, 79°44'W, 768 m.— UMML 30.8042, 2
specimens; Gerda Station G-857, 25°22'N, 80°03'W, 194-
186 m. — UMML 30.7538, 1 specimen; Gerda Station
G-834, 25°15'N, 80°10'W, 86-79 m.— UMML 30.7914, 1
specimen; Gerda Station G-1035, 24°34.7'N, 80°58.6'W,
254-358 m.— UMML 30.7770, 8 specimens; Gerda Sta-
tion G-970, 24°24'N, 82°08'W, 512 m.— UMML 30.8063,
1 specimen; Gerda Station G-969, 24°18'N, 82°33'W, 269-
402 m. — L'MML 30.7644, 1 specimen; Gerda Station
G-968, 24°17'N, 82°34'W, 499-503 m.— UMML 30.8062,
6 specimens; Gerda Station G-967, 24°15'N, 82°26"W,
499-503 m — IIMML 30.8065, 1 specimen; Gerda Sta-
tion G-1099, 24°12.5'N, 82°50'VV, 622 m.— UMML
30.8058, 1 specimen; Gerda Station G-861, 24°08'N,
8r36'W, 514-558 m.— USNM 421840, 5 specimens;
Waldo Schmitt Station 69, off Dry Tortugas, 455-655
m.— USNM 94947, 4 specimens; Blake Station 2, 23°14'N,
82°25'W, 1,472 m.— USNM 94948, 4 specimens; Blake
Station 21, 23°02'N, 83°13'W, 525 m.— USNM 859433,
1 specimen; Blake Station (data unrecorded), Yucatan
Channel, 1,170 m.— USNM 94058, 3 specimens; Alba-
tross Station 2150, 13°34'45"N, 8r21'10"W, 699 m.—
I'.i-
THE NAUTILUS, Vol. 105, No. 3
USNM 94106, 1 specimen; Albatross Station 2135,
19''55'5S"N, 75''47'07"W, 457 m.— USNM 429872, 5
specimens; USNM 429873, 2 specimens; USNM 429895,
2 specimens; Johnson-Smithsonian Deep-Sea Expedition
Station 94, 18°37'45"N, 65°05'00"W, 549-860 m.—
UMML 30.8339, 1 specimen; John Elliott Pillsbtiry Sta-
tion P-929, 15°29.5'N, 61°11.5'\V, 457-503 m.— USNM
94949, 2 specimens; Blake Station 211, 14°28'40"N,
61°06'08"W, 653 m.— UMML 30.8340, 8 specimens; John
Elliott Pillsbury Station P-905, 13°46.3'N, 61°05.4'W,
384-963 m.— UMML 30.8341, 13 specimens; John El-
liott Pillsbury Station P-904, 13°45.5'N, 6r05.7'\V, 201-
589 m.— UMML 30.8342, 1 specimen; yo/in Elliott Pills-
bury Station P-903, 13°44'N, 61°03.1'W, 231-430 m.
Remarks: Shells of Lamellitrochus inceratus are similar
in shape and sculpture to that of L. suavis, but are much
larger (up to 8.2 mm vs. 3.35 mm, respectively) and have
larger protoconchs (350-375 ^m vs. 325 nm, respective-
ly). Shells of L. pourtalesi (figures 4-6) are most similar
to those of L. inceratus, but are somewhat larger (to 10.3
mm), have much larger protoconchs (525-550 ^m), have
a narrower umbilicus (about 35% vs. 45-50% of shell
width, respectively), lack axial riblets on the first 0.5
whorl, the aperture is ovate rather than circular, and the
tubercles on the subsutural angulation and peripheral
carina are sharply conical and axialK elongate rather
than rounded and spirally elongate as in L. inceratus.
Lamellitrochus inceratus occurs off southern Georgia;
in the Straits of Florida from off Cape Florida to the
Dry Tortugas, and off Havana and Bahia Honda, Cuba;
in the Yucatan Channel; off Old Providence Island; off
southeastern Cuba; off northern Puerto Rico; and the
Lesser Antilles from Dominica to St. Lucia. This species
is usually collected in depths of about 250-550 m.
ACKNOWLEDGMENTS
I thank Riidiger Bieler (DMNH), the late Gilbert L. Voss
(UMML), and Robert Hershler and M. G. Harasewych
(USNM) for access to the collections under their care,
and Jose H. Leal for bringing the Brazilian specimens of
Lamellitrochus carinatus to my attention and providing
the SEM micrographs of one of those specimens. Earnest
Truby (Florida Marine Research Institute) assisted with
SEM operations and provided the final prints of the other
SEM micrographs. Marjorie Myers (Florida Marine Re-
search Institute) word-processed the various versions of
the manuscript. Thomas H. Perkins and William G. Ly-
ons (both Horida Marine Research Institute), and two
anonymous reviewers, provided valuable critiques of
previous drafts of this paper.
LITERATURE CITED
Abbott, H T 1974. .American seasheils, 2nii cil \ an Nostrand
Reinhold, New York, NY. fifi3 p
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from Japan. Annals and Maga/inc of Natural Histors.
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Oarcelli'S, .\. R. 19.53. .Nuevasespeciesde gastropodos inarnios
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Clencli, W J, and C G. .'\guayo. 1939. Notes and descriptions
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Dali, VV. H. 1881. Reports on the results of dredging, under
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Dall, W. H. 1889a. Reports on the results of dredging, under
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Coast Survey steamer "Blake", Lieut-Commander C. D.
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THE NAUTILUS 105(3):92-103, 1991
Page 92
Tridacna tevoroa Lucas, Ledua and Braley:
A Recently-described Species of
Giant Clam (Bivalvia; Tridacnidae) from
Fiji and Tonga
J. S. Lucas
Zoology Department
James Cook University
Townsville, Q. 4811, Australia
E. Ledua
Fisheries Division
Ministry of Primary Industry
Suva, Fiji
R. D. Braley
Zoology Department
James Cook University
Townsville, Q. 4811, Australia
ABSTRACT
Tridacna tevoroa Lucas, Ledua and Braley is the eighth extant
species of giant clam. It has only been found in the eastern Lau
Islands of Fiji and the northern Vava'u and Ha'apai islands of
Tonga. It inhabits relatively deep water (20+ m) compared to
other giant clam species and is apparently intolerant of shallow
conditions. Tridacna tevoroa occurs sympatrically with T. de-
rasa (Roding) and appears to be most closely related to it, both
belonging in the subgenus Persikima Iredale. The two species
are readily separated by the presence in T. tevoroa of a rugose
mantle, prominent guard tentacles on the incurrent aperture,
thinner valves, and colored patches on the shell ribbing near
the umbonal area. Tridacna tevoroa has some Hippopus-like
characters: shell pigmentation, absence of byssal gape, lack of
lateral mantle projection, and absence of hyaline organs in the
mantle. These necessitate modification of Rosewater's diag-
nostic features distinguishing the two genera of extant giant
clams, Tridacna Bruguiere and Hippopus Lamarck.
Key words: Giant clam, Tridacnidae, Tridacna. taxonomy,
key, Hippopus.
INTRODUCTION
Rosewater (1965) reviewed the confused taxonomy of
giant clams (Order Veneroida, Superfamily Tridacnacea,
Family Tridacnidae), recognizing six extant species in
two genera, Hippopus Lamarck, 1799 and Tridacna Bru-
guiere, 1797. He subsequently described a further Hip-
popus species (Rosewater 1982). Thus, after Rosewater's
pivotal work, the family consisted of seven extant species,
H. hippopus (Linnaeus, 1758), H. porcellanus Rosewa-
ter, 1982, T. gigas (Linnaeus, 1758), T. dcrasa (Rciding,
1798), T. squamosa Lamarck, 1819, T. maxima (Roding,
1798) and T. crocea Lamarck, 1819.
This taxonomic revision of the family Tridacnidae was
very timely, as it preceded the period of intense interest
in giant clams in recent years. Before 1970, there was
little research interest in giant clams, despite it being
known that giant clams are unique among bivalve mol-
lusks in having hypertrophied siphonal tissues packed
with symbiotic algae (zooxanthellae) (Lucas, 1988). Then,
in the 1970's, there was a surge of research. This resulted
partly from academic interest in their unique features,
but mainly from interest in farming giant clams. It was
recognized that giant clam stocks were being heavily
over-exploited through most of their geographic ranges
and that some species had been fished to extinction in
particular regions. Furthermore, it was found that giant
clams grow more rapidly than previously envisaged
(Munro & Gwyther, 1981). Farming them as a means of
supplying the demands for giant clam meat and other
products, and thus taking pressure off natural popula-
tions, became a desirable and feasible proposition.
Rosewater's taxonom\ of giant clams has been widely
accepted and validated through use in species' identifi-
cations in numerous studies. However, at an international
meeting on giant clams in 1988, the possibility of a rare,
new species in Fiji was raised (Lewis & Ledua, 1988).
This new taxon was subsequently found also in Tonga
and recentlv described as Tridacna tevoroa by Lucas et
al. (1990).
This paper provides further description of T. tevoroa,
including morphometric comparison with T. derasa. It
also considers the affinities of the new species within the
family Tridacnidae.
GENERA AND SUBGENERA OF
TRIDACNIDAE
In order to consider the affinities of T. tevoroa within
the family Tridacnidae, it is necessary to review Rose-
water's diagnoses of Hippopus and Tridacna. and sub-
genera of the latter. Rosewater (1965) recognized three
subgenera of Tridacna: Tridacna s.s. Bruguiere, 1797
(containing T. gigas). Persikima Iredale, 1937 (contain-
ing T. derasa). and Chametrachea Morch, 1853 (con-
taining T. squamosa, T. maxima, and T. crocea). As the
diagnostic features for the various taxa are not clearly
J. S. Lucas et ai, 1991
Page 93
Table 1. Diagnostic features of Tridacna and Hippopus frm
HoM-uater (1965), with additions and modifications
Feature
Tridacna
Uippupus
Hyaline organs in
mantle surface
B\ssal orifice
Tightly fitting teeth
along byssal orifice
Posterior adductor
muscle scar loca-
tion
Outer demibranch of
ctenidia
Klongate- triangular
shape'
Strawberry blotches
on earl\ portion of
shell
Mantle projects later-
ally over shell
margins^
Distinct ventral re-
gion of shells'
present (usually absent
present)'
well defined (poor- poorK defined
ly-well dehned)'
absent present
si bcliiiu! inid-liiic central
complete-incom-
plete
no
not present (usual-
1\ not present)'
yes (usually)'
not present
complete (?)-
yes (in large
specimens)'
present
present
' Modification reiiuired to accommodate T. tcvuroa.
- It is presumed that Rosewater was referring to the presence
or absence of a distal food groove in describing the <iuter demi-
branch as "complete' or "incomplete "; however, J. Norton
(personal cornnnmication) has found that the two Hippopus
species have no food groove on the outer demibranch and are
thus "incomplete .
■'Small H. porcellanus are semicircular.
^From Lucas (1988),
^ From this paper.
This region includes the byssal orifice and is outlined by
the ventral-most pair of rib-like radial folds. It is ap-
proximately ovate and flat in small specimens. In large
specimens it is heart-shaped and concave. This feature
is recognized in the common name for H. hippopus,
"horse's hoof " clam. The ventral region is closely applied
to the substrate in small juveniles attached by their bys-
sus. Thus, the long body axis of Hippopus juveniles is
strongly angled to the substrate in a characteristic fash-
ion.
While T. squamosa is included in the subgenus Cha-
metrachea, it shows little if any tendency to bore into
hard substrates in our experience. Furthermore, some
large specimens of T. squamosa lose their byssal attach-
ment, in accordance with the general pattern of large
size and loss of byssal attachment in tridacnids. All tri-
dacnid species are attached to their substrate with byssal
threads as juveniles; however, as they grow, the byssus
is apparently insufficiently strong to maintain the clam
in its vertical orientation (mantle uppermost, promi-
nently exposed to sunlight) against strong wave forces.
Instead, the clam greatly thickens the umbonal regions
of the valves. This basal weight, together with strongly
convex curvature of the valves near the umbos, provides
a strong righting force when the large clam is displaced
onto its side.
The features by which the other two members of Cha-
metrachea (T. crocea and T. maxima) differ from other
tridacnids, i.e., small size, permanent byssal attachment,
wide byssal orifice (which must be concealed or predators
will penetrate it), substrate erosion and numerous shell
scutes (for wedging and eroding), are a suite of func-
tionally interrelated features.
set out in Rosewater (1965), the features he considered
to be diagnostic are compiled in tables 1 and 2.
One feature of the Hippopus species was not empha-
sized by Rosewater. This is the well-defined ventral re-
gion of the valves, distinctly angled to the adjacent shell
curvature and almost parallel to the lateral body axis.
TERMINOLOGY
The terminology used in various publications to describe
the anatomical orientation of giant clams is inconsistent.
This arises from alternative interpretations of the clam's
unusual body form, with its greatly expanded siphonal
region. Yonge (1936, 1980) followed some earlier authors
Table 2. Diagnostic features of the three subgenera of Tridacna. derived from Rosewater (196.5)
Feature
Tridacna ,s.,'
Pcrsikinia
Chametrachea
Outer demibranch
Adult shell length
Byssal orifice
Adult attached to substrate
Coral boring
Lateral shape of valves
Deep radial folds on shell
Shell scutes
Guard tentacles on incurrent aperture
Ctenidia extend dorsallv
with food groove
large (to H- m)
small
no
no
equilateral
yes
only in small juveniles
no
no
w/o food groove
medium (to 500-(- mm)
small
no
no
umbo displaced backward
no
only in small juveniles
yes
yes
w/o food groove
small-med (150-400 mm)
med. -large
yes'
yes/no^
ca. equil. — umbo displaced
forward
no
yes
yes
no
' Not always for large T. squaniuau.
- Not for T. squamosa.
Faee 94
THE NAUTILUS, Vol. 105, No. 3
in considering that the expan.sion ot the giant clam's
siphonal region had occurred through the mantle/shell
rotating 180° about the viscero-pedal mass. In this view,
the relationship between the body and shell of giant
clams is the reverse of other bivalves. Stasek (1962, 1963)
concluded that there was no evidence for an altered
functional relationship between the body and the man-
tle/shell in giant clams; their unusual morphology results
from growth largely in a posterior direction, in contrast
to typical growth that is largely in a ventral direction.
In this paper we follow the anatomical orientation
terminology of Stasek (1962, 1963), and this is illustrated
in figure 1. In this terminology, ventral, posterior, etc.,
relate to the equivalent regions in typical bivalve mol-
lusks, not to the orientation of the adult tridacnid. This
terminology poses problems in referring to the relative
positions of structures within the body, e.g., to say that
one structure is posterior to another is quite imprecise
as the posterior region of the tridacnid occupies almost
70% of its total circumference (Stasek, 1962; figure 2).
Terms such as '"above", "below", "before" and "behind",
or equivalents, are suggested for the four directions in
sagittal plane, which would otherwise be "mid-posteri-
or", "anterior", "postero-dorsal" and "postero-ventral",
respectively. Stasek (1962) used terms such as "foremost"
and "upward".
The anatomical orientation terminology used by Yonge
(1936, 1980) and Rosewater (1965, 1982), and in Lucas
et al. (1990), is illustrated in figure 2. It reflects the
orientation of the adult tridacnid, but has no relationship
to the equivalent regions of typical bivalve mollusks.
Shell dimensions measured in this study are illustrated
in figures 3-5.
Tridacna (Persikima) tevoroa
Lucas, Ledua and Braley 1990
(figures 6-11, 16)
Tridacna sp. (cf. T. derasa), Lewis and Ledua 1988;82.
Tridacna tevoroa Lucas, Ledua and Bradley 1990:1.
Diagnosis: Lateral shell outline asymmetrical, with long
hinge line and short ventral margin. Byssal orifice poorly
defined, but without interlocking teeth. Shell with 6 or
7 rib-like radial folds usually having strawberry colored
patches near umbonal region, producing a striped ap-
pearance. Mantle surface rugose, with numerous pro-
tuberant lobes. Long, broad and prominent guard ten-
tacles on incurrent aperture.
Description: Shell: Reaching 550 mm length; suboval
in lateral outline, but strongly asymmetrical, with dorsal
region much longer than ventral region; shell height :
length ratio ca. 0.6:1 in large (>300 mm) specimens.
Umbo angle > 150°; umbo slightly opisthogyrous. Valves
moderately inflated, thin at margins but distinctly thick-
ened in umbonal region in large specimens. Shell colored
gra\-white externally, usually with strawberry colored
patches on rib-like radial folds in umbonal area (extend-
ing up ca. 100 mm from umbo), giving ribs a striped
appearance. Strawberry patches are not evident in some
old specimens (including holotype), apparently lost
through erosion or obscured by calcareous overgrowths
of umbonal region (c/. figure 1 1 ). Primary radial sculp-
ture consisting of 6 or 7 prominent folds, plus 3 short
thin ridges near ligament. Prominent rib-like folds grad-
ing into undulating folds near shell edge in large spec-
imens. Secondary radial sculpture consisting of fine ribs,
much stronger in fold interstices than on radial folds;
fine ribs not evident in distal shell region of large spec-
imens. Apparent remains of eroded scutes often present
on rib-like folds (figure 11) from umbo to extent of straw-
berry colored striping; occurring on white regions be-
tween colored patches. Other concentric sculpture lim-
ited to fine growth lines. Upper shell margins weakly
undulating, with 5 rounded medially-projecting pro-
cesses, representing extremities of rib interstices; upper
shell margins often thin and sharp (figure 8). Length of
hinge line ca. 0.6 x shell length. Umbo strongly back-
ward of mid-point of shell length. Cardinal tooth and
cardinal socket (figure 7) not strongly developed, even
in large specimens. Lateral hinge tooth little more than
a ridge. Ventral shell margin somewhat irregular and
slightly convex. Byssal orifice poorly defined; without
teeth; with ca. 5 rows of small plicae in byssal orifice
region. Interior of shell white; gloss\' distal to pallial line,
dull medial to pallial line. Hinge plate with pale brown
tinge. Pallial line strongly indented in ventral region.
Posterior adductor muscle scar large and strongK' indi-
cated with lines and ridges; ca. 70 mm high and 55 mm
wide in 435 mm shell length specimen; sited behind mid-
point of shell length but in front of umbo on long shell
axis. Posterior pedal retractor muscle scar adjacent to
foremost border of posterior adductor scar and much
smaller, <0.2x area of posterior adductor scar. Soft
parts: Mantle of subdued color, brown, gray-brown or
green-brown, uniform in color or mottled; without stri-
ations, but sometimes with pale lines following mantle
surface contours (figure 10). Mantle surface rugose, with
numerous protuberant lobes (figure 10). Mantle tissue
with few iridophores, without hyaline organs, and with
most zooxanthellae situated close to mantle surface (fig-
ure 16). Mantle not or barely extending lateralK over
shell edges. Expanded mantle broad and exposed by wide
gaping of valves. Incurrent aperture with long, broad
guard tentacles, usually white, but sometimes brown
(presumably through presence of zooxanthellae), often
projecting prominently out from aperture. Byssus absent
and foot vestigial in all large specimens (300-1- mm shell
length). Ctenidia demibranchs extending forwards be-
yond proximal oral grooves for ^-'t of their length.
Outer demibranch of similar size or slightly less than
depth of inner demibranch; outer demibranch may be
upwardly reflected in some regions and the ctenidial axis
is exposed in some regions. Inner demibranch with food
groove and outer demibranch without. Long distal oral
grooves lead from ctenidia to labial palp bases.
Morphometries: In comparing the morphometries of
T. tevoroa and T. derasa. seven parameters were mea-
POSTERIOR
... ( = siphonal region)
POSTERO-VENTRAL
(= hindmost)
POSTERO-DORSAL
(= foremost)
VENTRAL
( = pedal gape)
DORSAL
DORSAL
(= ligament region)
ANTERIOR
(= foremost end of pedal gape to umbo)
POSTERIOR
ANTERIOR
VENTRAL
"Fs" platform
umbonal region thickness I
\^ J umbo
Figures 1-5. Diagrams of anatomical terminology for giant clams and shell dimensions measured. 1. Orientation terminology
used in this paper (following Stasek, 1962), 2. Alternative orientation terminology (after Rosewater, 1965, 1982; Yonge, 1936, 1980).
3. Lateral view of valve. 4. Upper view of valves. 5. Cross-section of valve.
Page 96
THE NAUTILUS, Vol. 105, No. 3
**
--».<.
"?'"}'=»
Lir^
■:f^i
/
'■r'
W
#
J. "^^ .;>_.,■►«'' ~^ _
#•
Figures 6, 7. Lateral views of T. tevoroa holotype left valve. 6. E.xtenial surface, 7. Internal surface.
J. S. Lucas et ai. 1991
Page 97
Figures 8, 9. T. tevoroa holotype. 8. Upper view. 9. Umbonal view.
sured: 1. shell length (figure 3); 2. shell height (figure 3);
3. shell width of conjoined valves (figure 4); 4. shell weight;
5. underwater shell weight; 6. umbonal region thickness
(maximum distance through the umbonal region) (figure
5); 7. distal shell thickness (measured 25 mm in from the
margin of the medial radial fold). Only measurements
1, 2 and 3, and sometimes 7 could be made on living
specimens. The shell weight (4) in living specimens was
estimated from the underwater weight of the whole an-
imal, which was assumed to be entirely due to shell (=5),
the soft tissues having approximately neutral buoyancy.
shell weight = 1.611 x underwater weight
This relationship between shell weight and underwater
shell weight was found to be very regular (1.611 ± 0.008
s.d.) in T. derasa and T. tevoroa.
Parameters 2-7 were related to shell length (1), which
was used as the general measure of size. Least squares
regressions were calculated for each of parameters against
shell length for each species (table 3). The slopes and
intercepts for each pair of regressions, and their standard
errors, were compared by t-tests (2-tail) for statistically
significant differences. There were no significant differ-
ences between the intercepts for each pair of regressions.
Slopes were significantly different (P < 0.05) between
species for three of the regressions: shell width, umbo
Page 98
THE NAUTILUS, Vol. 105, No. 3
^:;^.'^
Figures 10, 1 1. Live T. tevoroa specimens. 10. Rugose mantle; ex, excurrent aperture; t. conspicuous guard tentacles on incurrent
aperture. 1 1. Umbonal regions. Note that the umbonal region of the specimen in the foreground is heavily eroded and encrusted,
showing little striping of the radial ribs, but striping and apparent eroded scutes are clear on the rear-left specimen (arrowed);
striping is clear on the rear-right specimen (arrowed).
J. S. Lucas et al., 1991
Page 99
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200 301) 100 r,00 600
Shell leiifith (nun)
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20
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200 300 (00 500
Shell length (mm)
600
40
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200 300 400 500
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Figures 15, 16. Cross-sections of surface layers of mantle
tissues. 15. r. derasa. 16. T. tevoroa. i, iridophores; z, zooxan-
tfiellae. Scale bars = 100 micrometers.
thickness and shell weight versus shell length (table 3).
These data are figured in figures 12-14.
Types: Holotype: Australian Museum, Sydney, Reg.
No. C164071. Whole specimen with soft tissue preserved
Figures 12-14. Morphometric comparisons of T. tevoroa (tri-
angles and dashed regression line) and T. derasa (circles and
continuous regression line). 12. Shell width versus shell length.
13. Umbo thickness versus shell length. 14. Shell weight versus
shell length.
Page 100
THE NAUTILUS, Vol. 105, No. 3
Table 3. Morphometries of T. tevoroa and T. derasa. Coefficients and standard errors (SE) for "a" (slope) and "b' (Y axis intercept)
for least squares regressions of Y = aX + b, where X is shell length (nun). Sample numbers (n) for T. tevoroa and T. derasa ranged
from 18-35 ami 9-16, respectively. P, significance level ol correlation; vs, very significant (P "C 0 01); *, slope coefficients differ
significantl)' between species at P < 0.05 level.
7'
ridacna tevoroa
Tridacna derasa
Y
a/b
Coeff.
SE
P
Coeff
SE
P
Shell width (mm)
a
0.307*
0.071
vs
0.613*
0.061
vs
b
45.0
29.4
0.2
-13.77
25.34
0.6
Shell height (mm)
a
0.548
0.075
vs
0.657
0.058
vs
b
23.05
31.20
05
-11.42
24.57
0.6
I'mbo thickness (mm)
a
0.026*
0.034
0.5
0.324*
0.034
vs
b
40.8
14.8
0.02
-54.93
1449
vs
Shell margin thickness (mm)
a
0 0072
0 0145
0.6
0031
0.008
vs
b
5.52
6.26
0.4
1.46
3.41
0.7
Shell weight (kg)
a
0.0649*
0.0053
vs
0.114*
0.012
vs
b
-17.17
219
vs
-30.29
4.78
vs
Underwater weight (kg)
a
0.0418
0 0040
vs
0.0716
0 0115
vs
1,
-11 33
1,59
\s
-IS 95
471
\s
in formalin (figures 6-9). Shell length 485 mm. Collected
at Cakau Tabu Reef, Southern Lau islands, eastern Fiji,
17°40'S, 178°32'W, A. Lewis and Filipe Viala, 26 January
1986. Paratypes: 1. Fiji Museum, Suva, Reg. No. C-001.
Whole specimen with soft tissue preserved in formalin.
Shell length 328 mm. Collected at Vatoa Island, Southern
Lau islands, eastern Fiji, 19°49'S, 178°13"W, E. Ledua
and M. Tavisa, 23 November 1989. Habitat details: at
24 m depth on gentle slope outside the reef on the south-
ern side of the island; only low, hard corals were present
and scattered in low numbers; very clear water. 2. Na-
tional Museum of Natural History, Smithsonian Insti-
tution, Washington, DC, USA, Reg. No. USNM 859439.
Specimen consisting of valves only. Shell length 389 mm.
Collection and habitat details as for paratype 1. 3. Mu-
seum of Tropical Queensland, Townsville, Australia, Reg.
No. MO 17791. Specimen consisting of valves only. Shell
length 413 mm. Collection and habitat details as for
paratype 1.
Type locality: Cakau Tabu Reef, Lau Islands, Fiji.
Distribution: Lau Islands, Fiji (known from Cakau Tabu,
Komo Island, Namuka Island, Vatoa Island and Vuata
Vatoa, 17°40'S-19°56'S, 178°13'W-178°32'W; Lewis &
Ledua, 1988). Islands within the Ha apai and Vava u
groups, Tonga.
Habitat: From the limited collection details, the habitat
of T. tevoroa appears to be along outer slopes of leeward
reefs, in very clear, oceanic water at 20-30 m. Tridacna
derasa is also found in this kind of habitat, and some
specimens may occur down to these depths in very clear
water.
Material examined for morphometries: T. tevoroa:
Fiji, Lau Island group: 440 mm, single valve; 392 mm,
single valve; 413 mm, valves only; 548 mm, valves only;
389 mm, valves only; 375 mm, live specimen; 354 nmi.
live specimen. Tonga, Vava'u Islands: 560 mm, live spec-
imen; 326 mm, live specimen; 420 mm, live specimen;
376 mm, live specimen; 313 mm, live specimen. Tonga,
Ha'apai Islands: 408 mm, valves only; 412 mm, valves
only. T. derasa: Fiji, Makogai Island: 538 mm, valves
only; 425 mm, valves only; 370 mm, valves only; 368
mm, live specimen; 348 mm, live specimen; 317 mm,
live specimen. Tonga, Tongatapu: 490 mm, valves only;
376 mm, valves only; 434 mm, valves only; 435 mm,
valves only; 323 mm, valves only; 294 mm, valves only;
291 mm, valves only; 474 mm, valves only. Tonga, Fale-
vai, Vava'u group: 415 mm, live specimen; 293 mm, live
specimen; 332 mm, live specimen; 471 mm, live speci-
men. Australia, Great Barrier Reef, Bramble Reef: 435
mm, valves only; 500 mm, valves only; 490 mm, valves
only; 440 mm, valves only; 470 mm, single valve; 303
mm, live specimen; 415 mm, live specimen; 435 mm,
live specimen; 340 mm, live specimen; 430 mm, live
specimen; 450 mm, live specimen; 465 mm, live speci-
men; 500 mm, live specimen.
Etymology: The species name is derived from the clam's
Fijian name, "tevoro " (devil), with a latin termination
(nominative, feminine).
Comparison: Tridacna tevoroa is most like T. derasa
in general appearance, with the latter species occurring
in shallower reef areas at all the localities where T. te-
voroa has been collected. The similarity of these species
goes beyond superficial appearance: in those features by
which the subgenus Persikima is distinguished from oth-
er subgenera of Tridacna (table 2). T. tevoroa belongs
with T. derasa in Persikima. It is, however, readily dis-
tinguished from T. derasa b\ a number of features re-
lating to both hard and soft anatomy (table 4). Among
these distinguishing features, the most conspicuous are
the rugose mantle, prominent guard tentacles on the
J. S. Lucas ct ai, 1991
Page 101
Table 4. Features which distinguish T. tevoroa from T. derasa.
(These features apply to specimens ca 250 mm shell lenijth
and greater.)
Feature
T. tfcoroa
T. dcrd.'ici
Byssal orifice
Umhniial region of
rib-like primary
f.ilds
Mantle
Iridophores in man-
tle tissue
Hyaline organs
Mantle projecting fie-
yond shells
Guard tentacles on
incurr. apert.
Demibranchs of cte-
nidia
Cardinal tocitli
poorK defined
striped with
color, appar-
entl\ with
eroded scutes
rugose, with
protuber-
ances
low density
(mantle dull
colors)
absent
not or barely
long, broad,
prominent
similar size
small, but clearly
defineil
uncoloretl. with-
out scutes
smooth, plain
high density
(mantle often
bright colors)
present
short, tlun, not
prominent
outer smaller
not prominent \er\ prominent
incurrent aperture, thinner valves, and colored patches
on the shell ribbing of T. tevoroa.
The three shell parameters versus shell length for which
regression slopes differ significantly between T. tevoroa
and r. derasa (table 3) are interrelated, i.e., they reflect
the thinner shell of T. tevoroa compared to T. derasa.
There is, however, little difference in shell thickness be-
tween medium-sized specimens of the two species, ca.
300 mm shell length; interspecific differences become
pronounced in the largest (and probably oldest) speci-
mens, 450+ mm (figures 12-14). The sharpness and thin-
ness of the shell margins of T. tevoroa, noted by Lewis
and Ledua (1988), are certainly evident in some speci-
mens (figure 8), but its shells soon thicken behind the
margin and there is no significant difference in regression
slopes between the species for distal shell thickness, 25
mm behind the margin, versus shell length.
Stasek (1962) described the ctenidia of some tridacnid
species from Eniwetok Atoll in detail, distinguishing T.
derasa and H. hippopus from other species in having
food grooves on both inner and outer demibranchs. Tri-
dacna derasa was further distinguished from other spe-
cies in having similar-sized demibranchs, with the outer
demibranch upwardly reflected, exposing the ctenidial
axis. In view of this reported distinctiveness of the cte-
nidia of r. derasa, it would seem to be appropriate to
compare these structures in T. tevoroa and T. derasa.
However, Rosewater (1965) described T. derasa as hav-
ing an "incomplete" outer demibranch, and recent stud-
ies of tridacnid morphology by J. Norton (personal com-
munication) have revealed that T. derasa specimens from
Fiji and Tonga, at least, have short outer demibranchs
without food grooves. The degree of upward reflection
of the outer demibranch and exposure of the ctenidial
axis vary between specimens; they ma\' partlv reflect
handling during dissection. Thus, the distinctiveness of
ctenidial morphology in T. derasa observed by Stasek is
questionable, and comparisons with T. tevoroa are made
on the basis of Rosewater s and Norton's observations.
Tridacna tevoroa has some features typical of Hip-
popus in the Hippopus /Tridacna separation based on
Rosewater (1965) (table 1 ). Its byssal aperture is indistinct
and it has reddish markings on the external surfaces of
the shells. Its dull mantle color, lack of hyaline organs
and mantle posture (only reaching the shell margins but
offset by wide gaping of the valves) are also like Hip-
popus but unlike Tridacna, sensu lata. However, these
features of T. tevoroa are never completely like those of
Hippopus. Its byssal aperture differs from the Hippopus
species in not having interlocking teeth, nor does it have
the distinctive "horse-shoe" ventral surface of the valves.
It is unlikely that attached juveniles of T. tevoroa orient
to the substrate in the characteristic manner of Hippopus
juveniles. While T. tevoroa shares shell color patches with
the Hippopus species, its colored stripes on the radial
ribbing are unlike the irregular color patches of Hip-
popus species. The mantle of T. tevoroa is distinct from
that of all other tridacnids.
It is conceivable that T. tevoroa represents a transi-
tional stage between these two genera, the divergence
of which goes back in the fossil record at least to the
lower Miocene (Rosewater, 1965). However, it comfort-
ably fits with T. derasa in the subgenus Persikima and
we do not propose to give it separate subgeneric or ge-
neric status.
One consequence of the discovery of T. tevoroa is the
need to make some modifications to the characters dis-
tinguishing Hippopus and Tridacna (table 1).
Small juveniles of T. tevoroa are unknown, but they
are likely to be distinctly unlike T. derasa and any other
tridacnid. This is because of the colored stripes on the
valve ribs and also the probable presence of scutes on
the ribs; these characters being present up to about 150
mm shell length. The distinctive appearance is lost in
larger specimens.
Remarks: Lewis and Ledua (1988) hypothesized that
the localized distribution of the "tevoro " clam in eastern
Fiji may represent a "relict population on the eastern
edge of both its own former range and that of the ap-
parently more abundant and successful T. derasa". Their
prediction, that other populations might be found, was
soon confirmed by collections in Tonga. The known dis-
tribution of T. tevoroa is still, however, limited to several
island groups over about 300 km distance in the south-
western Pacific. At none of these localities is it abundant:
in the \'atoa area, Fiji, Lewis and Ledua (1988) estimated
its relative abundance compared to T. derasa was on the
order of 1:50. It thus must be regarded as the giant clam
species most vulnerable to extinction. The previous bear-
er of this unfortunate distinction was H. porcellanus,
with a relatively restricted distribution centered on the
Page iU2
THE NAUTILUS, Vol. 105, No. 3
southern Philippines (Lucas, 1988), and a high demand
within the international shell trade despite CITES ({Con-
vention on International Trade in Endangered Species)
protection. Fortunately, the deep habitat of T. tevoroa
partially protects it from human predation, but there is
still some exploitation. This is evident from the fact that
there are local names for T. tevoroa in the three island
groups where it occurs (Lucas et a/., 1990) and there are
anecdotal reports of its consumption, i.e.. one diver on
N'ava'u Island, Tonga, reported having collected and eat-
en about 20 specimens. Fortunately, there is no major
trade in giant clam shells in Fiji or Tonga.
Tridacna tevoroa has only been collected from rela-
tively deep-water habitats for giant clams (20—30 m
depth). Their mantle tissue contains symbiotic zoo.xan-
thellae like other tridacnids, and presumably photosyn-
thetic products from these symbionts are used in their
nutrition. However, there are low light levels and a shift-
ed light spectrum at the depths they inhabit, and prob-
ably the extent to which autotrophy supports the clam's
nutrition will be reduced. Thus, T. tevoroa is expected
to have morphological adaptations to facilitate using the
low light intensities at these depths. Extending the mantle
laterally over the shell margins to increase its exposed
surface, as in other Tridacna species, is surprisingly not
shown by T. tevoroa. However, it achieves a large mantle
surface area through the mantle protuberances and wide
gaping of its valves. The dull mantle color is due to low
densities of iridophores (iridocytes) (c/. figures 15 and
16), with the brown coloration of zooxanthellae showing
through. Iridophores in giant clams contain thin reflect-
ing platelets (Kamishima, 1990). They are particularly
common near the mantle tissue surface in brightly col-
ored tridacnids, including some T. derasa (figure 15).
Most zooxanthellae are situated below the iridophore
layer in T. derasa. In T. tevoroa the zooxanthellae also
occur close to the mantle surface (figure 16); clusters of
zooxanthellae may occur in small elevations of the mantle
surface. The paucity of iridophores and shallow
distribution of zooxanthellae in the mantle of T. tevoroa
clearly reflect the need to maximize light reaching the
zooxanthellae. It has been shown that zooxanthellae pho-
to-adapt to low light levels (Mingoa, 1988) and this may
be important for T. tevoroa.
It seems that T. tevoroa are not only adapted to rel-
atively deep conditions, but they are intolerant of shallow
habitats. In early collections by Fiji Fisheries staff, these
clams were transported on the decks of Fisheries vessels
and kept in shallow conditions {ca. 3 m) at Makogai
Island. Rapid mortality occurred until the clams were
moved into deeper water {ca. 14 m). Solar UV radiation
penetrates clear tropical waters sufficiently to have sig-
nificant biological effects down to 20 m and it is possible
that T. tevoroa lacks or has insufficient of the U\' screen-
ing agents that protect organisms inhabiting the shallow
regions of coral reefs, e.g., Dunlap et al. (1988). Also, it
may have insufficient enzymes that deal with the toxic
oxygen levels generated by photosynthesis at high light
levels {e.g., Shick & Dykens, 1985).
KEY TO THE TRIDACNID SPECIES
There are now eight extant species of giant clams and
the key to species given by Lucas (1988) needs to be
expanded and modified to incorporate T. tevoroa.
1 Byssal orifice region of opposed valves with in-
terlocking teeth; distinct region of shell around
byssal orifice, outlined by ventral-most pair of
prominent radial ribs; mantle, when fully ex-
tended, not projecting laterally beyond shell
margins Hippopus ... 2
Byssal orifice of opposed valves without inter-
locking teeth; no distinct ventral region of shell
outlined by prominent radial ribs; mantle, when
fully extended, usually projecting laterally be-
yond shell margins Tridacna ... 3
2(1) Shells thick and strongly ribbed, with reddish
blotches in irregular bands; incurrent apertures
without guard tentacles H. hippopus
Shells in specimens less than about 200 mm shell
length not thick nor strongly ribbed and with
only faint reddish blotches; incurrent apertures
with guard tentacles H. porcellanus
3(1) Shell length of large specimens >550 mm,
sometimes greater than 1 m; with about four
elongate, interdigitating projections of each dis-
tal shell margin, being most elongate and acute
in large specimens; shell without scutes, except
for some tubular projections near umbo in very
small juveniles; mantle brownish, with numer-
ous iridescent blue-green circles T. gigas
Shell length rarely >550 mm; without elongate,
interdigitating projections on each distal shell
margin; mantle variably colored, without iri-
descent blue-green circles 4
4(3) Shell length up to 550 mm, occasionally larger;
upper region of large shells plain, without scutes
or strong ribs; hinge usually longer than half
shell length 5
Shell length usually <400 mm; upper shell re-
gion with scutes or eroded scutes; hinge equal
to or less than half shell length 6
5(4) Rib-like radial folds on shell without colored
patches; mantle without protuberances; incur-
rent aperture with inconspicuous guard tenta-
cles T. derasa
Rib-like radial folds on shell usually striped with
colored patches near umbo; mantle with pro-
tuberances; incurrent aperture with conspicuous
guard tentacles T. tevoroa
6(4) Shell approximately symmetrical about umbo
in lateral view, with hinge about half shell length;
scutes large and well-spaced both within and
between the radial rows; lateral distance be-
tween scutes in adjacent rows usually about the
same as scute width; byssal aperture narrow to
moderately wide; not embedded into substrate;
mantle usuallv of subdued and mottled color;
J, S. Lucas et ai, 1991
Page 103
incurrent aperture with distinct guard tentacles
T. squamosa
Shells usually asymmetrical about umbo in lat-
eral view, with hinge less than half shell length;
scutes usually low and often eroded, set close
together both within radial rows and between
rows; byssal aperture moderately wide to wide;
embedded or partK embedded into substrate;
mantle brightly colored; incurrent aperture with
indistinct guard tentacles 7
7(6) Shell length < 150 mm; shells not strongly asym-
metrical about umbo in lateral view; byssal ap-
erture wide; scutes eroded away except near
shell margin; occurs deepK embedded in reef
substrate T. crocea
Shell length of large specimens often > 150 mm;
shell often strongly asymmetrical about umbo
in lateral view; byssal aperture moderately wide
to wide; scutes present in substantial part of
upper shell region; occurs partially embedded
in reef substrate T. maxima
ACKNOWLEDGEMENTS
This research was part of an international project funded
by the Australian Centre for International Agricultural
Research (Project No, 8733). We thank the Fisheries
personnel in Fiji and Tonga, and local divers, for their
assistance with collecting; Dr John Norton for assistance
with the histology, the micrographs and for reading an
early version of the manuscript; Dr Winston Ponder for
advice and for providing the photographs of the holo-
type; Dr Ross Alford for advice on statistics and Mr
Jeremy Barker for assistance with the figures.
LITERATURE CITED
Dunlap, W, C, B E. Chalker, and W. M Bandaranayake.
1988. Ultraviolet light absorbing agents derived from
tropical marine organisms of the Great Barrier Reef Pro-
ceedings of the 6th International Coral Reef Symposium
3:89-93.
Kamishima, Y. 1990. Organization and development of re-
flecting platelets in iridophores of the giant clam, Tridacna
crocea Lamarck. Zool. Sci. 7:63-72.
Lewis, A. D. and E. Ledua. 1988, A possiljle new species of
Tridacna (Tridacnidae: Mollusca) from Fiji. /;i; Copland,
J. VV. and J. S. Lucas (eds.). Giant clams in Asia and the
Pacific. ACIAR Monograph No. 9, Australian Centre for
International Agricultural Research, Canberra, ACT, p.
82-84.
Lucas, J. S. 1988. Giant clams: description, distribution and
life-history. In: Copland, J. W. and J. S. Lucas (eds. ). Giant
clams in Asia and the Pacific. ACI.\R Monograph No 9,
Australian Centre for International Agricultural Research,
Canberra, ACT. p. 21-32
Lucas, J S., E. Ledua, and R. D Brale) 1990. A new species
of giant clam (Tridacnidae) from Fiji and Tonga. ACIAR
Working Paper No 23, Australian Centre for International
Agricultural Research, Canberra, ACT, 8 p.
Mingoa, S. S. M. 1988. Photoadaptation in juvenile Tridacna
gigas. In: Copland, J. W. and J. S. Lucas (eds). Giant
clams in Asia and the Pacific. ACIAR Monograph No. 9,
Australian Centre for International Agricultural Research,
Canberra, ACT, p. 145-150.
Munro, J L and J Gw>ther. 1981. Growth rates and mari-
culture potential of tridacnid clams. Proceedings of the
4th International Coral Reef Symposium 2 633-636.
Rosewater, J, 1965. The family Tridacnidae in the Indo-
Pacific. Indo-Pacific Mollusca 1:347-394.
Rosewater, J. 1982. A new species of Hippopiis (Bivalvia:
Tridacnidae). The Nautilus 96:3-6.
Shick, J. M. and J. A. Dykens. 1985. Oxygen deto.xification
in algal-invertebrate symbioses from the Great Barrier Reef
Oecologia 66:33-41.
Stasek, C, R. 1962. The form, growth, and evolution of the
Tridacnidae (giant clams). Arch. Zool. Exp, Gen. 101:1-
40.
Stasek, C. R, 1963, Orientation and form in the bivalved
Mollusca. J. Morph. 112:195-214.
Yonge, C. M. 1936. Mode of life, feeding, digestion and sym-
biosis with zooxanthellae in the Tridacnidae Scient Rep.
Gt. Barrier Reef Exped. 1:283-321.
Yonge, C. M. 1980, Functional morphology and evolution in
the Tridacnidae (Mollusca: Bi\alvia: Cardiacea), Rec. Aust,
Mus. 33:735-777.
THE NAUTILUS 105(3):104-115, 1991
Page 104
Dates of Publication and Supraspecific Taxa of
Bellardi and Sacco's (1873-1904) "I moUuschi dei terreni
terziarii del Piemonte e della Liguria" and
Sacco's (1890) "Catalogo paleontologico
del bacino terziario del Piemonte"
B. A. Marshall
National Museum of New Zealand
PO Box 467, Wellington, New Zealand
ABSTRACT
Dates of publication are determined for the serial parts of
Bellardi and Sacco's "I molluschi dei terreni terziarii del Pie-
monte e della Liguria " and its largely overlooked companion
series, the latter containing the earliest valid introductions of
many taxa. Genus group taxa introduced by Sacco (1890) in
the rare work "Catalogo paleontologico dei bacino terziario del
Piemonte' have been hitherto ignored or treated as nomina
nuda, although most meet criteria for availability and predate
introductions in Bellardi and Sacco's 1873-1904 work. Supra-
specific taxa introduced in these works are listed with biblio-
graphic references. Attention is drawn to some taxonomic prob-
lems that result from recognition of hitherto overlooked valid
introductions of names.
Key words: Bellardi; Sacco; compilation, dates; supraspecific
taxa; Tertiary fossils; Mollusca; Mastodon.
INTRODUCTION
Bellardi and Sacco s monumental work "1 molluschi dei
terreni terziarii del Piemonte e della Liguria " is one of
the most comprehensive treatments of a regional Tertiary
fauna ever published. Issued between 1873 and 1904 in
30 numbered quarto parts, it contains descriptions of
thousands of species and infraspecific taxa, numerous
genus group taxa, and several families and subfamilies.
The Bellardi and Sacco collection is now in the care of
the Museo Regionalc di Scienze Natural!, Turin (Ferrero
Mortara et al., 1982, 1984).
The primary objective of this contribution is to draw
attention to the fact that many names generally consid-
ered to date from this work were validly introduced
earlier by Sacco in a little-known companion series and
in the rare "Catalogo paleontologico del bacino terziario
del Piemonte".
Parts 1-8, 1 1 and 13a of "I mollusciii dei terreni ter-
ziarii del Piemonte e della Liguria " were published in
"Memorie della Reale Accademia delie Scienze di To-
rino ". It has been seldom appreciated that reprints of
these parts were actually published earlier than the par-
ent volumes (table 1). The remainder of the series (parts
9, 10, 12, 13b, 14-30) was published independently by
Carlo Clausen, Turin. In all assembled sets of the work
that I have seen (including the reprint by Atlante Ma-
lacologico, Rome, 1987-89), parts 1-8, 11 and 13a are
represented by reprints. Although identical in content,
the reprints differ from corresponding parts in the parent
journal in being independently paginated and in having
wrappers and title pages that were specially printed for
each part. Determination of even the pagination of each
part of the work was a laborious exercise, since there is
considerable inconsistency in pagination from part to
part in the parent journal, the reprints, the independentK-
published parts, and in Sacco's (1904:vii) compilation
(table 2). To avoid confusion with the companion series
(see below), reprints, parent journal and independently
published parts are referred to collectively as the "Me-
morie/Clausen " series. Page reference to the Memorie/
Clausen series in the following discussions and in table
3 refer only to the reprints, though both paginations are
indicated in table 2 and in the compilation of the work.
The companion series (hereafter referred to as the
"Bollettino " series) has the same title as the Memorie/
Clausen series and was issued serially in association with
parts 7-29. It was published in "Bollettino dei Musei di
Zoologia ed Anatomia Comparata della Reale Universita
di Torino ". The first two parts of the Bollettino series
(Sacco 1890c,e) are subtitled "nota preventiva" (intro-
ductory note), so the parts of this series were clearly
intended to precede corresponding parts of the Memo-
rie/Clausen series (see Sacco 1891a:l, footnote). Of the
22 issued parts of the Bollettino series, 17 were evidently
published prior to the corresponding Memorie/Clausen
parts, and 12 of these contain nomenclaturally valid in-
troductions of new taxa (see below). There are few ref-
erences to the Bollettino series in the Memorie/Clausen
series (Sacco 1891a:l footnote, 1891d, synonymies). Sub-
B. A. Marshall, 1991
Page 105
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Page 108
THE NAUTILUS, Vol. 105, No. 3
sequent references to the Bollettiiio series by other au-
thors are rare, and the few that I fiave been able to trace
{e.g., VVenz, 1938-44, Ferrero Mortara et al., 1984) refer
only to parts 7 and 8 (Sacco 1890c, e). Moreover, the series
has been overlooked in a compilation of his published
works (Anonymous, 1948).
While the Memorie/Clausen series was in progress,
Sacco (1890a) also published "Catalogo paleontologico
del bacino terziario de Piemonte". This is essentially a
tabular listing of the 5,472 species of Tertiary fossils
known from Piedmont. Although Sacco (1904:40, 74)
dated his new subgenera Simplicotaiirasia and Miocenia
from this work, the fact that other genus group names
are potentially available there has been entirely over-
looked.
DATES OF PUBLICATION
All reprints of parts 1-8, 11, and 13a of the Memorie/
Clausen series were clearly published before correspond-
ing parts in the parent journal (table 1).
Reprints of parts l-5[a-c] bear year dates on the title
pages and wrappers and are secondarily dated in the text
with a year, month and, with the e.xception of part 2, a
day. With the exception of part 1, the imprinted year
dates on the title page and in the text are accordant.
Although it is possible that the textual dates actually
represent dates of completion of the manuscripts, they
are interpreted as dates of publication in the absence of
evidence to the contrary (ICZN, 1985, Art. 21).
Part 1 (Bellardi, 1873) is almost invariably dated from
1872 {e.g., Sacco, 1889:155, 1904:vii; Wenz, 1944:1513)
in accordance with the date on the title page and wrap-
per. The text (p. 264), however, is dated June 10 1873
and there are references to a publication dated 1873 in
some synonymies {e.g., p. 130), so the 1872 date is clearly
an error.
Dates imprinted on the backs of the title pages of
reprints of parts 6, 7, 8 and 1 1 are unequivocally dates
of printing, as are dates on their parent volumes (table
1). Curiously, Sacco (1904:vii) dated part 6 from Feb-
ruary 1890, part 8 from August 1890, and part 11 from
February 1892, but these are surely errors since they
predate the printing dates. That Sacco's dating of August
1890 for part 7 is almost certainly incorrect is suggested
by the fact that the Natural History Museum, London
(BMNH) copy of the reprint was accessioned on Septem-
ber 1, 1890 yet all other parts were accessioned by that
library 2-8 months later than the imprinted dates.
The reprint of part 13a bears only the year of publi-
cation (1893), although from the statement on the title
page verso it must have been published after February
1893 and presumably before September 1893, the pub-
lication date of part 13b In the absence of conflicting
evidence, it is appropriate to accept Sacco's (1904:vii)
indication that it was published in April 1893 (received
BMNH August 14 1893).
For bibliographic and priority purposes, the day of
publication is interpreted as the last day of the stated
month (ICZN, 1985, Art. 21i). Days of publication of
reprints so interpreted (table 1) are shov\n in square
parentheses. Year dates associated with new supraspecific
taxa in the body of the text in both the Memorie/ (Clausen
and Bollettino series frequently do not correspond with
the actual year of publication and evidently represent
merely anticipated years of publication.
All parts of the Bollettino series bear imprinted dates
(day, month and year), and library accession date stamps
are accordant with their interpretation as dates of pub-
lication (table 1).
Sacco's "Catalogo paleontologico del bacino terziario
del Piemonte " was submitted to the general assembly of
the Societa Geologica Italiana on September 23 1889 for
publication in Bollettino della Societa Geologica Italiana
(BSGI — vol. 8(3), p. 545, 565). Presumably because of
its length it was decided to split the work into two parts,
which were published in BSGI volumes 8(3) and 9(2).
The work also exists reprinted in its entirety as part 3 of
Sacco's "II bacino terziario e quaternario del Piemonte"
(1889-90). Despite extensive enquiries with numerous
libraries, including the publishers and Societa Geologica
Italiana, I have been unable to determine the exact dates
of publication of the two forms of the Catalogo.
References to Sacco's article in BSGI vol. 8(3) have
been consistantly dated from 1889, but although parts
1-3 of this volume bear the date 1889 beside the volume
number, part 3 is dated 1890 at the foot of the page
(parts 1 and 2 are dated 1889). The latest date mentioned
in the text of part 3 (p. 582) is in a report on an excursion
that took place on September 26 1889, so it cannot have
been published before then. Strong evidence that part 3
was published early in 1890 is provided by the list of
new publications on the back wrapper (National Museum
of New Zealand, Wellington) of series 3, volume 1, num-
bers 3 and 4 ("Marzo e Aprile 1890") of the Bollettino
del Reale Comitato Geologico d'ltalia (BRCGI) in which
papers from BSGI volume 8(3) are dated 1890. The latest
date shown in BRCGI 1(3,4) and thus the earliest date
that it can be demonstrated to have been published (ICZN
Art. 21c) is April 30 1890 (p. [159]) (interpretation ac-
cordant with dates of publication of journals mentioned
on pp. 151, 153, 156, 157). That BSGI 8(3) was probably
published no later than early March 1890 is suggested
by the fact that the BMNH cop> was received on May
3 1890, yet other Italian works of the period (table 1)
were accessioned by this library 2 or more months after
publication (see below).
BSGI 9(2) is dated 1890 on the title page and was
evidently published mid-late 1890. L'nfortunately, the
earliest references to it that I have located all date from
early 1891 and are thus of little assistance (received
BMNH February 21).
The reprint was printed from the same plates as the
BSGI parts and is identical apart from being indepen-
dently paginated and in that page 78 was split for division
of the BSGI parts. It is dated 1889 on the title page and
1890 on the front v\ rapper (University Library, Stock-
holm), and bears the statement (p. [ii]) "Estratto dal Bol-
B. A. Marshall, 1991
Page 109
Table 3. List of molluscan supraspecific taxa introduced by Bellardi & Sacco (1873-1904) in "I molluschi dei terreni terziarii del
Piemonte e dejja l.iguria, " and by Sacco (1890) in "Catalogo paleontologicn dei baciiio terziario del Piemonte ."
Acrilloscala Sacco, 1890a:214(321) (1891b:[iv]; 1891d:86).
Adiscoacrilla Sacco, 1890a:21-l(321) (1891b:[iii]; 1891d:67).
Mectryom-lla Sacco, 1897b:100 (1897d;19).
Alvaniclla Monterosato in Sacco, 189.5b;76 (189.5d;25).
Amathinoides Sacco, 1896a:41.
Ampullonatica Sacco, 1890a:208(315) (1890e:40; lS91a:10,5),
Amussiopecten Sacco. 1897c:102 (1897e:53).
Ancillarina Bellardi, 1882:217^
Ancillina Bellardi. 1882:220,
Angulatumitrella Sacco, 1890b:36.
Anura Bellardi. 1873:201.
Archimcdiclla Sacco. 1895e:12.
Arcuaiumitrclla Sacco, 1890b:37.
Ascolatims Bellardi. 1884:41.
Asperarca Sacco. 1898d:10.
Aulachalia Sacco, ]893e:33.
Bivoniopsis Sacco, 1896a:1.5,
Brachelixclla Sacco, 1890a: 1 1.5(222) (1890b:34).
Callistotapes Sacco, 1900:52.
Cardiohicina Sacco. 1901a:89.
Centrucardita Sacco. 1899b: 14.
Ceromyclla Sacco. 1901a: 133.
Clanculella Sacco, 1896b:22,
Clinomitra Bellardi. 1888:10,
Clinurella Sacco. 1890a:117(224) (1890b:44).
Coccodentalittm Sacco. 18961:98 (1897a:110).
Conocerilhium Sacco. 1895a:72 {nom.nud.}; 1895c:22.
Contortia Sacco. 1894b:69 (1894d:48).
Costatoscala Sacco. 1891d:72,
Costoanachis Sacco. 1890b:57
Crassostrea Sacco, 1897b:99 (lS97d:15),
Ciihitoslrea Sacco, 1897b:99 (1897d:12),
Cyclodostomia Sacco. 1892a:53 {nom.nud.); 1892c:46.
Cyllenina Bellardi. 1882161
CYLLENINAE Bellardi. 1882: 159
Cymbiilostrea Sacco. 1897d:12,
Cyrsocrassiscala Sacco. 1891e:66.
Cytherocardia Sacco. 1900:5,
Denticuloglahella Sacco, 1890b:25.
Dertonia Bellardi. 1884:51.
Diptychomitra Bellardi, 1888: 10
DIPTYCHOMITRLNAE Bellardi, 1888: 10
Dtscosca/fl Sacco, 1890a:213(320) (1891b:[iii]; 189ld:.57),
Ditoma Bellardi, 1877295
Dizoniopsis Sacco, 1895a:74 (1895c:67).
Dolicholatims Bellardi, 1884:38,
Echinophoria Sacco. 1890a:88(195) (1890c:9; 1890d:39).
Ecostoanachis Sacco, 1890b:59,
Eolatirus Bellardi. 1884:15,
Eratotrivia Sacco, 1894c:62.
Euspirocrommhtm Sacco. 1890a:208(315) (Sacco, 1890e:42;
lS91b:[i]; 1891d:10).
FimbriateUa .Sacco, 1895e:36.
Flabellipecten Sacco, lS97c:102 (1897e:55).
Flabellulum Bellardi. 1873:32,
Flexopectcn Sacco. 1897c:102 (1897e:39).
Fulguroficus Sacco, 1890a:97(204) (1890e:27; 1891a:41),
Fusimorio Sacco, 18961:90 (1897a:25),
Fusoficula Sacco, 1890a:97(204) (nom. nud.); 1890e:26
(1891a:38),
Fusoterebra Sacco, 1891c:[vi] (1891e:59).
Caleodinopsis Sacco, 189.5b:77 (189.5d:28).
Caleudocassis Sacco, 1890a:85(192) (nom. nud.); 1890c:5
(1890d:18),
G.\LEODOLIIDAE Sacco. 1890e:21 (invalid— no type ge-
nus); 1891a:l,
Caleodolium Sacco, 1891a:4
Galeodosconsia Sacco, 1890a:93(200) (1890c:17; 1890d:69),
Camoplcura Bellardi, 1873:28,
Genoa Bellardi. 1873:205.
Gibbominliola Sacco. 1898a:41.
Gibbori.s.soia Cossmann in Sacco, 189.5d:34.
CMndoidca Sacco. 1896b:32
GdAndoidclla Sacco. 1896b:38
Gigantcotrochus Sacco. 1897a:4.
Gigantostrea Sacco. 1897b:99 (1897d:14).
Glabrondina Sacco, 1892c:49.
Glabropecten Sacco, 1897e:39, 73 (error for Flexopectcn).
Granosolarium Sacco, 1892b:56 (1892d:.59),
HALIIDAE Sacco. 1893d:64 (1893e:32) (=Halliidae Sisinon-
da. 1842),
Hordeulima Sacco. 1892a:52 (1892c: 17).
Jania Bellardi. 1873:147,
Jousseaiimia Sacco. 1894c:8,
Kleistopyrazus Sacco. 1895c:56
LACHESINAE Bellardi, 1877:150
Lazariella Sacco, 1899a:112 (1899b:21),
Ledina Sacco. 1898c:107 (1898d:.53).
Limatulella Sacco. 1898a: 16,
Lissochlamis Sacco. 1897c:102 (1897e:46),
Luponovula Sacco. 1894a:66 {nom. nud.); 1894c:44.
Macomopsis Sacco. 1901a: 107
Macrochlamis Sacco, 1897c: 101 (1897e:32),
Macrodostomia Sacco. 1892a:53 (nom. nud.); 1892c:43.
Macrurella Sacco. 1890a:117(224) (1890b:46),
Mantellina Sacco, 1904:148,
MargoDohUa Sacco, 1890a:94(201) (nom. nud); 1890c:20;
(1890d:82),
MATHILDIDAE Sacco, 1892c:27 (=Mathildidae Dall,
1889),
Mayeria Bellardi, 1873:156,
Megacardita Sacco. 1899a:lll (1899b:9).
Micromitra Bellardi. 1888:5,
Miocardiella Sacco, 1904:161,
Miocenia Sacco, 1890a: 100(207) (1904:40).
Mioporomya Sacco, 1901a:140,
Mitraefusm Bellardi, 1873:204.
MITROLIMNIDAE Sacco, 1904:88.
Modiohda Sacco. 1898a:41,
Monodontclla Sacco. 1896b:23,
Morionassa Sacco, 1890a:93(200) (nom. nud.); 1890c:18
(1890d:74).
Myrsopsis Sacco, 1900:57.
Myrteopsis Sacco, 1901a:96.
Ncoathleta Sacco. 1890a:152(259) (1890b:12).
Ncolatiru.s Bellardi, 1884:40.
Nodosolarium Sacco. 1892d:63,
Obliquarca Sacco. 1898c: 106 (1898d:16).
Oombobna Sacco. 1896b: 15,
Oopecten Sacco. 1897c:102 (1897e:54),
Oostrombus Sacco, 1893d:63 (1893e:13).
Ooterebralia Sacco. 1895c:51.
OrmastraUum Sacco, 1896b:15.
ORTHOMITRIXAE Bellardi. 1887a:3.
lu
THE NAUTILUS, Vol. 105, No. 3
Table 3. Continued
Orilmrclk Sacco, 1890a:l 17(224) (If'aObi.Sl).
Pantherinaria Sacco, 1894a:65 (1894c: 10,67).
Papillicardhim Monterosato in Sacco, lS99a:11.3 (1899b;44)
Parvamiissium Sacco, 1897c: 102 (1897e:48).
Parvivcnus .Sacco. 1900 4.5
Parvochlamys Sacco, 1904: 145.
Pectinatarca Sacco, 1898c: 106 (1898d:26).
Phornilellus Sacco, 1896b:35.
Pilhocerithium Sacco, 1895a:72 (1895c:28).
Plrsiolatirus Bcllardi. 1884:32.
I'LESIOMITHINAE Bellardi, 1887b:23.
Plicostoniia Montero.sato in Sacco, 1892c:46.
Poculina Bcllardi. 1873:33
Pohlgireulima Sacco, 1892c:10.
PwaJusta Sacco, 1894a:66 (1894c:33).
Pseudavena Sacco, 1896f:92 (nom. mid.), 1897a:39.
Pseudemarginula Sacco, 1897a:17.
Pseitdolalims Bellardi, 1884:46.
Pscudonina Sacco, 1896b:9.
PaeudonineHa Sacco, 1896b:51.
Pscudosthenorytis Sacco, 1891d:43.
Pscudotorinia Sacco, 1892d:66.
Pseiidoxyperas Sacco, 1901a:26.
Ptycheulimella Sacco, 1892a:53 (1892c:59).
Ptychocerithium Sacco, 1895a:72 (1895c:23).
Ptychomdania Sacco, 1895b:75 (1895d:7).
Ptychopotamides Sacco. 1895a:73 (1895c:44).
PurpurcUa Bellardi. 1882:193.
PI RPl RELLINAE Bellardi, 1882:193.
HLRPURINAE Bellardi, 1882:174 ( = Piirpurinae Lamarck,
1809).
Pyrogolampros Sacco, 1892b:54 (nam. nud.)\ 1892c:85.
Rliombomya Sacco, 190 la: 133.
Ringiculella Sacco, 1892b:55 (1892d: 16).
Ringiculocosta Sacco, 1892b:56 (1892d:33).
Ringiculospongia Sacco, 1892b:55 (1892d:15).
Ruuaidtia Bellardi, 1877:223.
Sainliopsis Sacco, 1898a:ll.
Sassia Bellardi, 1873:219.
Scalnella Sacco, 1890a: 118(225) (nom. nud.}; 1890b:54.
Scalaricardita Sacco, 1899b:23
Scaptorrhynchus Bellardi, 1873:12.
Scgucnziclla Sacco, 1904:123.
Sigaretotrcma Sacco, 1890a:207(314) (1890e:38; 1891a:97),
Sinipliiodnlinin Sacco, 1891a:13.
Simpli,ogl(d>ella Sacco, 1890b;21.
Simplirolanmsia Sacco, 1890a:124(231 ) (1904:74).
Spica Monterosato in Sacco, 1892a:53 (nom. nud.}; 1892c:
63.
Spineoterelna Sacco. I891c:[vi] (1891e:58).
Siazzania Sacco, 1890a: 138(245) (1890b:26).
Steirotuha Sacco, 189.5e:40.
Strigosella Sacco, 1896b:49.
Striolucina Sacco, 1901a:83.
Striolcrclnum Sacco, 1891c:[vi] (1891e:33).
Striolurlnmilla Sacco, 1892b:.55 (1892c:94).
Sulcoghidius Sacco, 1893d:63 (1893e:15).
Sulconuirinula Sacco, 1896f:95 (nom. nud.h !897a:80.
Sulcosulndaria Sacco, 1892a:52 (nom. nud.); 1892c:17.
Sulcoturhomlla Sacco, 1892b:55 (1892c:92).
Syndesmyella Sacco, 1901a:122.
Tflnra.sifl Bellardi, 1882:194.
TAURASINAE Sacco, 1904:74.
Tauraxinus .Sacco, 1901a:61.
TauTcmarginula Sacco, 1897a:13.
Taurofuri.s Sacco, 1895c:62.
Taurotapes Sacco, 1900:58.
Tecionatica Sacco, 1890a:205(312) (1890e:33; 1891a:81).
TEREBELLIDAE Sacco, 1893d:64. (=Terebellinae H.&.A.
Adams, 1854).
Tetrastomella Sacco, 1890a: 11 6(223) (1890b:41).
Timirclla Sacco, 1890a:l 17(224) (I890b:52).
Thiarinclla Sacco, 1890a: 118(225) (1890b:56).
Tiamccrithium Sacco, 1895a:72 (189.5c:35).
Tiarapirenella Sacco, 1895a:73 (189.5c:60).
Tomochiton Sacco, 1877a:90 (nom. nud.).
Torculoidella Sacco, 1895e:28.
TORNIDAE Sacco, 1896b:55.
Trochacerilhium Cossmann & Sacco in Sacco, !896t):60.
Trocholugurium Sacco, 1896a:27.
Tuhcrculodolium Sacco, 1891a:9.
Turritodoslomia Sacco, 1892a:53 (1892c:41).
Tylastmlium Sacco. 1896b:19.
Uromitra Bellardi, 1887b:23.
Variamussium Sacco, 1897c: 102 (1897e:49).
Ventricoloidea Sacco, 1900:31.
Vulgocerithium Sacco, 1895c:7.
lettino della Societa geologica italiana Vol. 8(3)". This
strongly suggests that the reprint was printed when it
was still assumed that the work was to be published entire
in BSGI 8(3) and thus before that part was published.
The date "1889" on the title page could indicate that it
was printed late in that year and that the wrapper was
dated 1890 when it was realised that the work would not
be circulated until 1890, or that BSGI 8(3) was to be
published in 1890. It seems more plausible, however, that
the reprint was printed in 1890, and that the 1889 date
was added to the manuscript by Sacco and/or simply
represents the nominal year for BSGI volume 8. The
earliest reference to the work (Sacco, 1890d:7, footnote —
"Boll. See. geol. it., 1889-90") appears in an introduction
dated February 5 1890. Since the work cannot be proven
to have been published in 1889, in the absence of evi-
dence to the contrarv the date of publication is inter-
preted as February ,5' 1890 (ICZN, 1985, .Art. 21c). This
conclusion is accordant w ith C. D. Sherborn's, who noted
"issued before the Bollettino" on the title page of the
BMNH copy of the reprint.
STATUS OF TAXA
Apart from supraspecific names (see below), parts 7 and
8 of the Bollettino series (Sacco 1890c,e) contain brief
Latin diagnoses of hundreds of species group taxa ac-
companied b\- specimen dimensions and type localities.
B. A. Marshall, 1991
Page 1 1 1
Although these descriptions are brief and unaccompa-
nied by illustrations, many meet criteria for availability
(ICZN, 1985: Art. 11, 12), so that approximately half the
total number of species group taxa more fulK described
and illustrated in the corresponding parts of the Me-
morie/Clausen series actually date from the prior Bol-
lettino parts (taxa in part 8 so dated by Ferrero Mortara
et al., 1984). The remainder of the species group names
first introduced in parts 7 and 8 of the Bollettino series
and all of those in subsequent parts are nomiija niida.
since they are mentioned by name onl\ .
Most supraspecific names first introduced in the Bol-
lettino series meet criteria for availability since they are
accompanied by one or more valid species names that
were introduced by earlier authors. Some of the generic
and subgeneric names, however, are nomina nitda. since
they are accompanied onh' by specific names (i.e., Sac-
co's) that are themselves nomina nuda. Two subfamily
names (Orthomitrinae and Plesiomitrinae) introduced by
Bellardi ( 1887a, b) are unavailable since they are not based
on a type genus, while Galeodoliidae is unavailable at
its first introduction (Sacco, 1890e:21) because it is un-
accompanied by a type genus (Galeodolium Sacco, 1891a:
4). Genus group names first introduced by Sacco (1890c,e,
1891b) in parts 7-9 of the Bollettino series lack type
species designations, while type species are designated
unequivocally in all of the following parts. Type species
for man\- genus group names introduced by prior authors
are indicated in parts 17-29 of the Bollettino series and
are thus potentially available as first subsequent desig-
nations.
All moUuscan species group names introduced by Sac-
co in "Catalogo paleontologico del bacino terziario del
Piemonte" are nomina nuda. Of the 26 genus group
names first introduced there, however, 21 meet criteria
for availability because they are accompanied by valid
specific names that were introduced in prior publications
(ICZN, 1985: Art. 12b(5)). The only species group name
that is validly introduced in this work (reprint p. 192) is
Mastodon arvernensis var. cantamessae Sacco, 1890
(Mammalia).
Attention is drawn to the fact that dating of genus
group taxa from the Bollettino series and the "Catalogo"
affects some currently accepted type species designa-
tions. For example, in cases where Sacco did not desig-
nate a type species in the Memorie/Clausen series, au-
thors have selected type species by subsequent
designation, but some of these designations are invalid
because the species were not among those included when
the genus was first introduced. These problems should
preferably be treated individually by specialists, and to
avoid influencing future decisions they are not consid-
ered here.
All molluscan supraspecific names introduced by Bel-
lardi (1873-1888) and Sacco (1890-1904) that I have
been able to trace are listed (table 3). For convenience,
page references are provided for all names in the Me-
morie/Clausen series (reprints) and in "Catalogo pa-
leontologico del bacino terziario del Piemonte , but onl\
to prior introductions (valid and invalid) in the Bollettino
series. Page references to the "Catalogo" include both
the reprint and (in parentheses) Bollettino della Societa
Geologica Italiana 9(2). Unless indicated as nomen nu-
dum or otherwise invalid, the first citation is interpreted
as the earliest nomenclaturally valid introduction of the
name.
Spelling of Lissochlamis, M.acrochl.\mis and
v.\riamvssium
In part 24 of the Memorie/Clausen series, Sacco (1897e)
altered the original spellings (Sacco, 1897c) of Lissochla-
mis, Macrochlamis, and Variamussum to Lissochlamys,
Macrochlamys and Variamussium. Although "chlamis"
is an incorrect transliteration of the Greek "chlamys" (a
garment), incorrect transliterations are not to be consid-
ered inadvertent errors (ICZN, 1985: Art. 32c), so Lis-
sochlamys and Macrochlamys are unjustified emenda-
tions (ICZN, 1985: Art. 33). That Sacco's (1897c) original
spellings of Lissochlamis and Macrochlamis were per-
haps intentional is suggested by the consistent usage (twice
each), as subgenera of Chlamys (sic. — three times). Ac-
cordingly, I follow Hertlein (1969) and Yokes (1980) and
interpret Lissochlamis and Macrochlamis as correct
spellings, despite the fact that Lissochlamys and Macro-
chlamys have been widely used.
The original spelling Variamussum has been entirely
overlooked, and Sacco's subsequent spelling has been
universally accepted. Although Variamussum is consis-
tently so-spelt (twice) in the original reference, it is ob-
viously a lapsus, so from the standpoint of nomenclatural
stability it is appropriate in this instance to accept Varia-
mussium as a justified emendation.
COMPILATION OF "I MOLLUSCHI DEI
TERBENI TERZIARII DEL PIEMONTE E
DELLA LIGURIA", AND "CATALOGO
PALEONTOLOGICO DEL BACINO
TERZIARIO DEL PIEMONTE"
It should be noted that the sections entitled "Catalogo
generale dei molluschi dei terreni terziarii del Piemonte
e della Liguria" occurring in each part of the Memorie/
Clausen series {i.e.. in both parent journal and reprints)
were reprinted independently with independent pagi-
nations and title pages. Identical to the originals, these
reprints were prepared for separate binding and are not
included here.
Bellardi, L. 1873. 1 molluschi dei terreni terziarii del
Piemonte e della Liguria. Parte 1: Cephalopoda,
Pteropoda, Heteropoda, Gasteropoda (Muricidae e
Tritonidae). Memorie della Reale Accademia delle
Scienze di Torino (2) 27, 1873:33-294 (reprint 264
p.), 15 plates (June 10 1873).
Bellardi, L. 1877. I molluschi dei terreni terziarii del
Piemonte e della Liguria. Parte 2: Gasteropoda
(Pleurotomidae). Memorie della Reale Accademia
Page 112
THE NAUTILUS. Vol. 105, No. 3
delle Scienze di Torino (2) 29, 1878:1-364 (reprint
364 p.), 9 plates (June 30 1877).
Bellardi, L. 1882. I nioUiischi dei terreiii terziarii del
Piemonte e della liguria. Parte 3: Gasteropoda (Buc-
cinidae, Cyclopsidae, Purpuridae, Corailiophilidae,
Olividae). Memorie della Reale Accademia delle
Scienze di Torino (2) 34, 1883:219-469 (reprint 253
p.), 12 plates (December 10 1882).
Bellardi, L. 1884. I molluschi dei terreni terziarii del
Piemonte e della Liguria. Parte 4: Fasciolariidae e
Turbinellidae. Memorie della Reale Accademia delle
Scienze di Torino (2) 37, 1886:3-62 (reprint 62 p.),
2 plates (December 1 1884).
Bellardi, L. 1887a. I molluschi dei terreni terziarii del
Piemonte e della Liguria. Parte 5[a]: Mitridae. Me-
morie della Reale Accademia delle Scienze di To-
rino (2) 38, 1888:79-166 (reprint 85 p.), 2 plates
(January 15 1887).
Bellardi, L. 1887b. 1 molluschi dei terreni terziarii del
Piemonte e della Liguria. Parte 5[b]: Mitridae (con-
tinuazione). Memorie della Reale Accademia delle
Scienze di Torino (2) 38, 1888:257-327 (reprint 72
p.), 2 plates (June 1 1887).
Bellardi, L. 1888. I molluschi dei terreni terziarii del
Piemonte e della Liguria. Parte 5[c]: Mitridae (fine).
Memorie della Reale Accademia delle Scienze di
Torino (2) 39, 1889:145-194 (reprint 52 p.), 2 plates
(September 20 1888).
Sacco, F. 1890a. Catalogo paleontologico del bacino
terziario del Piemonte. Reale Accademia dei Lincei,
Roma, 233 p. (reprint). Subsequently published in
Bollettino della Societa Geologica Italiana 8(3):281-
356, 9(2):185-340 (February 5 1890).
Sacco, F. 1890b. I molluschi dei terreni terziarii del
Piemonte e della Liguria. Parte 6: (Volutidae, Mar-
ginellidae, Columbellidae). Memorie della Reale
Accademia delle Scienze di Torino (2) 40, 1890:
295-368 (reprint 76 p.), 2 plates (April 6 1890).
Sacco, F. 1890c. I molluschi dei terreni terziarii del
Piemonte e della Liguria. Parte 7: (Harpidae e Cas-
sididae). Bollettino dei Musei di Zoologia ed Ana-
tomia comparata della Reale Universita di Torino
5(82): 1-20 (June 13 1890).
Sacco, F. 1890d. I molluschi dei terreni terziarii del
Piemonte e della Liguria. Parte 7: (Harpidae e Cas-
sididae). Memorie della Reale Accademia delle
Scienze di Torino (2) 40, 1890:469-560 (reprint 96
p.), 2 plates (June 18 1890).
Sacco, F. 1890e. I molluschi dei terreni terziarii del
Piemonte e della Liguria. Parte 8: Galeodoliidae,
Doliidae, Ficulidaee Naticidae. Bollettino dei Musei
di Zoologia ed Anatomia comparata della Reale
Universita di Torino 5(86):21-43 (August 12 1890).
Sacco, F. 1891a. I molluschi dei terreni terziarii del
Piemonte e della Liguria. Parte 8: Galeodoliidae,
Doliidae, Ficulidae e Naticidae. Memorie della Reale
Accademia delle Scienze di Torino (2) 41, 1891:225-
338 (reprint 114 p.), 2 plates (March 25 1891).
Sacco, F. 1891b. I molluschi dei terreni terziarii del
Piemonte e della Liguria. Parte 9: Naticidae (fine),
Scalariidae ed Aclidae. Bollettino dei Musei di Zoo-
gia ed Anatomia comparata della Reale Universita
di Torino 6(103):[i-iv] (May 29 1891).
Sacco, F. 1891c. [I molluschi dei terreni terziarii del
Piemonte e della Liguria] Parte 10: Cassididae (ag-
giunte), Terebridae e Pusionellidae. Bollettino dei
Musei di Zoologia ed Anatomia comparata della
Reale Universita di Torino 6 (103):[v-vi] (published
consecutively with Part 9) (May 29 1891).
Sacco, F. 1891d. I molluschi dei terreni terziarii del
Piemonte e della Liguria. Parte 9: (Naticidae (fine),
Scalariidae ed Aclidae). Clausen, Torino, 103 p., 2
plates (May 31 1891).
Sacco, F. 1891e. I molluschi dei terreni terziarii del
Piemonte e della Liguria. Parte 10: (Cassididae (ag-
giunte), Terebridae e Pusionellidae). Clausen, To-
rino, 68 p., 2 plates (September 30 1892).
Sacco, F. 1892a. I molluschi dei terreni terziarii del
Piemonte e della Liguria. Parte 11: Eulimidae e
Pyramidellidae (pars). Bollettino dei Musei di Zoolo-
gia ed Anatomia comparata della Reale Universita
di Torino 7(121):51-53 (April 30 1892).
Sacco, F. 1892b. [I molluschi dei terreni terziarii del
Piemonte e della Liguria]. Parte 12: (Pyramidellidae
(fine), Ringiculidae, Solariidae e Scalariidae (agg.)).
Bollettino dei Musei di Zoologia ed Anatomia com-
parata della Reale Universita di Torino 7(121 );54-
57 (Published consecutively with Part 11) (April 30
1892).
Sacco, F. 1892c. I molluschi dei terreni terziarii del
Piemonte e della Liguria. Parte 11: Eulimidae e
Pyramidellidae (parte). Memorie della Reale Ac-
cademia delle Scienze di Torino (2) 42, 1892:585-
682 (reprint 100 p.), 2 plates (June 30 1892).
Sacco, F. 1892d. I molluschi dei terreni terziarii del
Piemonte e della Liguria. Parte 12: (Pyramidellidae
(fine), Ringiculidae, Solariidae e Scalariidae (ag-
giunte)). Clausen, Torino, 88 p., 2 plates (June 30
1892).
Sacco, F. 1893a. I molluschi dei terreni terziarii del
Piemonte e della Liguria. Parte 13[a]: (Conidae)
(Fasciolo Primo). Memorie della Reale Accademia
delle Scienze di Torino (2) 44, 1894:1-54 (reprint
54 p.), 2 plates (April 29 1893).
Sacco, F. 1893b. I molluschi dei terreni terziarii del
Piemonte e della Liguria. Parte 13: (Conidae e Co-
norbidae). Bollettino dei Musei di Zoologia ed An-
atomia comparata della Reale Universita di Torino
8(153): 1-4 [NB— this part corresponds to 1893a and
1893c combined] (May 31 1893).
Sacco, F. 1893c. I molluschi dei terreni terziarii del
Piemonte e della Liguria. Parte 13[b]: (Conidae e
Conorbidae). (Fascicolo secundo). Clausen, Torino:
55-143, 9 plates (September 30 1893).
Sacco, F. 1893d. 1 molluschi dei terreni terziarii del
Piemonte e della Liguria. Parte 14: (Strombidae,
Terebellidae, Chenopidae, Haliidae, Cypreidae).
Bollettino dei Musei di Zoologia ed Anatomia com-
B. A. Marshall, 1991
Page 113
parata della Reale Universita di Torino 8(165):63-
64 (December 11 1893).
Sacco, F. 1893e. I molluschi del terreni terziarii del
Piemonte e della Liguria. Parte 14: (Strombidae,
Terebellidae, Chenopidae ed Haliidae). Clausen,
Torino, 40 p., 2 plates (December 30 1893).
Sacco, F. 1894a. I molluschi dei terreni terziarii del
Piemonte e della Liguria. Parte 15: Fam. C\praei-
dae Gray 1824. Bollettino dei Musei di Zoologia ed
Anatomia comparata della Reale Universita di To-
rino 9(171):65-67 (April 27 1894).
Sacco, F. 1894b. [I molluschi dei terreni terziarii del
Piemonte e della Liguria] Parte 16: Fam. Cancel-
lariidae H. e A. Adams 1853. Bollettino dei Musei
di Zoologia ed Anatomia comparata della Reale
Universita di Torino 9(171 ):68-70 (published con-
secutively with Part 15) (April 27 1894).
Sacco, F. 1894c. I molluschi dei terreni terziarii del
Piemonte e della Liguria. Parte 15: (Cypraeidae, ed
Amphiperasidae). Clausen, Torino, 74 p , 3 plates
(April 29 1894).
Sacco, F. 1894d. I molluschi dei terreni terziarii del
Piemonte e della Liguria. Parte 16: (Cancellariidae).
Clausen, Torino, 82 p., 3 plates (August 31 1894).
Sacco, F. 1895a. I molluschi dei terreni terziarii del
Piemonte e della Liguria. Parte 17: (Cerithiidae,
Triforidae, Cerithiopsidae e Diastomidae). Bollet-
tino dei Musei di Zoologia ed Anatomia comparata
della Reale Universita di Torino 10(197):71-74
(February 25 1895).
Sacco, F. 1895b. I molluschi dei terreni terziarii del
Piemonte e della Liguria. Parte 18: (Melaniidae,
Littorinidae, Fossaridae, Rissoidae, Hydrobiidae,
Paludinidae e Valvatidae). Bollettino dei Musei di
Zoologia ed Anatomia comparata della Reale L^ni-
versita di Torino 10(206):75-78 (May 30 1895).
Sacco, F. 1895c. I molluschi dei terreni terziarii del
Piemonte e della Liguria. Parte 17: (Cerithiidae,
Triforidae, Cerithiopsidae e Diastomidae). Clausen,
Torino, 86 p., 3 plates (June 30 1895).
Sacco, F. 1895d. I molluschi dei terreni terziarii del
Piemonte e della Liguria. Parte 18; (Melaniidae,
Littorinidae, Fossaridae, Rissoidae, Hydrobiidae,
Paludinidae e Valvatidae). Clausen, Torino, 52 p.,
1 plate (September 30 1895).
Sacco, F. 1895e. I molluschi dei terreni terziarii del
Piemonte e della Liguria. Parte 19: (Turritellidae e
Mathildidae). Clausen, Torino, 46 p., 3 plates (De-
cember 30 1895).
Sacco, F. 1896a. I molluschi dei terreni terziarii del
Piemonte e della Liguria. Parte 20: (Caecidae, Ver-
metidae, Siliquariidae, Phoridae, Calyptraeidae,
Capulidae, Hipponycidae, Neritidae, Neritopsidae).
Clausen, Torino, 65 p., 5 plates (May 31 1896).
Sacco, F. 1896b. I molluschi dei terreni terziarii del
Piemonte e della Liguria. Parte 21: (Naricidae,
Modulidae, Phasianellidae, Turbinidae, Delphinu-
lidae, Cyclostrematidae, Tornidae). Clausen, Tori-
no, 64 p., 4 plates [NB — p. 64 incorrectly numbered
•'65"] (September 30 1896).
Sacco, F. 1896c. I molluschi dei terreni terziarii del
Piemonte e della Liguria. Parte 19; (Turritellidae e
Mathildidae). Bollettino dei Musei di Zoologia ed
Anatomia comparata della Reale Universita di To-
rino 11(267):79-81 (December 14 1896).
Sacco, F. 1896d. I molluschi dei terreni terziarii del
Piemonte e della Liguria. Parte 20; (Caecidae, Ver-
metidae, Phoridae, Calyptraeidae, Capulidae, Hip-
ponycidae e Neritidae). Bollettino dei Musei di Zoo-
logia ed Anatomia comparata della Reale Universita
di Torino ll(267):82-84 (published consecutively
with Parts 19, 21, 22) (December 14 1896).
Sacco, F. 1896e. [I molluschi dei terreni terziarii del
Piemonte e della Liguria] Parte 21: (Naricidae,
Modulidae, Phasianellidae, Turbinidae, Delphinu-
lidae, Cyclostrematidae e Tornidae). Bollettino dei
Musei di Zoologia ed Anatomia comparata della
Reale Universita di Torino ll(267):85-88 (pub-
lished consecutiveh with Parts 19, 20, 22) (Decem-
ber 14 1896).
Sacco, F. 1896f. [1 molluschi dei terreni terziarii del
Piemonte e della Liguria] Parte 22: [untitled]. Bol-
lettino dei Musei di Zoologia ed Anatomia compara-
ta della Reale Universita di Torino ll(267):89-98
(published consecutively with Parts 19-21) (Decem-
ber 14 1896).
Sacco, F. 1897a. I molluschi dei terreni terziarii del
Piemonte e della Liguria. Parte 22: Gasteropoda
(fine) (Pleurotomariidae, Scissurellidae, Haliotidae,
Fissurellidae, Tecturidae, Patellidae, Oocorythidae,
Cyclophoridae, Cyclostomidae, Aciculidae, Trun-
catellidae; Acteonidae, Tornatinidae, Scaphandri-
dae, Bullidae, Cylicnidae, Philinidae, Umbrelli-
dae). — Pulmonata (Testacellidae, Limacidae,
Vitrinidae, Helicidae, Pupidae, Stenogyridae, Suc-
cineidae; Auriculidae; Limnaeidae, Physidae; Si-
phonariidae). Amphineura (Chitonidae). — Scapho-
poda (Dentaliidae). Clausen, Torino, 148 p., 10 plates
(March 31 1897).
Sacco, F. 1897b. I molluschi dei terreni terziarii del
Piemonte e della Liguria. Parte 23: (Ostreidae, Ano-
miidae e Dimyidae). Bollettino dei Musei di Zoolo-
gia ed Anatomia comparata della Reale Universita
di Torino 12(298);99-100 (June 11 1897).
Sacco, F. 1897c. [I molluschi dei terreni terziarii del
Piemonte e della Liguria] Parte 24; [untitled]. Bol-
lettino dei Musei di Zoologia ed .\natomia compara-
ta della Reale Universita di Torino 12(298); 101-102
(published consecutivelv with Part 23) (June 11
1897).
Sacco, F. 1897d. I molluschi dei terreni terziarii del
Piemonte e della Liguria. Parte 23: Pelecypoda (Os-
treidae, Anomiidae e Dimyidae). Clausen, Torino,
66 p., 11 plates (June 30 1897).
Sacco, F. 1897e. 1 molluschi dei terreni terziarii del
Piemonte e della Liguria. Parte 24; (Pectinidae).
Page 114
THE NAUTILUS, Vol. 105. No. 3
Clausen, Torino, 116 p., 21 plates (December 30
1897).
Sacco, F. 1898a. 1 molluschi del terreni terziarii del
Piemonte e della Liguria. Parte 25: (Spondylidae,
Radulidae, .\viciilidae, Vulsellidae, Pernidae, Pin-
nidae, Mytilidae, Dreissensiidae). Clausen, Torino,
76 p., 12 plates (August 31 1898).
Sacco, F. 1898b. I molluschi dei terreni terziarii del
Piemonte e della Liguria. Parte 25: (Spondylidae,
Radulidae, Aviculidae, Vulsellidae, Pernidae, Pin-
nidae, Mytilidae e Dreissensiidae). Boilettino dei
Musei di Zoologia ed Anatomia comparata della
Reale Universita di Torino 13(332): 103-105 (De-
cember 14 1898).
Sacco, F. 1898c. [I molluschi dei terreni terziarii del
Piemonte e della Liguria] Parte 26: (Arcidae, Pec-
tunculidae, Limopsidae, Nuculidae, Ledidae e Mal-
letidae). Boilettino dei Musei di Zoologia ed Ana-
tomia comparata della Reale Universita di Torino
13(332):106-108 (published consecutively with Part
25) (December 14 1898).
Sacco, F. 1898d. 1 molluschi dei terreni terziarii del
Piemonte e della Liguria. Parte 26: (Arcidae, Pec-
tunculidae, Limopsidae, Nuculidae, Ledidae, Mal-
letidae). Clausen, Torino, 92 p., 12 plates (December
30 1898).
Sacco, F. 1899a. 1 molluschi dei terreni terziarii del
Piemonte e della Liguria. Parte 27: (Unionidae, Car-
ditidae, Astartidae, Crassatellidae, Lasaeidae, Gale-
ommidae, Cardiidae, Limnocardiidae e Chamidae):
Boilettino dei Musei di Zoologia ed Anatomia com-
parata della Reale Universita di Torino 14(349):111-
114 (May 23 1899).
Sacco, F. 1899b. I molluschi dei terreni terziarii del
Piemonte e della Liguria. Parte 27: (Unionidae, Car-
ditidae, Astartidae, Crassatellidae, Lasaeidae, Gale-
ommidae, Cardiidae, Limnocardiidae, Chamidae).
Clausen, Torino, 102 p., 14 plates (September 30
1899).
Sacco, F. 1900. I molluschi dei terreni terziarii del
Piemonte e della Liguria. Parte 28: (Isocardiidae,
Cyprinidae, Veneridae, Petricolidiae, Cyrenidae,
Sphaeridae). Clausen, Torino, 98 p., 14 plates (April
29 1900).
Sacco, F. 1901a. I molluschi dei terreni terziarii del
Piemonte e della Liguria. Parte 29: (Donacidae,
Psammobiidae, Solenidae, Mesodesmidae, Mactri-
dae, Cardiidae, Myidae, Corbulidae, Glycymeridae,
Gastrochaenidae, Pholadidae, Teredidae, Crypto-
dontidae, Ungulidae, Lucinidae, Tellinidae, Scrobi-
culariidae, Cuspidariidae, Solenomyidae, Pandori-
dae, Verticordiidae, Lyonsiidae, Ceromyidae,
Arcomyidae, Anatinidae, Poromyidae, Clavagelli-
dae). Clausen. Torino, 216 p., 29 plates (June 30
1901).
Sacco, F. 1901b. I molluschi dei terreni terziarii del
Piemonte e della Liguria. Parte 28: [untitled]. Boi-
lettino dei Musei di Zoologia ed Anatomia compara-
ta della Reale Universita di Torino 16(409):115-118
(November 25 1901).
Sacco, F. 1901c. I molluschi dei terreni terziarii del
Piemonte e della Liguria. Parte 29: [untitled]. Boi-
lettino dei Musei di Zoologia ed Anatomia compara-
ta della Reale Universita di Torino 16(409):1 18-125
(November 25 1901).
Sacco, F. 1904. 1 molluschi dei terreni terziarii del
Piemonte e della Liguria. Parte 30: Aggiunte e cor-
rezioni. Considerazioni generali. Indice generale dell'
opera. Clausen, Torino, 203 -I- xxxvi p., 31 plates
(August 31 1904),
ACKNOWLEDGEMENTS
First and foremost I wish to express my extreme gratitude
to Mr R. J. Cleevely (Department of Palaeontology, The
Natural History Museum, London), who patiently at-
tended to many requests for library searches and infor-
mation, and without whose insightful assistance and ap-
preciation of esoteric bibliographic problems this work
would not have been possible. A great deal of additional
information was provided by Dr R. Giannuzzi-Savelli
(Palermo), Dr M. G. Harasewych and Dr A. Kabat (Na-
tional Museum of Natural History, Washington DC),
and Dr A. Waren (Swedish Museum of Natural History,
Stockholm). Of the numerous other individuals who re-
plied to my enquiries, the following provided substantial
information and assistance: Dr P. Bouchet (Museum Na-
tional d'Histoire Naturelle, Paris), Dr A. L. Cocuzzi (Ac-
cademia Nazionale dei Lincei, Roma), Dr E. Menesini
(Dipartimento di Scienze della Terra, Pisa), Dr C. Moran-
dini (Museo Friulano di Storia Naturale, Udine), Dr S.
Palazzi (Modena), Dr G. Pavia (Dipartimento di Scienze
della Terra, Torino), Dr M. Taviani (Istituto di Geologia
Marina, Bologna), and Dr A. Zuccari (Societa Geologica
Italiana, Roma).
For constructive comments on the manuscript I am
grateful to Dr A. G. Beu, Dr P. Bouchet, Mr R. J. Cleeve-
ly, Dr R. Gianuzzi-Savelli, Dr M. G. Harasewych. Dr R.
S. Houbrick, Dr A. Kabat, Dr G. Pavia, Mr R. E. Petit,
Dr W. F. Ponder and Dr A. Waren. Special thanks to
Kathleen Ryan for cheerful expertise with word pro-
cessing, and Manuela Angelo for translations.
LITERATURE CITED
Anonymous. 1948. Elencho delle publicazioni di Federico
Sacco, dottore in scienze, gia professore di geologia al
politecnico e di paleontologia all Unixersita di Torino.
Checchini, Torino 29 p.
Ferrero Mortara, E., L. Montefameglio, G. Pavia, and R Tam-
pieri. 1982. Catalogo dei tipi e degli esamplari figurati
della collezione Bellardi e Sacco. Part 1. Musei Regionale
di Scienze Naturali, Torino. Cataloghi 6: 1-327, 58 pis.
Ferrero Mortara, E., L. Montefameglio, M. Novelli, G. Opesso,
G. Pavia, and R. Tampieri 1984, Catalogo dei tipi e
degli esamplari figurati della collezione Bellardi e Sacco.
B. A. Marshall, 1991
Page 115
Part 2. Musei Regionale di Scienze Natural!, Torino, Ca-
taloghi 7:1-484, 56 pis.
Hertlein, L. G. 1969. Pectinidae. 7n. Moore, R. C. (ed.). Trea-
tise on invertebrate paleontology, N(l) Mollusca, 6 Bival-
via. Geological Society of America, Boulder, Colorado and
University of Kansas, Lawrence, p. 348-373,
ICZN. 1985. International code of zoological nomenclature,
3rd ed. International Trust for Zoological Nomenclature
and British Museum (Natural History), London, 338 p.
Voices, H E. 1980. Genera of the Bivalvia: a systematic and
bibiographic catalogue (revised and updated). Paleonto-
logical Research Institution, Ithaca, New York. 307 p.
Wenz, W. 1938-44. Gastropoda. 1: Allgemeiner Teil und
Prosobranchia. In: Schindewolf, O. H. (ed.). Handbuch
der Palaozoologie 6, Borntraeger, Berlin, xii + 1639 +
10 p.
THE NAUTILUS 105(3):116-117, 1991
Page 116
A Note on Supposed Homonyms of
Octopus australis Hoyle, 1885, with Comments on
Octopus caniphelli Smith, 1902 (Cephalopoda: Octopodinae)
Ronald B. Toll
Doparlment of Biology
The University of tlie South
Sewanee, TN 37375, USA
ABSTRACT
"Octopus australis (Massy, 1916)" and "O. australis (Benham,
1942)'", both from New Zealand, are determined not to be
homonyms of O. australis Hoyle, 1885, from Australia, as pre-
viously suggested. Examination of new material attributable to
Octopus camphclli Smith, 1902, from New Zealand, supports
the ta.xonomic distinction between O. camphclli and O. aus-
tralis. Massy 's, Benham's and Dell's material is probabK refer-
able to O. campelli.
Key words: Octopus australis; Octopus cainpbclli: Octopod-
Tait (1982) redescribed and reviewed the taxonomic po-
sition of Octopus australis Hoyle, 1885, originally de-
scribed from Port Jackson, New South Wales, Australia.
In his review of related taxa "described from New Zea-
land, Tait (1982:20) indicated that two nominal taxa,
"Polypus australis Massy [not Massey], 1916" and "Rob-
sonella australis Benham, 1942, " are junior homonyms
of Octopus australis Hoyle, following the placement of
Polypus and Robsonella into the synonymy of Octopus
s. s. by Robson (1929) and Pickford (1955), respectively.
Tait (1982:20) concluded that these two species require
renaming. Massy s (1916) account, however, clearly in-
dicated that she did not describe Polypus australis as a
new species, but actually attributed her specimens to
Hoyle s australis, and indeed cited Hoyle as the author.
Therefore, Polypus australis is simply a new combina-
tion proposed by Massy, with Hoyle retained as the au-
thor of the species [see ICZN, 1985: Art. 50(c)(ii)]. The
same is true for Robsonella australis, a new combination,
not a new species, proposed by Benham (1942) based on
his examination of new material.
Tait (1982:19) suggested, and I agree, that based on
characters of the liectocotylus, it is highly unlikely that
Octopus australis from Australia and (). campbelli Smith,
1902, from C;am|)bell Island, New Zealand are conspe-
cific, as proposed by Robson (1929:145). Tait further
stated (pg. 20) that ""all the New Zealand species pre-
viously considered to be synonyms of O. australis Hoyle
appear to be separate and distinct". The available data,
however, do not support this assertion. The type of O.
campbelli and the specimens attributed to Ho> le's aus-
tralis by Massy (1916), Benham (1942), and Dell (1952)
are all characterized by W-shaped funnel organs. Robson
(1929:190) gave the value 8.5 as the ligula length index
(LLI) of the type of O. campbelli. Benham's specimens,
remeasured by Tait, have a LLI of 6 to 10. Massy s
specimen has a LLI of 10.9; however, the proximal start-
ing point along the hectocotylus used to obtain this mea-
surement is unclear. Dell's specimens have a LLI of 5-
7. These overlapping data sets certainly cannot be used
to distinguish among taxa. In comparison, Tait (1982)
reported the LLI ior O. australis as 12-18. Massy 's (1916)
figure of the radula showed a symmetrical, pentacuspid
rachidian. Dell (1952) reported that the rachidian tooth
is asymmetrical (however, he gave the formula as ""A4",
probably an error due to the confusing situation created
by Robson s original designation of the letter A for sym-
metrical rachidian teeth, see Robson, 1925). This varia-
tion of the radula is within the range observed from new-
material of O. campbelli examined here (see below).
Therefore, based on these accounts there appears to be
no clear taxonomic separation among New Zealand spec-
imens.
For comparison to published accounts, I examined six
specimens of Octopus campbelli (4 males, ML 22-30
mm, all with spermatophores; 2 mature females, ML 25-
27 mm) from Portobello, New Zealand, deposited in the
collections of the Invertebrate Museum, Rosenstiel School
of Marine and Atmospheric Sciences, University of Mi-
ami, Florida (UMML 31.2447). The range of LLI is 6.5-
11.1 (mean = 8.48). Two inconsistencies exist between
the original description of the type of O. campbelli and
the specimens I have examined. The Portobello speci-
mens have gill lamellae counts of 7-8 (including terminal
lamella). Smith (1902) gave no data on the gills from the
holotype of (). campbelli. Robson (1929:190) stated that
the gill count of the holotype is ""probably ten", based
on his reexamination. Robson's 1929 monograph contains
certain factual inconsistencies (Voss, 1973). Indeed, Tait
R. B. Toll, 1991
Page 117
(1982:20) found that Robson, despite his assertion that
he had examined the radula from one of the syntypes of
O. australis, could not have done so because the buccal
mass had not been removed from either specimen at the
time Tait acquired them for examination. Therefore,
Robson s gill lamellae data, given with some degree of
uncertainty, should be discounted.
The second discrepancy concerns the greatly enlarged
and elevated suckers (seventh pair only) on the lateral
arm pairs of the type of O. camphcUi (SnI ~ 8.5; Sel ~
14.3). Three Portobello male specimens have moderately
enlarged suckers (SnI 8.2-10.4; Sel 10.7-12.2) which are
not distinctly elevated. In these specimens about three
pairs of suckers, beginning with the fifth pair, are en-
larged on the lateral arm pairs. Specially enlarged suckers
are not mentioned specifically in the accounts of Massy,
Benham, or Dell. While this disparity is curious, it could
be the result of differences in reproductive maturity. In
light of the other numerous similarities in morpholog\ ,
dermal texture, and absolute size, I do not consider this
difference to be substantiative.
The radulae of three of the Portobello specimens were
examined and showed considerable variation. The ra-
chidian tooth is strongK as\ mmetrical in one specimen,
weakK as\ mmetrical in the second, and varies from s\ m-
metrical in one region to asymmetrical along another
region in the third example Intraspecific (including in-
tra-radular) variation is knov\n to occur in Octopoda
(Robson, 1925, 1929; KAam, 1933, 1941; Voss, 1973; Toll,
1981, unpublished); therefore, differences in the sym-
metry of the radula can be unreliable in distinguishing
among related species.
Based on examination of all pertinent accounts, I find
that no homonyms of Octopus australis Hoyle, 1885
exist; therefore, no renaming needs to be take place.
Octopus australis Hoyle and Octopus carnpbclli Smith,
1902 appear to be distinct allopatric taxa, most easiK
differentiated on the basis of hectocotylus and penial
apparatus morphology, dermal texture and absolute size.
Furthermore, the specimens described by Mass\. Ben-
ham and Dell from New Zealand are consistent with the
characters of O. campbelli and I consider them to be
referable to it.
ACKNOWLEDGEMENTS
Nancy A. Voss, University of Miami, kindly loaned the
specimens of Octopus campbelli. Clyde F. E. Roper and
Michael J. Sweene\- made constructive comments on an
earlier version of the manuscript. This contribution was
supported, in part, by a grant from the National Science
Foundation (BSR 8508439) and general support for re-
search from The I'niversity of the South.
LITERATURE CITED
.Adam, VV, 19.33. Notes sur le.s Clephalopodes II. .Aiiomalie
de la radule chez Octopus vulgaris Lamarck et observa-
tions sur la seriatioii et I'asymetrie. Musee royal d'Histoire
naturelie de Beigit|iie 9(3), 8 p.
.\dam, W 1941. NotessurlesCephalopodes. XV. Sur la valeur
diagnostique de la radule chez les Cephalopodes Octo-
podes Musee roval d'Histoire naturelie de Belgique 17(38),
19 p.
Benham, W. B. 1942. The octopodous Molliisca of New Zea-
land. 1. The midget octopus of coastal waters. Transactions
of the Royal Society of New Zealand 72:226-236.
Dell, R, K. 19.52. The recent Cephalopoda of New Zealand-
Dominican Museum Bulletin No. 16, 1.57 p.
ICZN. 1985. International code of zoological nomenclature,
3rd ed L'niversity of California Press, 338 p.
Massy, .\. L. 1916, Cephalopoda. British Antarctic ("Terra
Nova") Expedition, 1910. Zoology 2(7):141-175.
Pickford, G. 1955. A revision of the Octopodinae in the col-
lections of the British Museum. Bulletin of the British
Museum of Natural Histor\ 3(3):151-167.
Robson, G. C. 1925. On seriation and as\ mmetr\ in the ceph-
alopod radula. Journal of the Linnean Societ\ of London,
Zoology 36:99-108,
Robson, G. C. 1929. A monograph of the Recent Cephalop-
oda. Part 1, Octopodinae, British Museum (Natural His-
tory), London, 236 p,, 7 pis.
Smith, E. A. 1902. Report on the collections of natural histor\
made during the vovage on the "Southern Cross," Vol.
VII, Mollusca, p, 201-213, 2 pis,
Tait, R, W, 1982, A taxonomic revision of Octopus australis
Hoyle, 1885 (Octopodidae: Cephalopoda), with a rede-
scription of the species. Memoires of the National Museum
Victoria, No, 43:1.5-24,
Toll, R B 1981, Bcnthoctopus oregonae, a new species of
octopod (Mollusca; Ophalopoda) from the southern Ca-
ribbean with a redescription of Bcnthoctopus januarii
(Hoyle, 1885). Bulletin of Marine Science 31(l):83-95.
Voss, G. L. 1973- Present status antl new trends in cephalopod
s\stematics. Svmposium of the Zoological Societv of Lon-
don No. 38:49-60.
THE NAUTILUS 105(3):118. 1991
Page 118
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November 25, 1991
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THEtyNAUTILUS
CONTENTS
Volume 105, Number 4
November 25, 1991
ISSN 0028-1344
Yuri I. Kantor
Alexander V. Sysoev
Mollusks of the genus Antiplanes (Gastropoda: Turridae) /^^•^'\
of the northwestern Pacific Ocean I '^ Z^- 119
A" / •. \
'. 1 : \
Two new species of Vexilhim from the western Pacific y \ ll-/^/^^
(Gastropoda: Costellariidae) - ,- ~ -, — <-" 147
Parasitic mites (Acari: UnionicoUdae) of fresh-water
mussels (Bivalvia: Unionidae) in the Duck and Stones
Rivers in central Tennessee 152
Volutes from Saya de Malha Bank: The saga of Lyria
surinamensis and a new species 159
Mistaken localities for some shells "from Surinam" 165
New species of Gaza, Mirachelus. Calliotropis, and
Echinogurges (Gastropoda: Trochidae) from the
northwestern Atlantic Ocean 166
Acroloxus coloradensis (Henderson), a rare North
American freshwater limpet 173
Gary Rosenberg
Richard Salisbury
Malcolm F. Vidrine
James L. Wilson
Philippe Bouchet
Patrice Bail
Takashi Okutani
James F. Quinn, Jr.
Andrew J. Paul
Hugh F. Clifford
SMITHSONIAN FUNDS FOR MALACOLOGY STUDENTS
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Applications should be sent to:
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Smithsonian Institution
Washington, DC 20560 USA
THE NAUTILUS 105(4): 119-146, 1991
Page 119
Mollusks of the Genus Antiplanes (Gastropoda: Turridae)
of the Northwestern Pacific Ocean
Yuri I. Kantor
Alexander V. Sy§oev
A. N. Severtzov (iistihite ol Animal
Evolutionary Morpholng\ and
Ecology of the
U.S.S.R. Academy of Sciences
Lenin Avenue, 33
Moscow 117071, I'.S.SR.
ABSTRACT
Miillusks of the genus Antiplanes are represented in the fauna
of the northwestern PaciBc Ocean by 11 Recent species and
one subspecies. Onl\ Antiplanes vinosa (Dall) is sinistral; the
remaining members of the genus are dextralK coiled The new
name Antiplanes gahhi is proposed for Plcnrotoma perversa
Gabb, 1865, the type species of Antiplanes and a junior hom-
onym of Pleurotoma perversa Philippi, 1847. Six species: An-
tiplanes abyssalis. A. dendritoplicata, A. obliquiplicata. A. ku-
rilensis. A. spirinae. and A. habei are described as new, as is
the subspecies A. motojimai aquilunalis. The paucity of di-
agnostic features together w ith the high degree of intraspecific
and ontogenetic variability in shell characters requires the use
of statistical analyses for delineation of species. Three groups
of species can be distinguished on the basis of shell sculpture
and the rates of growth of radular teeth. Antiplanes does not
occur in the Gulf of Alaska. Thus, there are no species in
common between the Asian (including Bering Sea) and the
western American (south of the Gulf of Alaska) faunas.
Key words: Ai^tiplanes. Turridae, northwestern Pacific; sub-
littoral, batlnal, radular growth.
INTRODUCTION
Gastropods of the genus Antiplanes are among the larg-
est and most abundant representatives of the family Tur-
ridae in the sublittoral and bathyal zones of the northern
Pacific Ocean. Although many Recent and fossil species
have been described, the validity of many of these ta.\a
is questionable. The present revision of this genus is lim-
ited to the fauna of the northwestern Pacific Ocean from
northern Japan to the Bering Sea, and is based on the
considerable material from this region that was available
to us.
MATERIALS AND METHODS
This study was based primarily on the collections of the
P Shirshov Institute of Oceanology of the U.S.S.R. Acad-
emy of Sciences (Moscow), the Zoological Museum of
the Moscow State University, and the Zoological Institute
of the U.S.S.R. Academy of Sciences (Leningrad). These
samples, containing about 280 lots and over 900 speci-
mens, were collected in the northwestern Pacific over a
period of 90 years. Comparative material from British
Columbia was kindly provided by Dr. R, Shimek (Mon-
roe, Washington). The s>ntypes of Antiplanes beringi
(Aurivillius) were made available by Dr. A. Waren (Na-
turhistoriska Riksmuseet, Stockholm). Photographs of the
types of VV, H. Dalls, E. A. Smith's and T. Habe's ta.xa
were provided by Dr. J. H. McLean (Los Angeles County
Museum of Natural History), Mrs. A. Thompson (British
Museum (Natural History)) and Dr. A. Matsukuma (Na-
tional Science Museum, Tokyo), respectively.
When studying shell variability we counted the num-
ber of spiral cords on the body whorl. Since the number
and degree of cord development on the anal fasciole
were quite variable, only cords below the fasciole were
counted. In some cases the relative height of the body
whorl was an important taxonomic character. In a few
cases in which the upper whorls were lost, ratios of the
body whorl height to the height of the last three whorls
were used. Ratios of the length of the marginal teeth of
the radula to shell height were calculated using the mean
length of at least ten marginal teeth, measured with an
ocular micrometer calibrated to 0.001 mm.
The following abbreviations for scientific research in-
stitutes are used:
BM(NH) — British Museum (Natural History), London
CAS — California Academy of Sciences, San Francisco
NRS — Naturhistoriska Riksmuseet, Stockholm
NSMT — National Science Museum, Tokyo
SBMNH — Santa Barbara Museum of Natural History
SDNHM — San Diego Natural History Museum
UCMP — Museum of Paleontology, University of Cali-
fornia, Berkeley
USNM — National Museum of Natural History, Smith-
sonian Institution, Washington, DC.
Page 120
THE NAUTILUS, Vol. 105, No. 4
Figure 1. Variability of anal sulcus form in Antiplanes saiic-
tiioannis (Smith, 1875). Growth lines along different whorls of
the same specimen are connected by dotted lines.
ZIN — Zoological Institute of the U.S.S.R. Academy of
Sciences, Leningrad
ZM — Zoological Museum of the Moscow State University
ACKNOWLEDGEMENTS
We thank Dr. Alexander Golikov, Zoological Institute,
Dr. Lev Moscalev, Institute of Oceanology, Dr. Dmitri
Ivanov, Zoological Museum of Moscow State University,
and Dr. Ronald Shimek, who kindly provided material
for this study. We are grateful to Dr. Anders Waren,
Naturhistoriska Riksmuseet, for the opportunity to study
the syntypes of Antiplanes heringi, and to Dr. James H.
McLean, Los Angeles County Museum of Natural His-
tory, Mrs. Anne Thompson, British Museum (Natural
History), and Dr. Akihiko Matsukuma, National Science
Museum, Tokyo, who kindly sent photos of type speci-
mens. We are also greatly indebted to Mrs. Saida I. Ja-
gudina, Mrs. Sofia I. Strelzova, and Mrs. Marina A. Dol-
golenko, laboratory assistants at the Zoological Institute,
for their assistance during our studies of their collections.
We want to thank Dr. M. G. Harasewych, Editor, for
his help in checking the manuscript and making it more
clear for the readers.
HISTORICAL REVIEW OF THE
GENUS ANTIPLANES
The first species presently referred to Antiplanes to be
described was Pleurotoma perversa Gabb, 1865. Dall
(1902) proposed the genus Antiplanes and described 17
Recent and fossil species from American and Asian wa-
ters (Dall, 1874, 1902, 1919, 1925). Bartsch (1944a,b)
described three additional American species and isolated
the dextral species of Antiplanes as a separate genus
Rectiplanes. Bartsch began a revision of these groups,
but this was never completed or published. Three Jap-
anese species were later described by Habe (1958) who
also established the subgenus Rectisulcus to include those
dextral species with well-developed spiral sculpture.
Antiplanes differs from most other members of the
family Turridae in containing sinistral forms. The tax-
onomic status of these sinistral forms has been variously
interpreted by recent workers. Based on the study of
extensive collections of fossils and Recent specimens from
California, Grant and Gale (1931) concluded that the
direction of shell coiling was subject to intraspecific vari-
ability and synonymized almost all species known at that
time with A. perversa. This view was shared by Golikov
and Gulbin (1977) and by Golikov and Scarlato (1985).
Bartsch (194a,b), however, considered sinistrality to be
of generic significance.
It was shown in recent malacological literature that
most sinistral forms, including those of Turridae, are
separate species (Tippett, 1983; Kantor, 1990). However,
sinistral individuals and even isolated sinistral popula-
tions lacking taxonomic status have been reported (e.g.,
the sinistral population of Conns mediterraneus, see
Donati et. al. 1984). Analysis of the available material
showed that sinistral Antiplanes from the northwestern
Pacific belong to a single separate species. We have not
found any examples of sinistral individuals or populations
within normally dextral species of Antiplanes.
SPECIES VARIABILITY AND CRITERIA
FOR SPECIES DIAGNOSES
In our revision of this genus, we encountered an unusu-
ally high degree of variability in many conchological
features, along with a general paucity of shell diagnostic
characters. Thus, characters of shell form and proportions
of its parts appeared to be taxonomically insignificant in
most cases. Hickman (1976) documented the taxonomic
importance of anal sinus shape in Turridae. Our analysis
of intraspecific and ontogenetic variation of this feature
in nearly 100 individuals of A. sanctiioam^is as well as
in other species showed a very high degree of variability
overlapping between species (figure 1).
Although we considered a great many potential con-
chological characters, the most taxonomically significant
feature was determined to be the number of spiral ribs
below the anal fasciole. Based on this character, two large
groups of species can be distinguished; one is character-
ized by complete or nearly complete absence of spiral
sculpture (A. ahyssalis n. sp., A. obliquiplicata n. sp., A.
dendritoplicata n. sp. and A. kawamurai). the other,
containing the remaining species treated herein, by the
presence of variably developed spiral sculpture. It should
be emphasized that the number of spiral ribs on the body
whorl, being an ontogenetically and geographically vari-
able feature, allow s onlv statistical differentiation of spe-
cies (figures 117, 118, i38).
Y I Kantor and A. V. Svsoev, 1991
Page 121
Figures 2-10. Radulae of Antiplanes spp. Shell lengths and
catalog numbers are given in parentheses. 2. Antiplanes den-
dritoplicata n. sp., paratype (28.3 mm, ZIN N 6/54581); 3. A.
obliquiplicata n. sp., paratype (40.3 mm, ZM N Lc 6899); 4.
A. spirinae n. sp., paratype (22,4 mm, ZM N Lc 6905); 5. A.
isaotakii (Habe, 1958) (30.8 mm); 6. A. hurilensis n, sp,, para-
type (28.8 mm, ZIN N 1/54583); 7, 8. A. habei n. sp. 7. holotype
(38.0 mm, ZM N Lc 6909); 8. paratvpe (23.1 mm, ZM N Lc
6910); 9, 10. A. vinosa (Dall, 1874). 9. (36.3 mm), 10. (28.6
mm). Scale bars all 100 ixm.
It is interesting to note that similar patterns of mor-
phological variation occur in different species. Specimens
of A. sanctiioannis from Sakhahn Bay have fewer ribs
on their body whorl than individuals from the Kurile
Islands. The same pattern is seen in A. vinosa. a sister
species of A, sanctiioannis.
The morphology of the radula offers additional char-
acters of taxonomic importance. Though the form of the
marginal teeth was similar in all species (figures 2-17),
rates of radular growth permitted species to be segre-
gated into three groups; those with rapidly, moderately
and slowly growing marginal teeth (figure 18). A geo-
graphical component to the variability of this character
was observed. However, comparisons of specimens of the
same size from the same locality showed distinct differ-
ences between species. In figure 18, points representing
different species from the same locality are connected
by dotted lines.
The groups of species isolated on the basis of radular
Figures 1 1-17. Radulae of Antiplanes sanctiioannis (Smith,
1875). The collection locality and shell length of individual
specimens follow 11. Sakhalin Bay, 45.0 mm. 12. Sakhalin
Bay, 41.9 mm; 13. Tatar Strait, 38.0 mm; 14. Sakhalin Bay,
36.0 mm; 15. Southern Kurile Islands, 34.5 mm; 16. Eastern
Kamchatka, 42.0 mm; 17. Tatar Strait, 25.0 mm. Scale bars
all \00 lum.
growth pattern also differed in the degree of spiral sculp-
ture. Species with the most slowly growing marginal
teeth (A. dendritoplicata n. sp. and A. obliquiplicata n.
sp.) generally lack spiral sculpture. Species with mod-
erate marginal tooth growth rates (A. sanctiioannis, A.
vinosa, A. spirinae n. sp., and A. hurilensis n. sp.) are
characterized by more numerous and less developed spi-
ral cords than the group of species with rapid marginal
tooth growth (A. habei n. sp. and A, isaotakii). As with
the variation in marginal tooth growth rates, species and
species groups based on shell sculpture can onK be dis-
tinguished statistically.
In some cases, the relative height of the body whorl
was also useful for distinguishing similar species. Pairs
of species such as A. sanctiioannis and A. hurilensis n.
sp. as well as A. dendritoplicata n. sp. and A. obliqui-
plicata n. sp. (figure 46) were readily distinguished by
this character
The systematics of the genus Antiplanes were difficult
to discern using traditional techniques due to high on-
togenetic and geographical variability of nearly all gen-
erally accepted diagnostic characters. We have been able
to distinguish close but distinct species by using graphs
Page 122
THE NAUTILUS, Vol. 105, No. 4
50
lA
lA
42
40
38
36
34
32
30
28
26
24
22
20
18
16
14
(mm;
0.13 .14 .16
.18
.20 .22
2U
26 .28
mm [
Figure 18. Rales of radular growth in species of Antiplanes.
Crossed circles — A. dendritoplicafa u. sp.; open squares — A.
spirinae n. sp.; open circles — A. obliquiplicata n. sp.; solid cir-
cles— A. sanctiioannis (Smith); open ovals — A. vinosa (Dall);
open triangles — A. isaotakii (Habe); solid triangles — A. habei
n. sp. The lines of regression: 1. A. dendritopHcata n. sp.; 2.
A. sanctiioannis (Smith); 3. A. vinosa (Dall); 4. A. habei n. sp.
Ll — mean length of marginal tooth, Hs — shell length.
and regression analyses of various characters relative to
shell height. This approach forms the basis of the follow-
ing systematic treatment of this genus.
SYSTEMATICS
Family TURRIDAE Svvainson, 1840
Subfamily Turriculinae Powell, 1942
Genus Antiplanes Dall, 1902
Dall, 1902513 (type species, by original designation— P/ph-
roloma (Surciila) perversa (Jabb, 1865).
Rectiplanes Bartsch, 1944b:59 (type species, iiy original des-
ignation— Pleuroloma (Anliplancs) santarosana Dall.
1902).
Rectiplanes (Rectisulcus) Habe, 1958:184 (type species, by
original designation — Rectiplanes {Rectisidcus) motoji-
mai Habe, 1958)
Diagnosis of ihe genus: Shell large (to 72 mm), dextral
or sinistr.il fusiform, covered by smooth, thick perios-
tracum. A.muI sculpture absent. Spiral sculpture may be
absent or coi: posed of ribs and dendritic threads (figure
Figure 19. Diagrammatic representation of the shell base of
Antiplanes habei n. sp. AF — anal fasciole, DL — dendritic ribs,
GL — growth lines, SR — spiral ribs.
19, DL) that may be obliquely and irregularly disposed,
with ramifying small threads radiating from extremities
of anal sinus growth lines. Anal sinus deep, located on
shoulder or near periphery . Siphonal canal long, occa-
sionally slightly curved. Operculum (figures 20-25) large,
drop-shaped, terminally nucleated, filling the aperture.
Radula "nontoxoglossate", consisting of proximally bi-
furcated marginal teeth and of vestigial rachidian teeth.
Type species: Pleitrotoma {Siircula) perversa Gabb,
1865, the originally designated type species of Anti-
planes, is a primary homonym of Pleurotoma perversa
Philippi, 1846. Grant and Gale (1931) considered P. per-
versa Gabb to be a synonym of P. voyi Gabb, 1869.
However, the original descriptions and illustrations dem-
onstrate that these are different species. Indeed, one of
them {P. perversa) is sinistral while the other {P. voiji)
is dextral. As no available name exists for Pleurotoma
perversa Gabb, 1865, we here propose Antiplanes gabin,
nomen novum, as a replacement name. S\ ntypes (UCMP
15929 and 31547) Catalina Island, California (Recent);
(UCMP 15930), Pleistocene of San Pedro, California
(Coan & Bogan, 1988).
Distribution: Species of Antiplanes live in the northern
part of the Pacific Ocean from California to British Co-
lumbia, along the American coast and from Honshu,
Japan to the Gulf of Anadyr, Bering Sea, along the Asian
coast, at sublittoral to upper abyssal depths. In the Bering
Sea, Antiplanes v\ere found in western, eastern and cen-
tral parts. Neither the literature nor samples at our dis-
posal contain records of Antiplanes from the Gulf of
Alaska. According to our data, western American (south-
ward of the Gulf of .■\la-ska) and .\sian (including the
Bering Sea) faunas have no species in conmion.
Y. I. Kantor and A. V. Svsoev, 1991
Page 123
Figures 20-25. Opercula of Antiplanes spp 20. A, obliquiplicata n. sp , holotype, ZM N Lc6898, 40.3 mm 21. A. dendritoplicata
n. sp., paratype, ZIN N 6/54581, 28.3 mm. 22. A. spirinac n. sp., paratype, ZM N Lc 6905, 16.3 mm. 23. A. habei n. sp., paratype,
ZIN N 1/54627, 33.3 mm. 24. A. vinosa (Dall), 36.3 mm. 25. A. sanctiioannis (Smith), 42.7 mm.
Fossil species of Antiplanes are known from the Eo-
cene of North America {Drillia cooperi Dickerson, ac-
cording to Grant and Gale, 1931). In the eastern Pacific,
the genus was frequently found in Pliocene deposits of
Oregon and Washington (Grant & Gale, 1931; Addicott,
1976) and in Pliocene and Pleistocene of California (Grant
& Gale, 1931). In the western Pacific, earliest occurrences
of the genus were recorded from the Middle Miocene of
Japan (A. sadoensis (Yokoyama) — Chinzei, 1959, and A.
sanctiioannis (Smith) — Amano, 1983). Species of Anti-
planes are abundant in Pliocene deposits of Honshu and
Hokkaido, Japan (Masuda & Noda, 1976). Antiplanes
voiji and a dextral Antiplanes sp. were found in the upper
Pliocene to lower Pleistocene of eastern Kamchatka (Pe-
trov, 1982).
Remarks: The genus Rcctiplanes was erected b\ Bartsch
(1944b) to contain dextral species only. No other char-
acters distinguishing Rectiplaiies and Antiplanes can be
identified. Since we do not consider direction of shell
coiling to be of more than a species level character,
Rectiplanes is regarded as a synonym of Antiplanes.
Page 124
THE NAUTILUS, Vol. 105, No. 4
Figures 26-29. Light (26, 27) and SEM (28, 29) micrographs of radulae, 26-28. Antiplanes saiictiioannis (Smith). 29. A. vinosa
(Dall).
Y. I. Kantor and A. V. Sysoev, 1991
Page 125
Figures 30-32. Diagrammatic representations of transverse
sections of the radula in the radiilar sac. 30. Olivella burcalis
Gohkov, 1967. .3 1 . Splcndrillia chatamcnsis S\ soe\' and Kantor,
1989. 32. Antiplanessanctiioanni.s (Smith. 1.S75). cf — "central
formation", ct — central tooth, It — true lateral tooth, "It" — the
so called "lateral" tooth of Turridae. mt — true marginal tooth,
"mt ' — the so called "marginal" tooth ol Turridae.
Habe (1958) established the subgenus Rectisulcus for
dextral species with well developed spiral ribs. As there
is a continuum in the degree of spiral rib development,
we follow Powell (1966), in synonymizing Rectisulcus
with Antiplanes.
Varying opinions e.xist as to the composition of the
transverse rows of teeth in the Antiplanes radula. A
reduced central tooth having the form of a narrow plate
pointed at both ends occurs in the eastern Pacific species
A. santarosana and A. thalaea (Powell, 1966. figures 57,
58). However, McLean (1971) indicated that eastern Pa-
cific species of Antiplanes lacked central teeth. Struc-
tures superficially resembling the central teeth of other
Turriculinae were seen in Antiplanes by means of light
microscopy (figures 26, 27). Only the anterior and parts
of the lateral borders of these optically transparent struc-
tures were clearly seen in unstained preparations (figure
26). When the radula was stained with Orange G and
Aniline Blue, these structures appeared to be paired and
symmetrical (figure 27). However, scanning electron mi-
croscopy showed that there were no distinct tooth plates
on the central part of the radular membrane (figures 28,
29). The membrane had only indistinct folds that are
similar to those of some other Turriculinae.
The manner in which the radular membrane is folded
within the radular sheath is significant for evaluating the
nature of these central structures. The radular membrane
of prosobranch mollusks is folded within the radular
sheath. These folds occur between groups of teeth, or
between separate teeth if the radulae are greatly oligo-
merized. Taenioglossate radulae, for example, have two
pairs of folds, one between the central and lateral teeth,
the other between the lateral and marginal teeth The
same pattern occurs in those Neogastropoda that have
five teeth per transverse row {Olivella. figure 30). How-
ever, in turrids with five teeth per transverse row (sub-
family Clavinae) there is only one pair of folds, placed
between the central and lateral teeth. We have also ob-
served this pattern in two species of Splcndrillia (figure
31).
Study of the radula of Antiplanes shows that the rad-
ular membrane has but a singular fold situated exactK-
along the middle of the "central structure." Moreover,
the membrane can be easily torn along this bend. In our
opinion this suggests that the "central formation" is a
rudiment of a pair of lateral teeth but not of the central
tooth. We propose that the central tooth was completely
reduced during evolution, and that the paired folds of
the membrane have then grown nearer and fused into
a single bend. Maes (1983) proposed a similar evolution-
ary pathway for Crassispira (Crassiclava) apicata (Reeve)
(figures 31, 32), and referred to these structures as "soft
lateral teeth". Although studies of the embryonic devel-
opment of the radula of Antiplanes would shed further
light on this question, such data is presently not available.
The competing hypotheses, one of weakening of the
central tooth and complete reduction of lateral teeth, the
other of loss of the central tooth along with regaining of
the weakened lateral teeth, cannot be fully resolved at
this time.
WESTERN PACIFIC SPECIES
ERRONEOUSLY ATTRIBUTED TO ANTIPLANES
Rectiplanes (?) yukiae Shikama, 1962. Shikama, 1962:
.50, pi. 3, figs. 3a,b, 4a,b.
Type locality: Off Choshi, eastern Honshu — Japan, 250-
300 fms. The species was subsequently reassigned to the
genus Bent hodaphne Oyama. 1962 (Okutani, 1964, 1968).
Rectiplanes (Rectisulcus) hayashii .Shikama, 1977. Shikama,
1977:19, pi, .3, fig. 12a,b.
Type locality: Ensyu-nada. south-western Japan This
species has well developed axial sculpture and is here
referred to the genus Comitas Finlay. 1926.
RECOGNIZED SPECIES OF ANTIPLANES
Antiplanes abyssalis new species
(figures .33-35, 45)
Material examined: Holotype: R/V 'Vityaz', 39th cruise,
sta. 5603, south-east of Simushir, Kurile Islands, 46°22'N,
1.5.3°03'E, depth 3,175-3,250 m, 15.vn.l966 (ZM N Lc
6889); parat\pe R/V "Vityaz', 14th cruise, sta. 2210, south-
east of southern Kamchatka, 50°01.8'N, 157°.39.2'E, depth
2,430-2,670 m, 24.VI.1953 (ZM N Lc 6890).
Description of holotype: Shell slender, with slightly
curved axis, consisting of 6.5 teleoconch whorls and at
least 1.5 heavily eroded protoconch whorls. Whorl profile
angulated at periphery , whorl shoulder concave. Sutures
deep. Body whorl high, comprises about 0.6 of shell height.
Shell base weakly convex, passing gradually and without
bending into a moderately elongate siphonal canal.
Growth lines distinct. Spiral sculpture nearly absent, rep-
resented by single somewhat wavy riblet on anal fasciole
and b\ marked dendritic ribs that reach upper and lower
sutures of spire whorls but disappear on shell base. Anal
sinus wide, rounded. Aperture wide, elongate-oval, poor-
Page 126
THE NAUTILUS, Vol. 105, No, 4
Y. I. Kantor and A. V. Svsoev, 1991
Page 127
ly differentiated from siphonal canal, outer lip attaclied
to columella at acute angle. Columella straight, covered
witfi narrow, white, glossy callus. Color light-yellow.
Periostracum thin, smooth, glossy, fitting closely to hy-
postracum.
Dimensions of holotype: Shell height 31 0 mm, body
whorl height 18.7 mm, aperture height 14.2 mm, shell
diameter 12.0 mm.
Remarks: The paratype differs from the holotype in
having a somewhat darker periostracum, a less concave
profile of the shoulder, and a wider siphonal canal. The
spiral riblet on the anal fascicle is weaker in the paratype.
This species differs from Antiplanes dendritoplicata
n. sp., its nearest relative, in lacking spiral ribs on the
shell base and siphonal canal, and from A. ohliquiplicata
n. sp. in having a higher body whorl as well as in whorl
profile.
Distribution: This species was found along the Kurile-
Kamchatka trench at depths of 2430-3250 m (figure 45).
Antiplanes dendritoplicata new species
(figures 2, 21, 36-44, 45)
IVlaterial examined: Holotype (ZM N Lc 6891) and 3
paratypes (ZM N Lc 6892):' R/V 'Vitvaz', 39th cruise,
sta. 5641, east of Iturup, southern Kurile Islands, 44°46'N,
148°45'E, depth 472-479 m, Sigsbee trawl, 10.1X1966;
paratypes: R/V 'Vityaz', 12th cruise, sta. 1779, north-
eastern Okhotsk Sea, depth 464 m, Sigsbee trawl,
13.x. 1952 (2 specimens— ZM N Lc 6893); 12th cruise,
sta. 1831, northwestern Okhotsk Sea, 56°57.5'N,
145°57.0'E, depth 196 m, Sigsbee trawl, mud, 15. X. 1952
(2 specimens— ZM N Lc 6894); 24th cruise, sta. 3578,
east of Honshu, 38°35.0'N, 142°53.3'E, depth 1,641 m,
Sigsbee trawl, clay, 11. V, 1957 (1 specimen — ZM N Lc
6895); 39th cruise, sta. 5638, east of Iturup, 44°36'N,
149°07'E, depth 1,675-1,845 m, Galathea trawl,
10.1X1966 (1 specimen— ZM N Lc 6896); 52th cruise,
sta. 6671, east of Honshu, 40°12.0'N, 143°35.8'E. depth
2,400-2,720 m, Sigsbee trawl, 23.VI.1972 (1 specimen—
ZM N Lc 6897); Okhotsk Sea, sta. 139/20, 55°45'N,
145°50'E, depth 218 m, sandy mud, 1. VIII. 1930, coll. P.
V. Ushakov (3 specimens— ZIN N 1/54576); R/V To-
porok', sta. 63, southeastern Sakhalin, 46°49.5'N,
143°52.3'E, depth 187 m, 4.IX.1947 (1 specimen— ZIN
N 2/54577); sta. 70, southeastern Sakhalin, 46°26.2'N,
143°22.0'E, depth 103 m, mud with gravel, 6.1X1947 (1
specimen— ZIN N 3/54578); sta. 13-a, northwest of Hok-
kaido, 44°47.6'N, 144°29.8'E, depth 700 m, 26.VIII.1948
Figure 45. Geograpiiir distribution oi Antiplana^ abyssalis u
sp. (open circles), A. dendritoplicata n. sp. (solid circles), A.
ohliquiplicata n. sp, (solid scjuares).
(3 specimens— ZIN N 4/54579); sta. 17, off Shikotan,
43°37.1'N, 147°02.8'E, depth 1,450-1,530 m, Sigsbee
trawl, mud, 28. VIII. 1948 (2 specimens— ZIN N 5/54580);
sta. 100, off Iturup, 44°42.0'N, 147°56.7'E, depth 299 m,
14.1X1949 (1 specimen— ZIN N 6/54581); sta. 101, off
Iturup, 44°20.8'N, 148°24.0'E, depth 414 m, 14.1X1949
(1 specimen— ZIN N 7/54582).
Description of holotype: Shell slender, fusiform, com-
posed of 5 teleoconch whorls and 1.5 heavily eroded
protoconch whorls. Surface of upper 2.5 teleoconch whorls
heavily eroded, rose-colored. Coiling axis slightly curved.
Body whorl high, convex, comprising 0.64 of shell height.
Teleoconch whorls convex, evenly rounded, divided by
deep sutures. Shell base weakly convex, almost flat, pass-
ing into long siphonal canal without pronounced bend.
Growth lines distinct. Spiral sculpture of indistinct, in-
terrupted riblets on anal fasciole (3-4 on body whorl, 2-
3 on penultimate whorl) and of moderately developed
dendritic ribs that reach sutures on spire whorls but be-
come smooth on shell base. Siphonal canal with 8 indis-
tinct spiral ribs separated by shallow grooves. Aperture
Figures 33-44. Shells of Antiplanes spp. 33-.35. Antiplanes abyssalis n. sp. 33. lioiot\pe, ZM N Lc 6889, 3L0 mm. 34. 35.
paratype, ZM N Lc 6890, .36.2 mm. 36-44. Antiplaties dendritoplicata sp. nov. 36, 37. Holotype, ZM N Lc 6891, 414 mm 38.
Paratype, off Shikotan, Kurile Lslands, ZIN N 5/54.580, 35.2 mm. 39. Paratype, eastward off Iturup, southern Kurile Islands, ZM
N Lc 6892, 32.6 mm. 40. The shell base of 42, note the spiral ribs on the canals. 41, 42. Paratype, northwest of Hokkaido, ZIN
N 4/54579, 37.0 mm. 43. Paratype, east of Chonshu, ZM N Lc 6897, 27.0 mm. 44. Paratype, northwestern Okhotsk Sea, ZM N
Lc 6894, 47.7 mm.
Pa,
12t>
THE NAUTILUS, Vol. 105, No. 4
(mm)
40
36 j
32
28
24-
20
16
0.66 .68
.70 .72
%
•76//,
w//y
Figure 46. Comparison of the relative height of the body
whorl of A. dendritoplicata n. sp. (open circles, 1 — line of
regression) and .4. obliquiplicata n. sp. (solid circles, 2 — line of
regression). The shell height is plotted along the abscissa, the
ratio between the bod\ \\ horl height to the height of the last
3 \\ horis is plotted along the ordinate.
high, elongate, oval. Columella slightly thickened by
white, glossy callus. Siphonal canal differentiated from
aperture by its parallel margins. Shell color light-olive.
Periostracum thin, glossy.
Dimensions of holotype: Shell height 41.4 mm, body
whorl height 26.6 mm, aperture height 20.7 mm, shell
diameter 16.0 mm.
Remarks: Paratypes have less convex whorls, more dis-
tinct and closely set riblets on the anal fasciole (up to 4
on spire whorls), a more weakly developed bend at junc-
ture of the body whorl periphery and the shell base, and
a shorter siphonal canal that is better differentiated from
the shell base. Ribs on the siphonal canal of paratypes
may be more numerous (up to 15) and extend onto the
shell base.
Radulae of 6 paratypes were examined (figure 2). Rad-
ular teeth are small, the shell height/mean marginal
tooth length ratio ranges from 159.6 (shell height 22.5
mm) to 266.5 (shell height 47.7 mm). Mean length of
marginal teeth increases slowly. The linear regression
equation is:
Y = 0.1052 -I- 0.0016 X,
where Y is the mean length of marginal teeth and X the
shell height.
Antiplanes dendritoplicata n. sp. is most similar to A.
obliquiplicata n. sp., and the variable shell forms of both
species overlap. However, A. dendritoplicata always has
ribs on the siphonal canal (figure 40). These ribs are
invariably absent in A, obliquiplicata (figure 52). These
two species can also be distinguished by significant dif-
ferences in the relative height of the body whorl (body
whorl height/height of last three last whorls — figure 46).
The relative height of the bod\ whorl decreases linearK
during growth. In young individuals (shell height <30
mm), differences between species are less clearly dis-
cerned, but these two species differ significantly in mean
values. The regression equations are:
A. dendritoplicata
Y = 0.7344 - 0.0011 X
A. obliquiplicata
Y = 0.7485
0.0004 X,
where Y is the relative height of the body w horl and X
is the height of 3 last whorls.
Distribution: This species was recorded from eastern
Honshu to the northern part of the Okhotsk Sea at depths
from 103 m (south-eastern Sakhalin) to 2,720 m (east of
Honshu) (figure 45).
Antiplanes obliquiplicata new species
(figures 3, 20, 45, 47-52)
Material examined: Holotype (ZM N Lc 6898) and 6
paratvpes (ZM N Lc 6899): R/V 'Vityaz', 24th cruise,
sta. 3577, E of Honshu, 38°40.1'N, 143°29.3'E, depth
3042 m, mud, 10. V. 1957; paratypes: R/V 'Vityaz', 5th
cruise, sta. 618, western Bering Sea, 57''03.5'N, 168°30.5'E,
depth 3,875 m, Sigsbee trawl, mud. 25. IX. 1950 (1 spec-
imen—ZM N Lc 6900); 14th cruise, sta. 2209, E of south-
ern Kamchatka, 49°46'N, 157°48'E, depth 3,660 m, Sigs-
bee trawl 23. V. 1953 (4 specimens— ZM N Lc 6901 ); 24th
cruise, sta. 3578, E of Honshu, 38°35.0'N, 142°53.3'E,
depth 1641 m, Sigsbee trawl, clav, 11. V. 1957 (1 speci-
men—ZM N Lc 6902).
Description of holotype: Shell slender, fusiform, com-
posed of 8 whorls. Three upper whorls heavily eroded.
Two upper whorls considerably smaller in diameter,
probably representing protoconch. Bod> whorl convex,
comprising 0.56 of shell height. Shell covered by thin,
olivaceous, somewhat glossy, non-exfoliating periostra-
Figures 47-38. Shells of Antiplanes spp. 47-52. Antiplanes obliquiplicata n. sp 47. 48. Holotype. ZM N LC 6898, 40.3 mm.
49. I'aratype, east of Choiishii, 7A1 N Lc 6899, 36.2 mm 50. Paratype, east of Chonshu. ZM N Lc 6902, 32.0 mm. 5L Paratype.
the same sample as 49, ZM N Lc 6899, 20.8 nnn 52. The shell base of liie hoiot\ pe. note the absence of spiral ribs on the canal.
53-58. Antiplanes kawamurai (Habe, 1958). 53, 54. Holotype, NSMT N 38598. 35.8 nun. 55. Off Shikotan, Kurile Islands, ZIN
N 56.322, 52.1 mm. 56, 57. East of C;honshn, 30.6 mm 58. Tiic same sample as 56, 57. 1 1.7 mm
Y. I. Kantor and A. V. Sysoev, 1991
Page 129
Page 130
THE NAUTILUS, Vol. 105, No. 4
cum. Teleoconch whorls highly convex, evenly rounded,
separated by deeply impressed suture. Shell base ex-
tremelv convex, giving rise to long, broad, curved si-
phonal canal. Growth lines thin, indistinct. Axial folds
oriented parallel to growth lines below anal fascicle on
upper whorls, absent on body whorl. Anal sinus deep,
wide, rounded. Spiral sculpture of well-developed den-
dritic threads that reach sutures on spire whorls and
siphonal canal on body whorl. Spiral riblets (2-3) can be
traced on anal fasciole of upper whorls. Spiral sculpture
absent on shell base antl siphonal canal. Aperture elon-
gate-oval, well differentiated from siphonal canal. Col-
umella concave, covered by white, glossy callus.
Dimensions of holotype: Shell height 40.3 mm, body
whorl height 22.7 mm, aperture height 18.0 mm, shell
diameter 15.5 mm.
Remarks: Radular teeth of the paratype (figure 3) are
small. Mean length of marginal teeth is 0.124 mm (shell
height/tooth length = 325). The shoulders and shell bases
of the paratypes are weakly angular; their whorls may
be more flattened. The anal fasciole and the part of the
whorl nearest to the upper suture may be covered with
thin spiral cords. The canal may be straighter and more
differentiated from the aperture.
This species is most similar to A. dendritoplicata, and
may be differentiated from that species as discussed in
the remarks for that species.
Distribution: Antiplanes obliquiplicata ranges from
Honshu to eastern Kamchatka at depths of 1,641 to 3,875
m (figure 45).
Antiplanes kawamurai (Habe, 1958)
(figures 53-58)
Rectiplanes kawamurai — Habe, 1958:181-184, text-fig. 1; Shi-
kama, 1962:50-51, pi. 3. figs. la,b, 2a; Okutani, 1964:423,
pi. 14. fig. 6; 1968:37, pi, 3, fig. 7.
Type specimens: Holotype (NSMT N 38598), shell height
35.8 mm; paratype (NSMT N 52671), shell height 34.8
mm.
Type locality: Off Choshi, Chiba Pref., Honshu, Japan,
in 183-274 m.
Material examined: Off Shikotan, Kurile Islands,
43°35.5'N, 147°20.5'E, depth 1,450-1,530 m, mud,
28.VII.1948 (1 specimen ZIN N 56322); E of Honshu,
Japan (4 specimens).
Description: Shell light greenish or brownish, broadly
fusiform, with flat-sided spire outline. Body whorl mod-
erately convex, tapering rapidly to join broad, short si-
phonal canal. Whorls weakly convex, angulated below
anal fasciole, nearly turreted in young specimens. Anal
fasciole usually concave, with spiral sculpture absent or
of indistinct threads. Dendritic cords well developed,
reaching suture on spire whorls, occasionally reaching
shell base. Aperture oval, broad, outer lip joining colu-
mella at right or obtuse angle. Shell height reaching 56
mm (Shikama, 1962).
Remarks: This species is easily distinguished from other
species of Antiplanes with weakly developed spiral
sculpture (A. dendritoplicata, A. obliquiplicata, A. abys-
salis) by its broad shell and angulated whorls.
Distribution: Antiplanes kawamurai inhabits the east-
ern coast of Japan from Sagami Bay to the southern
Kurile Islands (Shikotan Island), at depths of 1,520 m
(Sagami Bay— Okutani, 1968) to 183 m (type locality).
Antiplanes sanctiioannis (Smith, 1875)
(figures 1, 11-17, 25, 26-28, 59-88, 100-103)
Plenrotoma (?*) Sancti-Ioannis Smith, 1875:416-417.
PU'urotonia Beringi Aurivillius, 1887:377, Taf. 13, fig. 3.
Plenrotoma {Antiplanes) piona Dall, 1902:514.
Antiplanes piona Dall, 1925:4, pi. 21, fig. 5.
Figures 59-69. SheWs^A Antiplanes spp.59-b2. Plenrotoma sancliioanrnsSmUh. 1875. 59,60. Lectotype, BM(NH) N 1873.8.6,16,
39.2 mm. 61, 62. Paralectotypes, BM(NH) N 1873.8.6.16, 39.9, 23.2 mm, 63-66. Plenrotoma beringi Aurivillius, 1887. 63, 64.
Lectotvpe, NSR N 1557, 38.3 mm. 65, 66. Paralectotypes, NSR N 1557, 37.7, 16.6 mm. 67. Plenrotoma (Antiplanes) piona Dall,
1902, holotype, USNM 109179, 42,0 mm, 68. Antiplanes yessoensis Dall, 1925, holotype, USNM 111053, .39 mm, 69. Antiplanes
willetti Berry, 1953, holotype, 18.4 mm.
Kiguri-s 70-82. Shells of Antiplanes sanctiioannis (Smith, 1875). 70-73. Tatar Strait. Okhotsk Sea (70, 71—39.3 mm, 72—35.1
mm, 73—33,8 mm), 74. Sakhalin Bay, Norlheni Sakhalin, Okhotsk Sea (41 3 mm), 75. Ofi^ Paramushir, northern Kurile Islands
(.38.2 mm). 76. Off Onekotan, central Kurilt- Islands (37.2 mm). 77. Tatar Strait, Okhotsk Sea (32.0 mm) 78. Off lona Island,
northwestern Okhotsk Sea (30.4 mm). 79. Off llurup, southern Kurile Islands (32 I mm). 80. Off eastern Sakhalin, (38.9 mm). 81-
82. Northern part of Okhotsk Sea (81—34.3 mm, 82—40.6 mm).
Figures 83-99. Shells of Antiplanes spp. 83-88. Antiplanes sanctiioannis (Smith, 1875). 83. Tatar Strait. Okhotsk Sea (32.0
mm), 84. Bristol Bay, eastern Bering Sea (34,0 mm), 85-88. Bering Sea (85—31,7 m, 86, 87—29,6 mm, 88—39,9 mm), 89-93.
Antiplanes kurilensis n. sp, 89, 90. Holotyix-, ZM N Lc 6903. 54,6 mm, 91. Paratype, Kurile Islands, ZM N Lc 6904, 52,3 mm.
92. Paratvpe, east off Shumshu, northern Kurile Islaiuls. ZIN N 1 5458.3, 28,8 mm 93. Off Moneron Island, Tatar Strait, sublossil
specimen, ZIN N 2/54584, .35,4 mm 94-99. Antiplanes spirinae n. sp. 94, 95. Holotype, ZIN N 1 54585, 36.5 mm. 96-98.
Paratypes, of Simiishir, central Kurile Islands, ZM N Lc 6905 (96—22.4 mm. 97—14.2 mm, 98—16,3 mm), 99. The shell base of
the holotype.
Y. I. Kantor and A. V. Svsoev, 1991
Page 131
Page 132
THE NAUTILUS, Vol. 105, No. 4
Y. I. Kantor and A. V. Svsoev, 1991
Page 133
Page 134
THE NAUTILUS. Vol. 105, No. 4
Figure 100. Geographic distribution of Antiplanes sanctiioannis (Smith, 1875)
Antiplanes yessoensis Dall, 1925:4, pi 21, fig. 3.
Antiplanes (Rectiplanes) willetti Berrv, 1953419-420, pi 29,
fig. 2.
Antiplanes sadoensis Otuka, 1949:pl 13, fig. 16 {sensu auct.,
non Yokoyama, 1926).
Antiplanes thalaea Dall, 1919:37, pi 11, fig. 6 (sensu auct.,
non Dall, 1902:514),
Type specimens: Lectotype of P. sanctiioannis BM(NH)
1873.8.6.16, shell height 39.2 mm; paralectotypes
BM(NH) 1873.8.6.16, shell height 39.9 mm, 23.2 mm.
Type locality, about 100 miles south-eastward of Yesso
[Hokkaido, Japan].
Lectotvpe of P. beringi (here designated), NRS N 1557,
shell height 38.3 mm; paralectotypes NRS N 1557, 4
specimens. Type locality, 'Vega' Expedition, Bering Sea,
62°39'N, 177°05'W, 55 fms.
Holotype of P. piona USNM 109179, shell height 42
mm. Type locality, off S. coast of Kamchatka, 51°00'00"N,
157°48'00"E, in 96 fms, black sand, "Albatross", sta. 3644
fms.
Holotype of A. yessoensis USNM 111053, shell height
39 mm. Type locality, Japan Sea, S. coast of Hokkaido,
41°58'00"N, 142°30'30"E, in 404 fms, brown mud, "Al-
batross", sta. 5036.
Holotype of A. willetti CAS 064665, shell height 18.4
mm; paratypes— SBMNH No. 34539, and SDMNH
22856. Type locality, off Forrester Island, SW Alaska, 50
fms.
Material examined: Over 200 lots (about 700 speci-
mens) were studied.
Description: Shell very variable, from elongate fusiform
(figures 70, 77) to broadly fusiform (figures 81, 82), white
with periostracum from light yellow to dark-brown. Body
whorl variously convex. Relative height of body whorl
varies within wide range (0.54-0.67). Whorls evenly
rounded to angular at the shoulder and shell base, oc-
casionally, below periphery of spire whorls (form de-
scribed as A. yessoensis). Anal fasciole concave, flat, or
convex. Anal sulcus form varies considerably, even with-
in same specimen (figure 1 ). Spiral sculpture of numerous
relatively narrow ribs, usually weaker at upper part of
the whorl periphery and especially above anal fasciole.
Ribs with serrated borders where crossed by growth lines,
covered with secondary riblets that are usualK incon-
spicuous or absent. Dendritic ribs moderately to highly
developed only on anal fasciole, do not reach sutures on
spire whorls. Siphonal canal usually recurxed, varies from
short, broad (figure 83) to long, narrow (figure 81). Shell
height reaches 60 mm.
Figure 101. The egg capsule of Antiplanes sanctiioannis
(.Smith, 1S75).
Y. I. Kantor and A, V. Sysoev, 1991
Page 135
Figure 102. Comparison of the number of spiral ribs on the
body u horl below the anal fasciole in populations of Anttplanes
sanctiioannis (Smith) The shell height along the abscissa, the
number of spiral ribs along the ordinate Sakhalin Ba\ {open
circles, 1 — line of regression), east Sakhalin (solid triangles, 2 —
line of regression), and Tatar Strait (solid circles, 3 — line of
regression).
Radula relatively large, ratio between shell height and
mean marginal tooth length ranging from 85.4 (shell
height 20.3 mm) to 173.4 (shell height 35.9 mm) (figure
18). Length of marginal teeth increases rapidly during
ontogeny Relationship between marginal tooth length
and shell height nearly linear, expressed by regression
equation:
Y = 0.1401 + 0.0030X
Egg capsules, most containing eggs, attached to shells
of living A. sanctiioannis. Capsules were hemispherical
(diameter — 3 mm), with narrow flange at base. Surface
covered with minute concentric creases, containing
translucent, nearly circular apical plug (diameter — 1 mm)
(figure 101). Three capsules studied each contained up
to 6 developing embryos.
Remarks: Shell morphology of this species is highly
variable. The spiral ribs above the anal fasciole are usu-
ally absent, but may be poorly to well developed. Sculp-
ture on the anal fasciole may consist of broad low ribs
or very thin threads that may become obscure when
Figure 103. Comparison of the number of spiral ribs on the
bod\ whorl below the anal fasciole in populations of Antiplanes
sanctiioannis (Smith). The shell height along the abscissa, the
number of spiral ribs along the ordinate Southern Kurile Island
(solid squares), northern Kurile (solid circles), east Kamchatka
(open circles), and Bristol Bay (open triangles).
dendritic ribs are well developed. The dendritic ribs are
moderately to highly developed only on the anal fasciole
and do not reach the sutures of the spire whorls. Spiral
ribs become stronger, more widely spaced and usually
broader below the anal fasciole. The number of ribs
below the fasciole varies (3-12, usually 6-8, on the penul-
timate whorl) within populations and during ontogeny.
When secondary riblets are well developed, the primary
ribs tend to be low and uneven. Only a single specimen
from the Bering Sea had uniform, equally developed ribs
over the entire shell surface (figures 86, 87), and we refer
this specimen to A. sanctiioannis with some doubt.
Although A. sanctiioannis is highly variable, different
suites of morphological characters predominate in dif-
ferent portions of the species range. Among populations
from the vicinity of Sakhalin Island, those in the Tatar
Strait and adjacent waters have shells with dark color-
ation and distinct spiral sculpture that is more or less
uniformly developed over the entire shell surface but
that weakens slightly towards the suture (figures 70-73).
Dendritic ribs are well-developed in most specimens from
this area, but may be weak or limited to certain portions
136
THE NAUTILUS, Vol. 105, No. 4
Figure 104. Geographic distributinn of Anli)>l(incs kurilensis
n. sp. (open circles) and A. spirinac n. sp. (solid circles).
of the shell. Whorls are evenly rounded. The anal fasciole
is flat or rarely concave. Numerous spiral ribs distinguish
this population.
Specimens from Sakhalin Bay (northwestern Sakhalin)
tend to be relatively light in color, and often have a
convex anal fasciole and inconspicuous sculpture of fewer
spiral ribs (figure 74). Spiral ribs weaken towards the
periphery and dendritic ribs are usually well developed.
Intermediate forms occur along the eastern coast of
Sakhalin and in the adjacent waters of the Okhotsk Sea.
Specimens with angular shoulders and shell bases (figure
80) appear to be restricted to this region. Only there and,
rarely, in the central and northeastern parts of the Okhotsk
Sea do broadly fusiform shells with abruptK narrowing
siphonal canals occur (figures 81, 82).
When these populations are compared on the basis of
a quantitative character (number of spiral ribs below the
anal fasciole of the last whorl), the population from the
Tatar Strait (figure 102, black dots) overlaps little with
the population from Sakhalin Bay. However, specimens
from eastern Sakhalin occupy an intermediate position,
suggesting clinal variation. Specimens found along the
Kurile Islands usually have pronounced spiral sculpture
over the entire shell surface, including the anal fasciole.
Dendritic ribs are generally absent in these populations.
C;olor is light yellowish to yellowish brown. Specimens
with ver\ shallow sinuses were lound only near the Kurile
Islands (figure 1 top, left). Specimens from the Kurile
Islands, Sakhalin Bay, eastern Kamchatka, and the Be-
ring Sea have a similar number of the ribs on the body
whorl (figure 103).
Antiplanes sanctiioannis differs from A. obliquipli-
cata n. sp., A. dendritupticata n. sp., A. abyssalis n. sp.,
and A. kawamtirai in having well developed spiral sculp-
ture that is alwa> s present below the anal fasciole of the
spire whorls. Antiplatics hurilensis n. sp. can be distin-
guished from A. sanctiioannis by its higher spire (body
whorl height/shell height ratio in A. sanctiioannis ranges
from 0.542 to 0.667, mean value = 0.615 ± 0.004, while
in A. kurilensis it varies from 0.500 to 0.533, mean value
= 0.515 ± 0.005) and less elongate shell (shell diameter/
shell height ratio for A. kurilensis ranges from 0.310 to
0.380, mean value = 0.346 ± 0.008, while in A. sanc-
tiioannis it varies between 0.360 and 0.477, with a mean
value of 0.420 ± 0.003). Features that distinguish A.
sanctiioannis from A. spirinae n. sp. include the presence
of spiral ribs with secondary riblets, as well as propor-
tionally longer marginal teeth, which are evident when
comparing equal-sized specimens (figure 18).
Antiplanes sanctiioannis differs from A. isaotakii. A.
motojimai, and A. habei n. sp. in having more numerous
spiral ribs (figure 138), and ontogenetically more slowly
elongating marginal teeth.
Distribution: This widespread boreal species ranges
from Sagami Bay (Honshu) to the Gulf of Anadyr in the
northwestern Bering Sea, and has also been collected in
the Sea of Japan and the Sea of Okhotsk. The bath\ nietric
range of this species extends from 50 m (southern Sa-
khalin) to 1,530 m (southern Kurile Islands). Most spec-
imens are from depths of 100 to 300 m (figure 100).
Ai}tipla7ies kurilensis new species
(figures 6, 89-93, 104)
Material examined: Holotype (ZM N Lc 6903): R/V
Gydrobiolog' Iturup Island (Kurile Islands), depth 55-
125 m, 1982; paratypes: R/\' 'Gvdrobiolog'. Kurile Is-
lands, 1982 (1 shell— ZM N Lc 6904); R/V "Lebed", sta.
Ill, E off Shumshu, northern Kurile Islands, 50°46.9'N,
157°13.4'E, depth 103-104 m, muddy sand, 22. VII. 1954
(1 specimen— ZIN N 1/54583); R/V 'Vityaz', 5th cruise,
sta. 529, off Medny Island, Commander Islands, Bering
Sea, 54°25.0'N, 168°16.4'E, depth 110 m, Sigsbee trawl,
14. VIII. 1950 (5 specimens— ZM N Lc 6907); Expedition
of the Institute of Marine Biology of the USSR Academy
of Sciences, sta. 42, off Moneron Island, Tatar Strait,
46°16.7'N, 141°10.6'E, depth 115 m, 30.VI1I.1972 (1
subfossil specimen — ZIN N 2/54584).
Description of holotype: Shell elongate-fusiform, with
very high spire consisting of 10 whorls. Surface of first
3 whorls eroded. Body w horl moderately convex, com-
prising 0.51 of shell height. Shell light-brown, covered
with thin, smooth, partly eroded periostracum. Teleo-
conch whorls evenly convex, enlarge slowly. Suture shal-
lowly impressed, nearly canaliculate. Incremental growth
lines thin, distinct, producing finely rugose appearance
on shell surface. Spiral sculpture of narrow, non-den-
dritic, slightly wavy cords, 25 between sutures on spire
whorls, about 50 on body whorl below anal fasciole.
Chords evenly developed over entire shell surface. Surface
of cords with pronounced growth lines. Base of shell
Y. I. Kantor and A. V. Svsoev, 1991
Page 137
weakly convex, gradually passing into short, straight,
obliquel) truncated siphonal canal. Aperture oval, nar-
row. Inner lip slightly curved, covered with very thin
callus. Callus better developed on columella. Aperture
comprises 0.35 of shell height.
Dimensions of hololype: Shell height 54.6 mm, body
whorl height 27.7 mm, aperture height 19.2 mm, shell
diameter 17.6 mm.
Remarks: Paratypes have less conspicuous spiral sculp-
ture, especially on the upper part of the whorls, and less
numerous spiral ribs below the anal fasciole of the body
whorl (<40 in paratype with shell height of 52.3 mm).
Weak dendritic threads may be present on or above the
anal fasciole. Only a single specimen from off Shumshu
Island had moderately developed dendritic threads on
the anal fascicle.
The radula (figure 6) of a paratype (shell height 28.8
mm) was studied. Marginal teeth were 0 221 mm long
(shell height/marginal tooth length = 130.3).
Antiplanes kurilensis is most closely related to A. sanc-
tiioannis, but differs in having a higher spire (body whorl
height/shell height = 0.500-0.533, mean value 0.515 ±
0.005 for A. kurilensis, 0.542-0.667, mean value 0.615
± 0.004 for A. sanctiioannis).
Distribution: This species occurs in the northern and
southern Kurile Islands and off Medny Island at 55-125
m. One subfossil specimen was found near Moneron Is-
land (figure 104). A specimen resembling this new species
conchologically was found in Pliocene deposits of north-
eastern Honshu, and was referred to Antiplanes (Rec-
tiplanes) sadoensis (Yokoyama) by Chinzei (1959:pl. X,
figs. 6-7, not figs. 8-9).
Antiplanes spirinae new species
(figures 4, 22, 94-99, 104)
Material examined: Holotype (ZIN N 1/54585): R/V
'Lebed', sta. 122, off Shumshu Island, northern Kurile
Islands, 50°32.5'N, 157°27'E, depth 280 m, sand, gravel
1. VIII. 1954; paratypes: R/V "Odyssey", 34th cruise, sta.
47, off Simushir, central Kurile Islands, 46°44.9'N,
155°2I.0'E, depth 450-480 m, 30.XII.1984 (3 speci-
mens—ZM N Lc 6905, shell heights 22.4; 16.3; 14.2 mm).
Description of holotype: Shell thin, high spired con-
sisting of 9 preserved whorls. Two earliest whorls lost.
Teleoconch whorls evenly rounded, weakly convex, wid-
est below shoulder. Sutures shallow. Bod\ whorl rather
convex, comprising about Vz shell height. Coiling axis
curved. Shell dark olivaceous; shell base, portions of anal
fasciole lighter. Periostracum thin, finely granulated,
peeling. Growth lines very thin, often darker than shell
surface. Spiral sculpture on anal fasciole of 3-4 wide,
unequal spiral cords somewhat broken by growth lines.
Spiral cords less conspicuous toward body whorl. Den-
dritic threads stronger on anal fasciole, but generally
much weaker than in other species of Antiplanes. Re-
maining surface of shell whorls covered by wide, flat-
tened cords that become less conspicuous in later whorls,
nearly absent on body whorl. Entire shell surface covered
by microscopic secondary spiral threads, most conspic-
uously on shell base (figure 99). Shell base weakly convex,
joining short siphonal canal, broken at tip. Aperture short,
narrow, oval Outer lip joining columella at acute angle.
Parietal wall, columella covered by narrow, thin, white
callus.
Dimensions of holotype: Shell height 36.5 mm, body
whorl height 17.9 mm, aperture height 12.2 mm, shell
diameter 12.0 mm.
Remarks: The paratypes have flatter whorls and less
developed spiral cords on the shell surface and on the
anal fasciole. The secondary threads are better developed
and dendritic threads are practically absent in the para-
types.
Radulae of two paratypes were studied (figure 4). The
mean length of the marginal teeth of the specimen with
a shell length of 16.3 mm was 0.166 mm (shell height/
marginal tooth length length = 98.2). The corresponding
values for the 22.4 mm specimen are 0.169 mm and
132.5.
Antiplanes spirinae is most similar to A. sanctiioannis,
but differs in having a narrower shell with a higher spire
and less convex whorls, as well as spiral sculpture of
poorly developed spiral cords and well developed sec-
ondary threads. The marginal teeth of A. spirirtae are
generalK shorter than in A. sanctiioannis (figure 18,
specimens from the same localities connected by a dotted
line).
Etymology: This species is named in honor of Dr. N.
Spirina, who collected the holot\pe.
Distribution: This species is known from the northern
and central Kurile Islands at depths of 280-480 m (figure
104).
Antiplanes vinosa (Dall, 1874)
(figures 9, 10, 24, 29, 105-120)
Pleurotonia vinosa Dall, 1S74:.553.
Pleurotoma (Antiplanes) vinosa Dall, 1902:514, pi. 34, fig. 4.
Antiplanes kamchatica Dall, 1919:33-34, pi. 10, fig. 1.
Antiplanes eontraria Yokoyama, Kira, 1965:100, pi. 36, fig. 2.
Type specimens: Holotype of Pleurotoma vinosa, USNM
220899, shell height 36 mm. Type locality — Kyska Har-
bour, Great Kyska Island, Aleutian Islands, 10 fms.
Holotype of Antiplanes kamchatica, USNM 225255,
shell height 52 mm. Type localit\ — south-eastern coast
of Kamchatka, 100 fms.
Material examined: 40 samples (more than 150 speci-
mens) were studied.
Description: Shell sinistral, variable, usually with high
spire. Color dark-brown to light yellow. Body whorl com-
prises 0.53 to 0.61 of shell height. Shell axis often curved.
Whorls weakly to moderately convex, evenK rounded.
Growth lines inconspicuous. Spiral sculpture of numerous
Page 138
THE NAUTILUS, Vol. 105, No. 4
Y. I. Kantor and A. V. Sysoev, 1991
Page 139
Figure 117. Comparison of the number of spiral ribs on tfie
body wfiorl below the anal fascicle in Sakhalin Bay populations
of Antiplanes vinosa (Dall) (solid circles, right regression line)
and A. sanctiioannis (Smith) (open circles, left regression line).
The shell height along the abscissa, the number of spiral ribs
along the ordinate
variably developed wavy spiral cords spanning shell sur-
face, most pronounced on siphonal canal. Cords some-
times indistinct on and above anal fasciole. Dendritic
threads variable in development, usually present on the
anal fasciole, but not reaching suture on spire whorls.
Aperture narrow, oval, usually well-differentiated from
long, straight siphonal canal.
Radula medium sized for genus, shell height/mean
marginal tooth length varies from 111.6 (shell height 21.2
mm) to 158.4 (shell height 48.8 mm). Increase in mar-
ginal tooth length with shell growth similar to that of A.
sanctiioannis; regression equation: Y = 0.0942 -I- 0.0043X
(figure 18).
Figure 118. Comparison of the number of spiral ribs on the
body whorl below the anal fasciole in the Kurile populations
of Antiplanes vii^osa (Dall) (solid circles, 2 — line of regression)
and A. sanctiioannis (Smith) (open circles, 1 — line of regres-
sion). The shell height along the abscissa, the number of spiral
ribs along the ordinate.
Remarks: Antiplanes vinosa is the only sinistral species
of Antiplanes in the northwestern Pacific Ocean. Anti-
planes sanctiioannis is the most closely related dextral
species. The number of spiral cords below the anal fas-
ciole on the body whorl of A. vinosa is greater than in
A. sanctiioannis (figure 117). Moreover, the spire of A.
vinosa is generally higher than that of A. sanctiioannis,
even in sympatric specimens (figure 119). In Sakhalin
Bay, the ratio body whorl height/shell height is 0.57-
0.62 (mean 0.589 ± 0.006) for A. sanctiioannis. and
0.53-0.57 (mean 0.553 ± 0.006) for A. vinosa. Regression
equations are Y = 0.5852 - 0.0007X and Y = 0.6346 -
Figure§ 105-116. Shells of Antiplanes vinosa (Dall, 1874). 105. Hoiotype of Plcuwtoma vvwsa Dall, 1874, USNM 220899, 36
mm 106. Hoiotype of Antiplanes kamchatica Dall, 1919, USNM 925255, 52 mm. 107. Sakhalin Bay, Okhotsk Sea, 45 2 mm.
108. Tatar Strait, 28 5 mm 109. Central Kurile Islands, 51 7 mm 110. Off Moneron Island, Tatar Strait, 42 0 mm 111. Off
Paramushir, northern Kurile Islands, 46.5 mm. 112. Northern Okhotsk Sea, 53.7 mm. 113. Off Shikotan, southern Kurile Islands,
40,9 mm. 114. Eastern Kamchatka, .35 3 mm. 115. Off Onekotan Island, central Kurile Islands, 29.7 mm 116. Southern Kurile
Islands, 31.3 mm.
Page 140
THE NAUTILUS, Vol. 105, No. 4
(mm)
60-1
50
(0
30
20
0.50 .52 .54 .56 .58 .60 .62
%
Figure 119. Comparison of the relative height of the body-
whorl of Antiplanes vinosa (Dall) (solid circles, 2 — line of re-
gression) and A. sanctiioannis (Smith) (open circles, 1 — line of
regression). The shell height along the abscissa, the ratio be-
tween the body whorl height and the shell height along the
ordinate.
0.00 13X respectively, where Y is the ratio of body whorl
height to shell height and X is the shell height.
Distribution: Antiplanes vinosa was recorded from the
northeastern part of Honshu (Kira, 1965), the northern
Japan Sea to the northern Okhotsk Sea, and from the
Bering Sea (figure 120).
Antiplanes isaotakii (Habe, 1958)
(figures 5, 121, 123-125, 137, 138)
Rectiplanes {Rectisulcm) isaotakii Habe, 1958:182, 185-186,
text. fig. 2; Okutaiii 1964:424.
Type specimens: Holotype, NSMT 38591, shell height
28.5 mm; paratype, NSMT 52684. Type locality, of Cho-
shi, Chiba Pref., Honshu.
Material examined: Off Kurile Islands (1 specimen); off
southern Kurile Islands (1 specimen).
Description: Shell light-olivaceous, elongate, high spired.
Body whorl weakly convex, comprising 0.57-0.62 shell
height. Spire whorls weakly to moderately convex, slight-
ly angulated at periphery, flattened at anal fasciole. Su-
ture shallow. Siphonal canal long, slightly curved. Ap-
erture narrow, short, comprising 0.44-0.52 of shell height.
Outer lip joining columella at nearly right angle. Spiral
sculpture well developed, comprised of 2-3 flattened to
convex cords between suture and anal fasciole, <5 cords
on fasciole, sometimes with traces of dendritic threads,
20 wide, rounded cords, uniformly developed over entire
body whorl below anal fasciole. Earlier whorls with 4-
5 cords below anal fasciole.
Radula of 30.8 mm specimen was studied (figure 5).
Marginal teeth were large (mean length = 0.289 mm,
shell height/tooth length ratio = 106.6) (figure 18).
Remarks: Antiplanes isaotakii is conchologically sim-
ilar to some forms of A. sanctiioannis, but differs in
having longer marginal teeth and fewer spiral cords that
are more convex, wider, and more evenly spaced. It
differs from A. habei in having fewer spiral cords and a
more acutely angled periphery.
Distribution: Antiplanes isaotakii was recorded off
Chiosi, Honshu, in Sagami-Bay and off the southern Ku-
rile Islands in 200-780 m (figure 137).
Antiplanes motojimai motojimai (Habe, 1958)
(figures 122, 137)
Rectiplanes (Rectisutciis) motojimai Habe, 1958:182-185, fig.
3; Okutani, 1964:424. pi. 4, fig, 7,
Type specimens: Holotvpe, NSMT 30600, shell height
38.2 mm; paratypes, NSMT 30600a-b, shell heights 37.0,
35.5 mm. Type locality — off Chiosi, Chiba Prefecture,
Honshu, Japan.
Description: "Shell elongate with about eight whorls
which are moderately inflated but deeply constricted at
the suture, covered with the olivaceous periostracum;
surface sculptured with the spiral cords, two or three of
which are contained in the area between the rather broad
fasciole and the upper suture and also two or three in
the area between the fasciole and the lower suture; body
whorl very large and stout, occupying about two-thirds
of the shell length, the periphery of which is rounded
and the base is marked by the spiral cords all over;
aperture wide, the outer margin broadly sinuated in
V-shape at a little above the periphery; canal broad and
short; columellar margin white." (Habe, 1958:184).
Remarks: This species is most similar to A. isaotakii.
but differs in having a broader shell and fewer spiral
cords (21 cords below the anal fasciole on the body whorl
of the holotype) that become narrower towards the si-
phonal canal.
Distribution: This subspecies occurs from Sagami Bay
to the Sea of Kashima Nada (eastern Honshu), in 550-
870 m (Okutani, 1964). It is not represented in collections
of USSR institutions.
Antiplanes motojimai aquilonalis new subspecies
(figures 126, 127, 137)
Material examined: Holotvpe (ZM N Lc 6906): R/V
'Adler', 18th cruise, sta. 169, eastern Bering Sea, 62°59'N,
Y, I. Kantor and A. V. Sysoev, 1991
Page 141
Figure 120. (Jeographic distribution of Antiplanes vinosa (Dall).
171°05'W, depth 46 m, 1983; paratypes: R/V 'Vityaz',
2nd cruise, sta. 20. southern Okhotsk Sea, 45°01.1'N,
144°21.0'E, depth 429-404 m, muddv sand and gravel,
10. VIII. 1949 (empty shell— ZM N Lc 6908); R/V 'Aka-
demik Oparin', 7th cruise, sta. 17, north-eastern Okhotsk
Sea, 56°00'N, 146°29'E, depth 397 m, 1988 (empty shell—
ZIN N 1/54624).
Description of holotype: Shell broad, consisting of 6
whorls, with earliest 1.5 whorls lost, next 3 whorls eroded.
Spire whorls evenly convex, with narrow, flattened sub-
sutural band above anal fasciole. Suture deeply im-
pressed. Shell color dark olive. Periostracum thick, glossy.
Growth lines numerous, some raised, rough, forming
raised flat tubercles along lower parts of whorls. Spiral
sculpture well developed, with 4-5 distinct, flattened ribs
below anal fasciole on spire whorls. Body whorl with two
narrow cords adjacent to anal fasciole, next 4 cords 1.5
times as broad, remaining 10 cords becoming progres-
sively narrower towards siphonal canal. Cords flat, with
serrated borders at intersections with growth lines. Anal
fasciole with 2-3 thin threads, separated by wide inter-
spaces. Dendritic threads well developed, reaching su-
ture on spire whorls, disappearing on shell base. Anal
fasciole with very thin secondary threads, especially on
spire whorls. Siphonal canal broken. Shell base eroded.
Aperture narrow, oval.
Dimensions of holotype: Shell height 35.5 mm, body
whorl height 21.6 mm, aperture height 14.2 mm, shell
diameter 16.5 mm.
Remarks: Paratypes are very similar to the holotype,
but differ in having three ribs above the anal fasciole.
This new subspecies differs from the nominatypical one
in spiral sculpture: while Antiplanes motojimai moto-
jimai has equally developed cords on the periphery of
the whorls and on the anal fasciole, Antiplanes moto-
jimai aquilonalis lacks wide ribs on the fasciole. These
two subspecies differ in their geographical distributions.
Distribution: This new subspecies was recorded in the
Okhotsk and Bering Seas at depths from 46 m (Bering
Sea) to 550 m (southern Okhotsk Sea) (Fig. 137).
Etymology: aquilonalis Latin — northern, from Aquilo
Latin — northern wind.
Antiplanes habei new species
(figures 7, 8, 23, 128-135, 137, 138)
Material examined: Holotype (ZM N Lc 6909): R/V
'Vityaz', second cruise, sta. 100, central part of the Okhotsk
PUL
THE NAUTILUS, Vol. 105, No. 4
Figures 121, 122. Shells of A;!np/(i«« spp. 121. Rectiplanes
isautakii liahe. 19.58. holot\pe, NSMT \ .526S4. 28..5 mm. 122.
Rectiplanes (Rectisulcus) molojimai Habe, 1958, holotype,
NSMT N 38600, 38.2 mm.
Sea, 54°35.0'N, 149°49.5'E, depth 543 m; paratypes: R/V
"Vityaz', 12th cruise, sta. 1853, central part of the Okhotsk
Sea,' 55°38,5'N, 143°01.0'E, depth 389 m, mud (1 spec-
imen—ZM N Lc 6910); R/V "Adler", 18th cruise, sta. 42,
Okhotsk Sea, 49°12.7'N, 144°57.6'E, depth 245-260 m
(1 specimen— ZM N Lc 6911); R/V 'Gagara', sta. 217,
Okhotsk Sea, 51°35'N, 154°46'E, depth 410 m, (1 spec-
imen—ZIN N 3/54625); R/V 'Gagara', sta. 252, Okhotsk
Sea, 55°32.4'N, 149°14'E, depth 335 m (3 specimens—
ZIN N 2/54626); R/V 'Toporok', sta. 17, off Shikotan,
43°32.5'N, 147°20.5'E, depth 1,450-1,530 m (2 speci-
mens—ZIN N 1/54627).
Description of holotype: Shell elongate-fusiform, with
high spire and high, weakly convex body whorl that
comprises 0.57 of shell height. Shell of seven whorls, with
at least one upper whorl lost. First preserved whorl erod-
ed. Shell yellowish, with darker spiral band below anal
fasciole. Periostracum thin, gloss>. Whorls weakK con-
vex, with flattened periphery nearly parallel to shell axis.
Sutures impressed, shallow. Incremental growth striae
thin, inconspicuous, some marked by color. Spiral sculp-
ture of inconspicuous dendritic thread-like riblets on anal
fasciole (4-5 on penultimate whorl, 3 on body whorl)
reaching sutures on spire whorls and disappearing at
periphery of body whorl. Ribs, 2-3 above fasciole, 4-5
below fasciole on three upper whorls, up to 9 on other
spire whorls, 31 below fasciole on body whorl, becoming
more distinct toward siphonal canal. Five ribs below anal
fasciole flattened and of equal width, subsequent ribs
more narrow and raised. Shell base flattened, joining
short siphonal canal with distinct bend. Aperture narrow,
short, comprising 0.43 of shell height. Outer lip joins
columella at obtuse angle. Callus thin, narrow, white,
glossy, covers columella and parietal wall. Siphonal canal
poorK' differentiated from aperture.
Dimensions: Holot\pe, shell height 38.0 mm, body whorl
height 21.6 mm, aperture height is 16.3 mm, diameter
13.6 mm.
Remarks: Paratypes have slightly more convex and
rounded whorls, wider apertures, more distinct spiral ribs
below the anal fasciole, and wide, flat ribs on the fasciole
that are absent on the body whorl. Limited material does
not allow an evaluation of the range of intraspecific vari-
ability. We provisionally refer to this species two spec-
imens from off Shikotan that differ from other paratypes
in having; rounded or angulated, but never flattened
whorls, a longer siphonal canal, high anal sinus apex,
different geographic (southern Kuriles rather than the
Okhotsk Sea) and bathymetric (1,450-1,530 m vs. 335-
575 m) distributions.
The radulae of four specimens were studied (figures
7, 8). The marginal teeth are large; the ratio between
the shell height and the tooth length varies from 76.3
(the shell height 14.2 mm) to 118.9 (37.8 mm). Their
length rapidly increases with shell growth according to
the regression equation:
Figures 12,3-136. Sht-lis of Antiplanes spp 123-12.5. Antiplanes isaotahii (Habe. 1958). 123. 124. ■' Kurilt- Islands, 31 0 mm
125. Soiitlu-rii Kurik- Islands. 21.3 mm. 126, 127. Anliplunes ruotojimai aqtiilonalis n. subsp. holotype, ZM N Lc 6906, 35.5 mm
128-135. Antiplanes hahci n sp. 128. llolotvpe, ZM N Lc 6909, 38 0 mm 129. 1.30. Paratvpe. Okhotsk Sea, ZM N Lc 6911.
29 0 mm. 131, 132. Central part of Okhotsk Sea, ZM N Lc 6910. 23 0 mm 133-135. Off Shikotan. Kurile Islands, ZIN N 1/54627
(133—33.5 mm, 134, 135—23.2 mm). 136. Antiplanes IniHmoides Dall. 1919 Holot\pe, 31 mm.
Y. I. Kantor and A. V. Svsoev, 1991
Page 143
Page 144
THE NAUTILUS, Vol. 105, No. 4
Figure 137. Geographic distribution of Antiplanes isaotakii (Habe) (solid circles), A. motojimai motojimai (Habe) (open circles)
A. motojimai aqiiilonalis n. subsp. (solid squares), and A. hahei n. sp. (open circles).
Y = 0.1290 + 0.0052X.
Antiplanes hahei is most similar to A. sanctiioannis, but
differs in having more convex, wider and less numerous
ribs (figure 138) that are nearly equal over the body
whorl. The ribs of A. sanctiioannis are considerably more
pronounced on the shell base and the siphonal canal.
This new species has large marginal teeth (figure 18),
which is most clearly seen when comparing equal-sized
specimens from the same locality (the respective points
are connected by dotted line in figure 18). Antiplanes
hahei also resembles A. isaotakii, but differs from it in
having a deeper suture, more convex whorls, more nu-
merous ribs on the body whorl, as well as in lacking the
angular periphery.
Distribution: Antiplanes hahei was recorded in the
Okhotsk Sea and the southern Kurile Islands at depths
from 335 m (central part of the Okhotsk Sea) to 1,530
m (off Shikotan) (figure 137).
Etymology: This new species is named in honor of Dr.
Tadashige Habe, who has considerably widened our
knowledge of the north-western Pacific species of An-
tiplanes.
Antiplanes hulimoides Dall, 1919
(figure 136)
Dall. 1919:.34, pi II. fig, 7; Dall, 192,5:4, pi .31, fig. 2.
Material examined: Holotype (USNM 111051), U.S.
Bureau of Fisheries, Station 4772 on Bowers Bank, Bering
Sea, 54°30'30"N, 179°13'00"E, in 344 fathoms, green-
brown sand, bottom temperature 38.1°F, 6. IV. 1906.
Description: "Shell elongate, decollate. Whorls six or
more, four remaining. Suture distinct, not appressed.
Whorls moderateK convex, smooth, with pale polished
greenish periostracum, in spots minutely granulose, ap-
parently from some wrinkling of the periostracum. Anal
sulcus wide, shallow, bareK forming a fasciole. Outer lip
thin, sharp, moderately produced. Inner lip with a thin
white layer of callus. Columella straight, with an oblique
anterior attenuation. Canal wide, hardly differentiated.
Length of three complete whorls, 31 mm. Length of last
whorl, 23 nun. Diameter at apex, 5 mm. Maximum shell
diameter, 15 mm."
Remarks: So far as we know , this species is known only
from the holotype, which is figured and described here.
Y. I. Kantor and A. V. Sysoev, 1991
Page 145
(mmf
^5
M)
35
30
25
20-
20
30
i.0
50
K
Figure 138. Comparison of the number of spiral ribs on the
body whorl below the anal fascicle in populations of Antiplanes
isaotakii (Habe) (open squares), A. habei n. sp. (solid circles,
1 — line of regression), and A. sanctiioannis (Smith) (open cir-
cles, 2 — line of regression). The shell height along the abscissa,
the number of spiral ribs along the ordinate.
The bulimoid shape and gUstening greenish periostra-
cum makes this species quite unique.
LITERATURE CITED
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THE NAUTILUS 105(4): 147-151, 1991
Page 147
Two New Species of Vexillum from the Western Pacific
(Gastropoda, Costellariidae)
Gary Rosenberg
Academy of Natural Sciences
19th and the Parkway
Philadelphia, PA 19103
Richard Salisbury
8807 Craydon Drive
Boise, ID 83704
ABSTRACT
Vexillum brunneolinea n sp. is described from the Palau Is-
lands and compared to Vexillum acuminatum (Gmelin, 1791)
and V. semisculptum (Adams & Reeve, 1850)^ It is the first
member of its genus reported in which some individuals lack
apertural lirations. Vexillum elliscrossi n sp is described from
Hawaii and central Pacific guyots and compared to Vexillum
daedalum (Reeve, 1845) and Vexillum xenium Pilsbry, 1921.
Key words:
Vexillum.
Costellariidae; Hawaiian Islands; Palau Islands;
INTRODUCTION
We first became aware of the two Vexillum species de-
scribed in this paper more than twelve years ago. Our
research in the literature at intervals during those years,
involving study of the original descriptions of more than
two thousand living and fossil species of miters (mitrids
and costellariids), failed to uncover names for these taxa,
although during that period both species were illustrated
under erroneous names. We have not assigned the species
we describe to subgenera because the limits of the sub-
genera of Vexillum are poorly defined and will surely
change as anatomical knowledge of the group increases.
On the basis of shell characters, it is possible that Vex-
illum brunneolinea n. sp. will be assigned to the subgenus
Costellaria, and V. elliscrossi n. sp. to the subgenus Pusia.
SYSTEMATIC DESCRIPTIONS
Family Costellariidae MacDorald, 1860
Genus Vexillum Roding, 1798
Vexillum brunneolinea new species
(figures 1-3)
Vexillum (Costellaria) acuminaturr^ i politum "Reeve" Tur-
ner, 1989, pp. 12, 28, pi. 5, fig. 21, right hand specimen
only; not Mitra politum Reeve, 1844
Description: Adult shell length ranges from 17.4 to 25.5
mm; width ranges from 28 to 33 percent of length, and
spire height from 37 to 42 percent of shell length. A
summary of measurements is given in table 1. The shell
is white with 29 to 57 fine brov\ n axial lines visible through
a pale yellow periostracum. In live-collected specimens
the periostracum is greenish-yellow and opaque. The
green color may be due to the presence of commensal
algae, as it rapidly fades after collecting.
The protoconch and first three or four teleoconch whorls
are dark rust in color. The protoconch is smooth, about
0.5 mm in diameter and consists of about 2.2 whorls; the
teleoconch consists of 8.9 to 10.5 convex whorls. The
upper whorls have 12 to 20 axial ribs per whorl, typically
13 to 16. These ribs become obsolete by the penultimate
whorl, which is smooth. On the upper three teleoconch
whorls, there are 3 or 4 spiral grooves, which in some
cases increase to five in number on succeeding whorls.
These grooves are visible as pits between the axial ribs,
but do not cross the ribs; they usually become obsolete
by the penultimate whorl, but are still faintly visible on
the body whorl in some specimens. There are three to
seven spiral cords above the fasciolar ridge, and four to
six below it on the siphonal fasciole. In some specimens
the cords on the fasciole are obsolete or absent. The
aperture and columella are white. There are four pli-
cations on the columella; the strongest is the posterior-
most, which ranges up to 0.40 mm in width distalK' in
adult specimens. There are 0-13 apertural lirations. In
some specimens the lirations are of normal strength for
Vexillum, in others they are shortened, reduced in num-
ber or in length, or, in seven specimens, are entirely
absent. To make sure that lirations were lacking, a high
intensity light was shone through the body whorl opposite
the aperture. This procedure shows any trace of a liration
in high relief, and none were detected in these specimens.
The lack of apertural lirations has not been documented
in any other Vexillum species.
The radula of the holotype is preserved on a slide
prepared at the Academy of Natural Sciences (ANSP)
between 1955 and 1958 by the late Virginia Orr Maes;
it is typical of Vexillum species as illustrated by Cer-
nohorsky (1970). The radula is fragmentary, with only
26 rows of teeth represented. Its width is 0.14 mm. The
spacing between rows is about 0.025 mm, so there would
have been about 40 rows of teeth per millimeter of ribbon
length. The rachidian teeth have eight to ten cusps; the
Page 148
THE NAUTILUS, Vol. 105, No. 4
Figures 1-3. Vexillum hrunneolinea new species. 1. Holotype, length 25.5 mm, ANSP 217492, with periostracum. 2. Paratype,
length 21 mm. Salisbury collection, periostracum removed ,3. ,.\pertural detail of liolotype, aperture length 115 mm Figures 4,
5. Vexillum acuminatum (GmeHn, 1791) 4. Form polilum (Reeve, 1844), length 20.5 mm, ANSP 28635, Philippines. 5. Apertural
detail of large individual, aperture length US. 3 mm, ANSP 29699. Sri Lanka Figures 6-9. Vexillum clli.scrossi new species 6.
Holotype, length 18.5 mm, KPHM 219991. 7. Paratype, ape.x, coated with magnesium o.xide, IMT Sta. 73-15. 8. Same specimen
G. Rosenberg and R. Salisbury, 1991
Page 149
lateral teeth are smooth. The rachidians are 0.08 mm in
length, the laterals 0.06 mm; the ratio of their lengths is
1.33. Radular statistics are all within the ranges reported
by Cernohorsky (1970:10) for Vexillttrn.
Type material: HOLOTYPE. ANSP 217492 (radula slide
937), 25.5 X 7.5 mm. PARATYPES. ANSP 202887 (1
specimen); ANSP 203082 (3 specimens); ANSP 203341
(2 specimens); ANSP 203787 (2 specimens); ANSP 217471
(2 specimens, bodies in ethanol); ANSP 382159 (1 spec-
imen). Salisbur\' Collection, 7 specimens. Paratypes total
eighteen specimens.
Type locality: Dredged in sand at 9 meters, inside the
outer reef 12.9 km WNW of Koror Island, Palau Islands,
Western Carolines. 7°24'N, 134°21'E. Collected by A. J.
Ostheimer, 3rd; Station 441-6, 21 August 1955.
Distribution: All known specimens have been taken in
the Palau Islands. Data for ten paratypes at ANSP that
were dredged by A. J. Ostheimer, 3rd in 1955 are as
follows:
ANSP 202887— Sta. 452. at 31 meters, 3.2 km NE of
Gamudoko Island off Urukthapel Island, 24 August.
7°17.5'N, 134'=2I.5'E.
ANSP 203082— Sta. 436, in sand at 27 meters, Malakal
Harbor, 20 August. 7°19'N. 134°27'E.
ANSP 203341, 217471— Sta. 361, in clean fine sand at 8
to 12 meters, in bay on inner edge of barrier reef 1.6
km south of West Passage, Babelthuap Island, 16 Julv.
7''31'N, 134°29'E.
ANSP 203787— Sta. 450, at 33 meters, east of Yoo Pas-
sage, 4.8 km NE of Eil Malk, 24 August. 7°12'N,
134°25.7'E.
The other paratypes were collected by Dieter Cosman
using SCUBA in April 1978 in sand at 5.5 meters in the
rock islands around Koror. One of these is ANSP 382159,
the other seven are in the Salisbury collection. Turner
(1989, pi. 5, fig. 21, right-hand specimen) published a
color photograph of a 24 mm specimen in the Ted Baer
collection taken at 10-20 meters in sand and fine rubble
in Palau.
Etymology: "Brunneolinea" is derived from Latin
brunneus (brown) and linea (line) referring to the brown
axial lines characteristic of the species. It is a noun in
apposition, and so does not change endings to match the
gender of the genus.
Discussion: This species was considered a variant of the
poiitum form of Vexiltum acuminatum (Gmelin, 1791)
by Turner (1989, p. 12). Vexillum brunneolinea is easily
distinguished from typical Vexillum acuminatum as the
latter is brown in color, has a light band below the suture
and has axial ribs on the body and penultimate whorls.
Vexillum acuminatum form poiitum (Reeve, 1844) (fig-
Table 1. Measurements (mm) and counts of conchological
features of Vexillum brunneolinea n. sp. Number of specimens
varies because in some cases values for juvenile, damaged and
worn specimens were omitted or could not be determined.
Mean
Range
N
Siiell length
21.3
17.;3-25.5
10
Bods whorl length
13.0
10.1-15.6
10
Bod\ whorl width
6.4
5.0-7.6
10
Spire length
8.3
7.1-9.9
10
Aperture length
9.5
7.3-11.5
10
.\perture width
2.3
1.7-2.8
10
Plica 1 width
0.32
0.20-0.40
10
Teleoconch whorls
9.6
8.9-10.5
10
Ribs, whorl 1
12.8
12-14
14
Ribs, whorl 2
13.6
12-16
19
Ribs, whorl 3
14.5
13-17
19
Ribs, whorl 4
14.9
13-18
19
Ribs, whorl 5
16.0
13-19
18
Ribs, whorl 6
15.9
13-20
17
Protoconch width
049
0.47-0.52
15
Protoconch whorls
2 2
2.1-2,4
6
ure 4) is smooth on these whorls, and sometimes develops
a pattern of brown axial lines on a grayish-brown back-
ground. However, these lines are broader and more wide-
1\ spaced than those in V. brunneolinea. The lines are
darkly pigmented areas along growth lines of the shell,
whereas the axial lines in V. brunneolinea are somewhat
flexuous and in some cases bifurcate; they do not follow
the growth lines.
Another feature that separates these species is that the
posteriormost columellar tooth in fully mature individ-
uals of V. acuminatum is broader than that in V. brun-
neolinea, and is often grooved at the top, giving it a
biplicate appearance (figure 5). This tooth is convex in
outline in V. brunneolinea. Vexillum acuminatum has
been recorded from Palau, so V. brunneolinea cannot
be considered a geographical variant of it.
Some forms of Vexillum semisculptum (.Adams &
Reeve, 1850) are also morphologically similar to V. brun-
neolinea. The species is usually ribbed, but as in V.
acuminatum, smooth variants are known. However, the
coloration of V. semisculptum is a uniform greenish,
bluish or grayish brown with a white line below the
suture. The aperture is brown except for the white line
visible there also. Its color pattern differs strikingly from
that seen in V. brunneolinea.
Vexillum elliscrossi new species
(figures 6-9)
Pusia daedala "Reeve," Kosuge, 1979, p. 27, pi. 6. fig. 28, not
Mitra daedala Reeve, 1845.
Pusia sp., Salisbury, 1981, p. 6, figs. 8.\-B (in color).
as fig. 7, length 12.8 mm. 9. Paratype, length 17.2 mm, Salisbury collection Figure 10. Vexillum xcnium Pilsbry, 1921, holotype,
length 17.8 mm, ANSP 116983. Note: All figures except 3, 5 and 7 are printed at the same magnification.
Page 150
THE NAUTILUS, Vol. 105, No. 4
Table 2. Measurements (mmj and counts of conchological
features of Vexillum elliscrossi n. sp. Number of specimens (N)
varies because in some cases values for juvenile, damaged and
worn specimens could not be determined.
Mean
Range
N
Shell length
17.4
16.9-18.5
Body whorl length
10.7
9.8-12.0
4
Bod\' whorl w idth
6.3
5.8-6.8
4
Spire length
6.3
6.2-6.4
3
Aperture length
8.6
7.7-9.5
4
Aperture width
2.5
2.5-2.7
4
.\pertural lirations
9.3
6-13
6
Plica 1 width
0.38
0.30-0.46
4
Teleoconch whorls
7.5
7.0-7.8
3
Ribs, whorl 1
13.7
12-15
3
Ribs, whorl 2
14.6
12-16
5
Ribs, whorl 3
15.0
12-16
5
Ribs, whorl 4
16.2
12-19
5
Ribs, whorl 5
17.4
14-22
5
Ribs, whorl 6
18.4
16-23
5
Ribs, whorl 7
20.0
18-23
3
Protoconch width
0.62
0 60-0.64
3
Protoconch whorls
44
44-45
2
Description: Shell length of adult specimens ranges from
16.9 to 18.5 mm; bod\' whorl width and spire height both
range from 35 to 38 percent of shell length. A summary
of measurements is given in table 2. The shell is white
with four brown bands on the body whorl, two of which
are visible on the spire whorls. The band below the suture
is sometimes faint and is lightest near the suture; the
band below it is the darkest of the four. Fresh specimens
are suffused with lavender, which fades rapidly after
collecting. The axial ribs are white and cut the brown
bands into a series of tightly spaced blotches. The ap-
erture and columella are white. All specimens studied
have large, diagonal scars where breaks in the shell were
repaired.
The protoconch is amber in color and appears smooth
under a light microscope. It is tall (0.90-0.95 mm), con-
sisting of about 4.4 whorls. There are up to eight teleo-
conch whorls. These are convex and slightly terraced.
The number of ribs on the early teleoconch whorls ranges
from 12 to 16, and increases to about 20 on the body
whorl. Spiral grooves are visible between the ribs. These
are faint on the first three teleoconch whorls and increase
in strength towards the body whorl. From 7 to 9 grooves
are visible on the spire whorls and 24 to 28 are visible
on the body whorl. The grooves on the body whorl are
stronger and more widely spaced along the siphonal ca-
nal. There is no siphonal notch and consequently there
is no fasciolar ridge or distinct siphonal fasciole. There
are four plications on the columella; the strongest is the
posteriormost, which is up to 0.5 mm wide. There are 6
to 13 apertural lirations.
Type material: HOLOTYPE BPBM 219991, 18.5 x
6.8 mm. PARATYPES. IMT, 3 uncatalogued specimens;
Salisbur) Collection, 1 specimen; Leonard Hill Collec-
tion Cs-173, 1 specimen, figured by Salisbury (1981).
Type locality: Off VVaikiki, Oahu, Hawaii, at 180 meters
on a sand bottom, collected bv the "Pele Expedition" on
21 March 1965.
Distribution: The three paratypes in IMT were cited
bv Kosuge (1979), from stations 73-3 (29°47.0'N;
179°04.7'E), 73-4 (29°47.4'N; 179°02.9'E), and 73-15
(26°18.9'N; 174''30.9'W) on central Pacific guyots west
of Midway Island. We did not examine a fourth speci-
men, figured by Kosuge (pi. 6, fig. 28), from station 73-
25 (29°48.0'N; 179°01.3'E) as it was not included in the
material loaned to us. The IMT specimens were dredged
in March 1973 at depths from 67 to 267 m. The paratype
in the Salisbury collection was dredged off Keehi Lagoon,
Oahu by E. R. Cross. The paratype in the Hill collection
was taken off Honolulu at about 90 meters in rock and
coralline algal rubble.
Etymology: We are pleased to name this species in
honor of Ellis R. Cross, editor emeritus of the Hawaiian
Shell News, who collected one of the paratypes. His work
with deep-dredged shells has contributed greatly to
knowledge of the molluscan fauna of the Hawaiian Is-
lands.
Discussion: Kosuge (1979) identified Vexillum ellis-
crossi as Vexillum daedalum (Reeve, 1845), but it is
easily distinguished from that species. Vexillum daeda-
lum has a brown or ashy green shell with a single white
peripheral band, whereas V. elliscrossi has a white shell
with four brown bands. The aperture and columella are
brown in V. daedalum and white in V. elliscrossi. The
axial ribs on the body whorl in V. daedalum number 10
to 15 (Cernohorsky, 1972), whereas V. elliscrossi has 18
to 23 ribs. V. daedalum has a siphonal notch and fasciole
whereas V. elliscrossi lacks the notch and fasciole. The
shell that Kay (1979) illustrated from Hawaii as Vexillum
sp. cf. rufofilosum (E. A. Smith, 1876) may be a deep
water form of Vexillum daedalum. Cernohorsky (1972)
and Turner (1989) considered V. rufofilosum to be syn-
onymous with V. daedalum. Turner illustrated the ho-
lotypeof V. rufofilosum. Vexillum oiiiscinum (Lamarck,
1811) might prove to be an older name for V. daedalum.
Another species that occurs in Hawaii that might be
confused with Vexillum elliscrossi is V. xenium Pilsbry,
1921 (figure 10). The holotype of V. elliscrossi was orig-
inally catalogued at the Bishop Museum as "V. (C.) xeni-
um?" but comparison to the holot\pe and paratype of
V. xenium (ANSP 116983) shows that the species are
easily distinguished. The apertural lirations in V. xenium
are interrupted; in V. elliscrossi they are continuous.
Vexillum xenium has a siphonal notch and fasciole; these
are lacking in V. elliscrossi. The brown peripheral band
in V. xenium is strongest on top of the spiral cords and
lighter in the grooves. There are several narrow bands
of brown on top of individual cords elsewhere on the
whorl; these are darkest where the\ cross the ribs and
lighter or absent in the interspaces. In contrast, the brown
G. Rosenberg and R. Salisbury, 1991
Page 151
bands in V. elliscrossi do not cross the ribs, but cover the
spiral cords and grooves uniformly, Vexillum xcniiim
also has scattered brown subsutural blotches; these blotches
are absent in V. elliscrossi.
ACKNOWLEDGEMENTS
We thank Dieter Cosman for sharing his knowledge of
Vexillum brunneolinea with us. We regret that we could
not name the species for him, but there is already a
Vexillum cosmani Kay, 1979, and the International
Commission on Zoological Nomenclature discourages
giving species in the same genus or in allied genera names
honoring the same person [ICZN Recommendation D(I)6].
We thank Robert H. Cowie of the Bernice P. Bishop
Museum (BPBM), Leonard C. Hill (Miami, Florida), and
Sadao Kosuge of the Institute of Malacology, Tokyo (IMT)
for providing information and loaning specimens.
LITERATURE CITED
Adams, A. and L. Reeve. 1848-1850. Mollusca. In: Adams,
A. (ed.). The zoology of the voyage of H.M.S. Samarang;
under command of Captain Sir Edward Belcher, London.
P. 1-24, pis, 1-9 (1848), p. 25-87, pis. 10-24 (1850).
Cernohorskv, W. O. 1970. Systematics of the families Mitri-
dae &i Volutomitridae (Mollusca: Gastropoda), Bulletin of
the .\uckland Institute and Museum, no, 8 iv, 190 p.
Cernohorskv, W, O. 1972, Marinesheilsof the Pacific, volume
II, Pacific Publications, Sydney, 411 p., 68 pis.
Gmelin, J, F, 1791. Caroli a Linne Systema naturae per regna
tria naturae. Edition decima tertia. Lipsiae, 1(6), Vermes,
pp, 3021-3910.
Kay, E. A. 1979. Hawaiian marine shells. Reef and shore
fauna of Hawaii. Section 4: Mollusca, Bernice P, Bishop
Museum Special Publication 64(4), xviii, 653 p.
Kosuge, S. 1979, Report on the Mollusca on guyots from the
central Pacific collected by 2nd and 3rd cruises of R/V
Kaiyomaru in 1972 to 73 with descriptions of twelve new
species. Bulletin of the Institute of Malacology, Tokyo,
l(2):24-35, pis. 5-6.
Lamarck, J. B. P. A. de M. de. 1811. Sur la determination
des especes parmi les animaux sans vertebres, et particu-
lierement parmi les mollusques testaces. .\nnales du Mu-
seum d'Histoire Naturelle (Paris) 17:195-222,
MacDonald, J. D. 1860, Further observations on the meta-
morphosis of Gasteropoda, and the affinities of certain
genera, with an attempted natural distribution of the prin-
cipal families of the order Transactions of the Linnean
Society of London 23:69-80. 1 table,
Pilsbry, H, A, 1921, Marine mollusks of Hawaii, VIII-XIII,
Proceedings of the Academv of Natural Sciences of Phil-
adelphia 72:296-328, pi. 12^
Reeve, L. 1844-1845. Monograph of the genus Mitra. Con-
chologia Iconica 2, pis, 1-27 (1844), 28-39 (1845).
Roding, P. F, 1798, Museum Boltenianum sive catalogus ci-
meliorum e tribus regnis naturae quae collegerat Joa, Fried
Bolten. Pars Secunda Johan, Christi, Trappii, Hamburg,
viii, 199 p
Salisbury, R, 1981, Little known miters of Hawaii II. Ha-
waiian Shell News 29(2):6, I color plate.
Smith, E. A. 1876. A list of marine shells, chiefly from the
Solomon Islands, with descriptions of several new species.
Journal of the Linnean Society of London. Zoology. 12:
535-562, pi. 30.
Turner, H. 1989. llncommon and new mitriform gastropods
from the Indo-Pacific. Part 1. Swiss Federal Research In-
stitute for Forest, Snow and Landscape, Birmensdorf. 35
p., 8 color plates.
THE NAUTILUS 105(4):152-158, 1991
Page 152
Parasitic Mites (Acari: Unionicolidae) of Fresh-water
Mussels (Bivalvia: Unionidae) in the Duck and
Stones Rivers in Central Tennessee
Malcolm F. Vidrine
Louisiana State University
at Eunice
P.O Box 1129,
Eunice, LA 70535 USA
James L. Wilson
Tennessee State University
Tenth and Charlotte,
Nashville, TN 37203 USA
ABSTRACT
The Duck and Stones Rivers in central Tennessee were sampled
for fresh-water mussels in 1956 and 1962. Twenty-nine species
of Stones River mussels were examined for parasitic mite in-
festations Twenty mussel species were infested by a total of
14 species of mites (Unionicola). Of the 552 mussel specimens
examined, 321 (58.2%) were infested with one or more mite
species. In the Duck River, 20 species of mussels were examined
for parasitic mite infestations. Thirteen mussel species were
infested by a total of 11 species of mites (Unionicola). Of the
637 mussel specimens examined, 381 (59.8%) were infested
with one or more mite species. Mussel-mite associations are
listed. These data provide a unique insight into a native fauna
that may now be irrecoverably damaged, at least in the im-
pounded areas of Stones River.
Key words: Unionidae; mussels; Unionicolidae; Unionicola;
mites; Stones River; Duck River; Tennessee.
INTRODUCTION
Fresh-water mussels (Bivalvia: Unionidae) form a ma-
jor part of the macrobenthos in streams and lakes of
North America. Although these mussels are commonly
parasitized by aquatic mites (Acari: Unionicolidae), few
published studies of geographical co-occurrence of the
host and parasite species exist. Classical works, especially
those of Wolcott (1899) and Marshall (1933), provide a
few host records and no comprehensive mite species in-
cidence records for the stations sampled. In the first con-
cise work on the Tennessee mussel mites, Mitchell and
Wilson (1965) describe 5 new species of mites from 5
mussel species in the Duck River. Before the 1980s, the
taxonomy of unionicolid mites was very poorly under-
stood; recent works (Vidrine 1980, 1985a, 1986d, 1987,
1988), however, propose 53 new species and a revision
of the supraspecific systematics. Vidrine (1989), Vidrine
and Vidrine (1987), and Vidrine, Bouchon and Poirrier
(1986) discussed incidence of mussel-mite associations in
Arkansas, Louisiana, Mississippi, and Texas. Earlier at-
tempts to describe mussel-mite community structure bv
Dobson (1966), Calnan (1976) and \idrinc (1974) were
hampered by the lack of recognition and definitive de-
scriptions of many of the mite species encountered.
This paper describes the mussel-mite associations from
collections made by JLW (junior author) in the Stones
River (Cumberland River drainage) and the Duck River
(lower Tennessee River drainage) in central Tennessee,
during 1956 and 1962. These data provide a unique
insight into a native fauna of the Stones River (impound-
ment in 1968: J. Percy Priest Lake) and the Duck River
(impoundment in 1976: the Normandy Project).
MATERIALS AND METHODS
During the summers of 1956 and 1962, mussels were
collected from the following 8 stations in the Stones Riv-
er:
1 . Stones River at lower end of Charlton Ford off Couch-
ville Pike, Davidson Co., Tennessee. 25 August 1956
(now under backwater of J. Percy Priest Lake).
2. Stones River ca. 200 m above Couch\ille Pike Bridge,
Davidson Co., Tennessee. 18 June, 16 and 29 July
1962 (now under backwater of J. Percy Priest Lake).
3. Stones River ca. 100 m above bridge on Stewart's
Ferry Road, southeast of Donelson, Davidson Co.,
Tennessee. 14 August 1962 (now under backwater of
J. Percy Priest Lake).
4. Stones River ca. 1.0 km below "Old Jefferson ' Bridge
ca. 4.5 km east of Smyrna and 1.3 km north of Hwy.
6306, Rutherford Co., Tennessee. 17 .August 1962 (now
under backwater of J. Percy Priest Lake).
5. Stones River ca. 30 m below junction of East Fork
and West Fork Stones Rivers and 1.3 km upstream
from "Old Jefferson" bridge and ca. 4.5 km east of
Sm\rna and 1.3 km north of Hwy. 6306, Rutherford
Co., Tennessee. 20 August 1962 (now under back-
water of J. Perc\' Priest Lake).
6. East Fork Stones River below bridge below dam at
Walterhill on Hwy. 231, ca. 0.8 km north of Mur-
freesboro, Rutherford Co., Tennessee. 20 August 1962
M, F. Vidrine and J. L. Wilson, 1991
Page 153
(above impoundment and free flowing; near station
E3 of Schmidt (1982)).
7. East Fork Stones River below bridge on Hw). 6189,
ca. 4.5 km northeast of Readyville (on Hwy. 705),
Rutherford Co., Tennessee. 23 August 1962 (above
impoundment and free flowing; near station E14 of
Schmidt (1982)).
8. East Fork Stones River ca. 30 m below mill dam (still
operating) and ca. 1.3 km east of Lascassas and ca.
3.0 km south of Rte. Tenn. 96, Rutherford Co., Ten-
nessee. 23 August 1962 (above impoundment and free
flowing; near station E6 of Schmidt (1982)).
Mussels were also collected from the following 10 sta-
tions in the Duck River:
1. Duck River below bridge on Hwy. 41 north of Man-
chester, Coffee Co., Tennessee. 31 July 1962 (above
backwater of Normandy Reservoir and free flowing:
J. Jenkinson, personal communication, 1990).
2. Duck River below bridge on Hw\'. 231 in Shelby ville,
Bedford Co., Tennessee. 26 July 1962 (free flowing,
ca. 45 km below the dam of Normandy Reservoir,
and minimally impacted by cold water discharge
from the impoundment: J. Jenkinson, personal com-
munication, 1990).
3. Duck River below bridge on Hw\. 31a south of
Chapel Hill, Marshall Co., Tennessee. 5 July 1962
and 26 July 1962 (free flowing: J. Jenkinson, personal
communication, 1990).
4. Duck River ca. 200 m below dam (Lillard Mill — an
old hydroelectric dam) and 0.7 km N of Hwy. 4277
at Milltown, Marshall Co., Tennessee. 25 June and
July 1962 and 26 August 1956 (free flowing and with
a large mussel community: J. Jenkinson, personal
communication, 1990).
5. Duck River ca. 100 m below old mill dam below
bridge on Hwy. 431, Maury-Marshall Co. line, Ten-
nessee. 27 July and 29 August 1962 (free flowing and
with a large mussel community, old dam now washed
out: J. Jenkinson, personal communication, 1990).
6. Duck River above bridge on Rte. Tenn. 7 at north-
western city limits of Columbia, Maury Co., Ten-
nessee. 2 and 29 August 1962 (free flowing with a
large mussel community: J. Jenkinson, personal com-
munication, 1990).
7. Duck River below bridge at Williamsport on Rte.
Tenn. 50, Maury Co., Tennessee. 2 .\ugust 1962 (free
flowing but with few or no mussels: J. Jenkinson,
personal communication, 1990).
8. Duck River below Wm. Erwin McEwen Bridge at
Rte. Tenn. 50, Maury-Hickman Co. line, Tennessee.
23 July 1962 (free flowing but with few or no mussels:
J. Jenkinson, personal communication, 1990).
9. Duck River ca. 100 m downstream from bridge at
western city limits of Centerville on Rte. Tenn. 50,
Hickman Co., Tennessee. 23 July and 6 August 1962.
Obviously polluted by sewage in 1962 (still free flow-
ing and polluted by sewage and industrial wastes.
with few or no mussels: J. Jenkinson, personal com-
munication, 1990).
10. Duck River below bridge on Rte. Tenn. 50, ca. 5 km
west of Centerville, Hickman Co., Tennessee. 6 Au-
gust 1962 (still free flowing but with few or no mus-
sels: J. Jenkinson, personal communication, 1990).
.\ total of 557 mussels from the Stones River, and 637
mussels from the Duck Ri\ er was examined by dissection,
for adult and n\ mphal mites by JLW. All but 5 of these
mussels were identified by Henry van der Schalie and
Ralph Sinclair. The unidentified mussels (all from Stones
River) are excluded from consideration here. The mites
of each individual mussel were stored in separate, la-
belled vials of Koenike's fluid. The mites were identified
by MF\' in 1987. Mussel names have been updated to
conform with the concepts of Burch (1975) and Davis
and Fuller (1981). The mussels were deposited in the
collections of the Museum of Zoology at the University
of Michigan, Ann Arbor.
RESULTS
Fourteen species of Unionicola were found in 20 of
29 species of fresh-water mussels from the Stones River.
Of the 552 mussels examined, 321 (58.2%) were infested
with one or more species of Unionicola: 268 had one
species, 40 had two species, and 13 had three species.
Results of these examinations are summarized in table
1. The mussel diversity and abundance conform gener-
ally with those reported by Wilson and Clark (1914) and
Schmidt (1982).
Eleven species of Unionicola were found in 13 of 20
species of fresh-water mussels from the Duck River. Of
the 637 mussels examined, 381 (59.8%) were infested
with one or more species of Unionicola: 372 had one
species, 7 had two species, and 2 had three species. Results
of these examinations are summarized in table 2. The
mussel diversity and abundance conform generalK- with
those reported bv Isom and Yokelv (1968), H. van der
Schalie (1973), Ahlstedt (1986), and Jenkinson (1988).
Mussel-mite associations can be classified as resident,
transient, or vagrant (Vidrine 1989), as adapted from
Treat's (1975) definitions for Lepidoptera-mite associa-
tions. All 15 species of Unionicola reported here are
naturally resident mites in the encountered mussel hosts,
as inferred from associations encountered in other studies
(Vidrine 1985b, 1986a,b,c, 1987, 1989). The few vagrant
associations encountered here are naturally incidental
and/or inevitable artifacts of sample handling.
The following associations were encountered in the
Stones River (site of natural infestation within branchial
chamber of hosts is provided within parentheses) (% in-
fection of a specific mussel species by a specific mite
species is provided in parentheses):
1. Unionicola dimocki Vidrine 1986b (gills) (83.1%)
and U. smithae Vidrine 1986a (mantle and foot)
(6.8%) were found in Lasmigona costata (Rafinesque
Page 154
THE NAUTILUS, Vol. 105, No. 4
Table
Sumniarv of miisscltiiitf associations in the Stones River, Tennessee, based upon collections in 19.56 and 1962
Mussels
Mites
Number of mussels examined (in bold)
Number of mussels infested by parasitic mites
Stations
Total
Anodonta grandis Say 1829
I'nionicold smilhai' X'idrine 1986a
Lasmigona costala (Rafinesque 1820)
U. dimocki X'idrine 1986b
V. smithac X'idrine 1986a
Amblema plicata Say 1817
U. amandita Mitchell and XVilson 1965
U. tupara Mitchell and XVilson 1965
U. serrata (XX'olcott 1898)
Amblemn gignntea (Barnes 1823)
('. tupara Mitchell and XX'ilson 1965
Fusconnin flnva (Rafinesque 1820)
['. parheri X itirine 1987
Elliptio dilntata Rafinesque 1820
V. scrrala (Wolcott 1898)
U. tupara Mitchell and XVilson 1965
E. crassidens (Lamarck 1819)
Cyclonaias tuberculata (Rafinesque 1820)
U. sakantaha Mitchell and Wilson 1965
U. abnormipes (X\'olcott 1898)
U. causeyae X'idrine 1985b
U. serrata (XVolcott 1898)
U. vamana Mitchell and XVilson 1965
Quadrula pustulosa (Lea 18.31)
['. causeyae X'idrine 1985b
Orthonymus cylindrica (Say 1817)
I', sakantaka Mitchell and XX'ilson 1965
Tritogonia verrucosa (Rafinesque 1820)
I', vamana Mitchell and XX'ilson 1965
Actinonaias carinata (Barnes 1823)
U. hoesei X'idrine 1986c
v. abnornnpes (XVolcott 1898)
U. serrata (Wolcott 1898)
A. pectorosa (Conrad 1834)
U. hoesei X'idrine 1986c
U. serrata (Wolcott 1898)
U. abnormipes (Wolcott 1898)
Lampsilis orbiculata (Hildrelh 1928)
['. hoesei X'idrine 1986c
r. abnormipes (XVolcott 1898)
L. ovata (Say 1817)
U. hoesei X'idrine 1986c
v. almormipes (XVolcott 1898)
U. serrata (Wolcott 1898)
L. fnsrioln Rafinesque 1820
Ohnvnrin subrotiinda (Rafinesque 1820)
Truncitia trunratn Rafinesque 1820
T. donatifurntis (Lea 1828)
Plagioln lenior (Lea 1840)
/'. interruptii (Rafinesqu<- 1820)
I'otaniiliis purpitralus (l.aniurek 1819)
P. alatus (Say 1817)
(/. hoesei X'idrine 1986c
U.fulleri Vidrine 1986c
U. australindistincta Vidrine 1985b
V. serrata (Wolcott 1898)
II
8
26
2
17
12
13
5
3
3
1
17
10
2
2
4
3
1
1
3
1
37
2
2
35
18
2
6
5
1
19
1
1
1
1
6
6
13
10
5
2
15
10
4
4
1
14
14
4 —
10
8
1
18
15
6
— — 1
15 — —
1
2
3
34
13
U
1
4 —
4 —
1 —
1
1
55
51
i
22
15
2 — —
118
98
8
151
15
53
11
23
17
1
1
35
1
2
2
26
24
2
6
2
1
22
1
1
1
7
6
16
12
5
3
33
23
8
4
4
1
2
26
22
6
3
4
1
1
1
1
4
1
16
11
4
5
1
M. F. Vidrine and J. L. Wilson, 1991
Page 155
Table
Continued
Mussels
Mitfs
Number of mussels exaniiiied (in bold)
Number ot mussels infested b\' parasitic mites
Stations
["otal
Ligumia recta (Lamarck 1819)
U. serrata (VVolcott 1898)
[/. ahnormipes (Wolcott 1898)
Villosa taeniata (Conrad 1834)
('. hoesei Vidrine 1986c
Medionidus conradicus (Lea 1834)
['. hciulrixi N'idrine 1987
Leptoden fragilis (Rafinesque 1820)
['. hoesei Vidrine 1986c
Ptychobranchus fasciolare (Rafinesque 1820)
U. hoesei Vidrine 1986c
C. can.seycw Vidrine 1985b
Obliquaria reflexa Rafinesque 1820
Totals ol' mussels
2
2
3
3
11
1
146
1 —
1 —
9
3
2 —
J9
()2
86
6
1
1
12
2
3
3
2
2
33
8
1
1
1820); a single vagrant specimen of V . smithae was
found in Anodonta grandis Sa> 1829.
2. Unionicola amandita Mitchell and Wilson 1965 (gills)
was found only in Amblema plicata Say 1817 (10.0%).
3. Unionicola tupara Mitchell and Wilson 1965 (gills)
was found in A. plicata (35.1%) and Amblema gi-
gantea (Barnes 1823) (73.9%).
4. Unionicola sakantaka Mitchell and Wilson 1965
(gills) was found in Cyclonaias tubercidata (Rafin-
esque 1820) (92.3%) and Orthonynms cylindrica (Say
1817) (100.0%).
5. Unionicola vamana Mitchell and Wilson 1965 (gills)
was found in Tritogonia verrucosa (Rafinesque 1820)
(85.7%); a vagrant specimen was found in C. tuber-
cidata.
6. Unionicola hoesei Vidrine 1986c (gills), a common
parasite of 15 genera in the Lampsilini (Unionidae:
Ambleminae) in North America (Vidrine 1986c), was
found in Lampsilis ovata (Say 1817) (84.6%), Lamp-
silis orbiculata (Hildreth 1828) (25.0%), Villosa
taeniata (Conrad 1834) (41.7%), Actinonaias cari-
nata (Barnes 1823) (75.0%), Actinonaias pectorosa
(Conrad 1834) (69.7%), Leptodea fragitis (Rafin-
esque 1820) (100.0%), Ptychobranchus fasciolare
(Rafinesque 1820) (24.2%) and Potamilus alatus (Say
1817) (68.8%) (all lampsilinines) in the Stones River.
7. Unionicola fidleri Vidrine 1986c (gills) (25.0%) and
Unionicola australindistincta Vidrine 1985b (man-
tle and foot) (31.3%) were found only in P. alatus.
8. Unionicola hendrixi Vidrine 1987 (gills) was found
only in Medionidus conradicus (Lea 1834) (100.0%).
9. Unionicola parkeri N'idrine 1987 (gills) was found
only in Fusconaia flava (Rafinesque 1820) (100.0%).
10. Unionicola causeijae Vidrine 1985b (mantle and foot)
was found in C. tubercidata (23.1%), P. fasciolare
(3.0%), and Quadrula pustulosa (Lea 1831) (4.5%).
11. Unionicola ahnormipes (Wolcott 1898) (mantle and
foot) was found in A. carinata (31.3%). A. pectorosa
(12.1%), L. ovata (23.1%), L. orbiculata (50.0%) and
Liqumia recta (Lamarck 1819) (16.7%); a vagrant
specimen was found in C. tubercidata.
12. Unionicola serrata (Wolcott 1898) (labial palps) was
found in A. carinata (18.8%), A. pectorosa (24.2%),
A. plicata (7.3%), C. tuberculata {1.1%), Elliptio di-
latata Rafinesque 1820 (2.9%), L. ovata (11.5%), L.
recta (16.7%), and P. alatus (6.3%).
The following associations were encountered in the
Duck River (site of natural infestation within branchial
chamber of hosts is provided w ithin parentheses) (% in-
fection of a specific mussel species by a specific mite
species is provided in parentheses):
1. Unionicola dimocki (gills) (89.5%) and U. smithae
(mantle and foot) (12.3%) were found only in L. cos-
tata.
2. Unionicola amandita (gills) was found only in A.
plicata (48.7%).
3. Unionicola tupara (gills) was found in A. plicata (0.3%)
and A. gigantea (84.2%).
4. Unionicola sakantaka (gills) was found in C. tuber-
culata (9O.7%0 and O. cylindrica (100.0%); a vagrant
specimen was found in T. verrucosa.
5. Unionicola vamana (gills) was found in T. verrucosa
(83.0%); vagrant specimens were found in C. tuber-
cidata (1) and Q. quadrula (Rafinesque 1820) (5).
6. Unionicola vikitra Mitchell and Wilson 1965 (gills)
was found in Q. quadrula (47.6%) and C^. pustulosa
(100.0%); a vagrant specimen was found in C. tuber-
culata.
7. Unionicola hoesei (gills) was found in Lampsilis fas-
ciola Rafinesque 1820 (50.0%), Villosa fabalis (Lea
1831) (81.3%), A. pectorosa (62.5%), L.fragilis (8.3%),
and P. alatus (30.0%) (all lampsilinines).
Page 156
THE NAUTILUS, Vol. 105, No. 4
Table 2. Siiminar> of mussel-mite associations in the Duck Ri\er, Teiuiessee, based upon collections in 19.56 and !9fi2
Number of niu§§eU examined (in biild)
Number of mussels infested h\ parasitic mites
Stations
Mussels
Mites 12 3 4 .5 fi 7 8 9 10 Total
Anodonia grandis Say 1829 — — — — — I — — — — 1
Alasniidonta marginatn Say I8I*J — — — — 2 — — — — — 2
Lasrnigona complanntn {Barne> 1823) — — — — — o — — — — a
L. cosrafa (Rafinesque 1820) — II 18 14 14 _____ 57
Uniotncola dimochi Mdrine l9H6h — 8 18 11 14 _____ 51
['. smithae \idrine 1986a — 14 1 ]_____ 7
Strophitus undiilatus (Say 1817) — — — — — — — — 3 — 3
.4m6/pma/»/ic«fn Say 18i7 — 3 29 170 74 42 — — — — 318
r. «"m;i(/i7« Mitcliell and WiLson 1965 — 2 17 90 45 i _ _ _ _ 155
C. (K/wro Mitchell and Wilson 1965 _____ i____ i
.4. gigantea (Barnes 1823) _ _ _ 5 10 4 — — — — 19
C;. (i/paru Mitchell and Wilson 1965 _ _ _ 4 1(1 2 — — — — 16
Cyc/onaias fubercu/afa (Rafinesque 1820) — 1 28 13 9 I 2 — — — 54
U. sakanlaka Mitchell and Wilson 1965 _ 1 27 12 9 — — — — — 49
U. vikitra Mitchell and Wilson 1965 __!_______ 1
U. varnana Mitchell and Wilson 1965 — — — — — 1 — — — — 1
Tritogonia verrucosa (Rafinesque 1820) — — 19 20 6 5 — — 3 — 53
v. varnana Mitchell and Wilson 1965 — — 19 17 6 2 — — — — 44
U. sakantaka Mitchell and Wilson 1965 ___ ]______ i
Orthonymus cylindrica (Say 1817) — — 2 — — — — — — — 2
U. ,sa/ca;i(a/l-a Mitchell and Wilson 1965 __2 — — — — — — — 2
Quadrula quadrtila (Rafinesque 1820) — — — — — 60 — 1 2 — 63
C. (iAi7ra Mitchell and Wilson 1965 _ _ _ _ _ 30 _ _ _ _ 30
['. varnana Mitchell luul Wilson 1965 — — — — — 5 — — — — 5
Q. pustulosa (Lea 18SI) ___ 1_ i____ 2
U. vikitra Mitchell and Wilson 19(i5 ___ i_ i____ 0
Elliptio crassidens (Lamarck 1819) — — — — — 1 — — 2 — 3
Lampsilis ovatn (Say 1817) — — — — — — — — 1 — 1
L. teres (Rafinesque 1820) _____ 2 — — — — 2
L. fasciola Rafinesque 1820 6 — — — — — — — — - — 6
I', huesei V'idrine 1986c 3 — — — — — — — — — 3
ViUosafabalis (Lea lii3\) 16 — — — — _____ 16
r. /ioesei \idrine 1986c 13 — — — — _____ 13
Actinonaias pectorosa (Conrad 1834) 5 — 1 1 1 — — — — — 8
U. hoesei V'idrine 19S6c 4 — 1 — — — — — — — 5
Lepfodea/ragi/is (Rafinesque 1820) ____ 1 2 — — 9— 12
V. hoesei V'idrine 1986c ____ i_____ 1
Potnmi7us a/afus (Say 1817) ___ 3 i___5110
C. /irw.vn \idrine 1986c — — — 3 ______ 3
U./ii/Zfri \idrine 1986c ___ 2 i_____ 3
V. australindistincta V'idrine 1985b — — — 1 — — — — — — 1
U. serrata (Wolcott 1898) — — — ]______ 1
Totals of mussels 27 15 97 227 118 124 2 1 25 1 637
8. Unionicola fulleri (gills) (30.0%), U. serrata (labial
palps) (10.0%), and U. australindistincta (mantle and
foot) (10.0%) were found only in P. alatus.
The following mussels were not found parasitized by
mites in either or both of the rivers studied: Alasniidonta
marginata Say 1819, Lasmigona complanata (Barnes
1823), Strophitus undulatus (Say 1817), Elliptio cras-
sidens (Lamarck 1819), Lampsilis teres (Rafinesque
1820), Obovaria subrotunda (Rafinesque 1820), Trun-
cilla truncata Rafinesque 1820, Truncilla donaciformis
(Lea 1828), Plagiola lenior (Lea 1840), Plagiola inter-
rupta (Rafinesque 1820), Potamilus purpuratus (La-
marck 1819), and Obliquaria reflexa Rafinesque 1820.
DISCUSSION
Intra-host regulation of a mite-parasite load, perhaps
through rivalry of sexual competitors, as described by
Mitchell (1965), was apparent in several mite species
encountered here. In the Stones River, Unionicola aman-
M. F. Vidrine and J. L. Wilson. 1991
Page 157
dita, U. dimocki, U. hoesei, U. sakantaka. U. vamana,
V. hendrixi and V. parkeri t>picall\- occurred in small,
characteristicaih discrete numbers within each host: a
single male, and one or two females usually infested the
gills of their mussel host. One to four individuals of V.
tupara and L'. fulleri (all males, all females, or combi-
nations of both sexes) infested the gills of their hosts. One
or two females of U. smithae were usually found on the
mantle or foot of infested hosts, while males were absent,
probabK due to seasonal reasons. One to four specimens
of the gill mites V. tupara and I', fulleri were found in
each host. One or two indi\ iduals of the less host-specific
labial palp parasite, U. serrata, were found in 7 genera
of mussels. One to five individuals of U. abnormipes, U.
causeijae and U. australindistincta, which infest the
mantle and foot of their hosts, were found in a variety
of amblemine hosts. The large number of individuals
(range 1-65, but most often l)oi U. abnormipes infesting
individual hosts may represent an absence of the strict
territorialit) apparent in the above associations.
In the Duck River. Unionicola amandita, U. dimocki,
V. hoesei, U. sakantaka. U. vamana and V. vikitra typ-
ically occurred in small numbers within each host: a
single male, and one or two females infesting the gills
of their individual hosts. One to nineteen. usualK 5.
individuals of U. tupara and V. fulleri infested the gills
of their hosts. One to five females of U. smithae were
found on the mantle or foot in each infested host, while
males were not found. Onl\ a single male of U. serrata,
a labial palp mite, was found. A single Potamilus alatus
had one male and two females of V. australindistincta,
a mantle and foot parasite.
In both the Stones River and the Duck River. V . aus-
tralindistincta was found in P. alatus. These are new-
host and localit\' records for L'. australindistincta, which
has been previously known mainly from other species of
the genus Potamilus and Leptodea fragilis in .\rkansas,
Louisiana, Mississippi, and Te.xas (Mdrine 1986b). The
closely related mite species, L'. indistincta (Wolcott 1898).
is the usual parasite of P. alatus, both in the upper Mis-
sissippi and upper Tennessee river drainages (Clinch and
Powell Rivers) (N'idrine, 1980; Wolcott. 1899). The ad-
dition of P. alatus to the host list for L'. australindistincta
neither confirms nor refutes Mdrine's (1985b) contention
that U. australindistincta is a southern form of U. in-
distincta. In this sampling. onl\ a single, unparasitized
P. purpuratus (a common host of U. australindistincta
south of Tennessee) was found in the Stones River.
The data in tables 1 and 2 illustrate the incidence of
specific mite taxa in specific mussel groups (subfamilies
Anodontinae and Ambleminae). The anodontine mussels,
for example, are infested by distinctly different mantle
and gill mites than are amblemine mussels. This phe-
nomenon was noted by Vidrine and Bereza (1978) and
reinforced by Vidrine (1980, 1986a,b,c,d, 1987. 1989).
These data suggest that Unionicola tend to be found
in the more abundant mussel species in a habitat. .^
critical threshold of host population size may be an oblig-
atory prerequisite for a population of a specific, extant
mite species to be sustained locally. Small and unpar-
asitized populations of certain mussel species may be
naturally below a threshold assuring sur\ival of mites
capable of parasitizing them, or artificially so as a result
of human impact, .although found parasitized elsewhere,
the following species were locally Unionicola-iree in this
study: E. crassidens was found parasitized in neither the
Duck or Stones Rivers; L. fasciola, O. subrotunda, T.
truncata, T. donaciformis, P. purpuratus, and O. refiexa
were not found parasitized in the Stones River; A. gran-
dis, A. marginata. L. complanata, S. undulatus, L. ova-
ta, and L. teres were not found parasitized in the Duck
River. Whether or not this local phenomenon is due to
naturalK small or to damaged populations of these mus-
sels, is unknown.
Some mussel species ma\- have never achieved critical
population sizes that could realize the very evolution of
a mite into a parasitic relationship with them. The t%p-
ically small populations of mussels in the genus Plagiola
may exemplify this phenomenon, and may be culpable
in hastening the extinction of several species of Plagiola
in the last centur>-, little doubt as a result of human
impact. Plagiola were never found parasitized by Unio-
nicola in the field or in museum holdings thus far bv
MFV, and no published records are extant.
The data derived from these collections are historicalK
important because all mussels that were found were col-
lected, providing a rough estimate of the relative species
abundance and incidence in these damaged and/or im-
pacted faunas. Also, these data are enriched by a mite-
mussel parasitization profile, providing another dimen-
sion to the systematics and natural history of these faunas,
and an insight into a diverse and differentially specialized
host-parasite complex. The Cumberlandian mussel fauna
and much of the lower Tennessee River fauna, now large-
ly decimated down to their few remaining refugia, his-
torically represented the most diverse svmpatric fresh-
water mussel fauna known on Earth. Insights into the
mite-mussel coevolutionar\' strategies and into the deli-
cate population control mechanisms that are necessar\
to maintain such a diverse community are now largely
unattainable from these mussel communities, as the nec-
essary sampling of mussels is untenable in the few viable
populations, or impossible in those extirpated by im-
poundment. Recent mussel studies (Ahlstedt 1986; Jen-
kinson 1988; Schmidt 1982) do indicate that diverse com-
munities still exist in portions of both the Duck and the
Stones Rivers. The presented baseline data sets will, we
hope, provide some corroborative elements to future
studies of the now-relict Cumberlandian and other mus-
sel faunas and their parasites and perhaps engender in-
terest from ecologists who seek to explain the intricacies
that drive the complex biotic relationships within benthic
invertebrate communities.
ACKNOWLEDGEMENTS
We sincereK thank Daniel J. Bereza for his assistance
in the preparation of this paper. John J. Jenkinson, Ten-
nessee X'alley Authority, provided valuable information
on the present conditions of the rivers.
Page 158
THE NAUTILUS, Vol. 105, No. 4
LITERATURE CITED
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Burch, J. B. 1975. Freshwater Unionacean clams (Mollusca:
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THE NAUTILUS 105(4):159-164. 1991
Volutes from Sava de Malha Bank:
The Saga of Lijria siirinamensis and a New Species
Page 159
Philippe Bouchet
Museum national d'Histoire
naturelle
55 rue Buffon
75005 Paris, France
Patrice Bail
2 square LaFontaine
75016 Paris, France
ABSTRACT
The supposedly fulgorariine volute Dallivoluta surinamensis
Okutani, 1982 is shown to have been described from an erro-
neous type locality in the Caribbean. It actually originates from
Saya de Malha Bank, in the Indian Ocean, where it is probably
endemic. Dallivoluta Okutani, 1982 is regarded as a synonym
of Lyria Gray, 1847. It is further suggested that Murex suri-
namensis Okutani, 1982 was also erroneousK- localized and
actually originates from the same submarine Indian Ocean
bank. Lyria doutei n.sp. is described from Saya de Malha, and
compared with other species in the group of L. lyraeformis.
Endemism on Saya de Malha is discussed in connection with
the history and isolation of the bank.
Key words: Indian Ocean; Mascarene Plateau; Caribbean; Vol-
utidae; endemism.
INTRODUCTION
In the last few months, collectors in Western Europe
have obtained from Russian sources specimens of two
remarkable volute species trawled by Soviet research
vessels on Saya de Malha Bank, in the Indian Ocean.
Nothing similar had been described from the Indo-Pa-
cific, and the Sa\ a de Malha shells appeared to represent
two new species of Lyria. when we noticed the identitv'
of one of them with Dallivoluta surinamensis Okutani,
1982. We present below the reasons why we believe that
D. surinamensis was erroneously localized in the original
description, we synonymize Dallivoluta with Lyria, and
we describe the second Lyria as a new species.
The species Dallivoluta surinamensis Okutani, 1982:
The volute Dallivoluta surinamensis was described as a
new genus and species, based on a single dead shell
(figure 1 ) and fragments collected by Japan Marine Fish-
ery Resources Research Center (JAMARC), supposedly
off the coast of Surinam in December 1978 (Okutani,
1982:115). The type locality was given as 11°03'N,
61°01'W, depth 128 m. Okutani (in Takeda & Okutani,
1983:290) provided an abridged description and color
photographs of the holotype. No further original infor-
mation on Dallivoluta surinamensis has been published
since the original description.
Okutani (1982) compared Dallivoluta with Tractolira
Dall, 1869 and especially Fulgoraria Schumacher, 1817,
which is probably the reason why Poppe (1987:14) placed
Dallivoluta in the subfamily Fuigorariinae.
The type locality: The t\ pe locality indicated by Oku-
tani (1982) is not situated off Surinam, as the specific
name would imply. The geographical coordinates indi-
cate a position 700 km to the northwest of Surinam,
between the islands of Trinidad and Tobago, off Vene-
zuela. The depth (128 m) is in the range of depths found
around these islands. The type locality of Murex suri-
namensis Okutani, 1982, described in the same paper,
is 10°46'N, 61°32'W, 94 m. These coordinates indicate a
position on the island of Trinidad, not in the seas around
it.
In fact, neither of these localities falls within the area
encompassed by the JAMARC study area off Surinam
and French Guiana (Takeda & Okutani, 1983:9, 11), and
there are grounds to suspect that they are erroneous.
Yokes (1990a:4, footnote; 1990b:6) had expressed doubts
on the type locaHty of Murex surinamensis, and sug-
gested that "opportunity for mi.xed locality data is cer-
tainly present" (Vokes, 1990a).
Identity of Dallivoluta surinamensis with the Saya de
IVIaIha volute: The volutes obtained from Soviet sources
belong to two different species of Lyria. Of one of these,
we have seen two complete shells, empty but very fresh
and probably live taken, said to have been collected by
Soviet research vessels on Saya de Malha: one is in the
Doute collection (Germany); the other is in the second
author s collection (figure 2). In addition, we have seen
photographs of a third specimen from another private
collection. Comparison of this material with Okutani's
(1982, 1983) description and figures (this paper figure 1)
leaves no doubt as to their identity. The spiral cords are
more distinct on the spire whorls of the newly collected
specimen, but this is undoubtedK due to the better pres-
ervation of this shell.
We therefore conclude that the type locality is erro-
neous. When the geographical coordinates of the type
locality are read with South instead of North, and East
instead of West {i.e.. 11°03'S, 61°01'E, instead of 1 1°03'N,
Page 160
THE NAUTILUS, Vol. 105, No. 4
Figure 1. Lyria surinamensis (Okulani, 1982). Holotype (from
Okutani, in Takeda & Okutani, 1983). Height 105 mm.
61°0rW), the position given is precisely correct for Saya
de Malha Bank, in the Indian Ocean! We also suggest
that the type locality of Murex surinamensis is similarly
inaccurate, and should be read 10°46'S, 61°32'E, also on
Saya de Malha Bank. This would confirm the observation
by Ponder and Yokes (1988;12) and Yokes (1990a:4,
1990b:6) that there are no species of Murex s.s. in the
New World.
It would thus appear that the Surinam material pro-
vided to Dr. Okutani by JAMARC had unfortunately
been contaminated by specimens trawled from the In-
dian Ocean.
Dallivoluta surinamensis and Murex surinamensis will
add to the list of species with unfortunate geographical
specific names, such as the Caribbean Cassis madagas-
cariensis Lamarck, 1822, originally believed to come
from Madagascar, or Punclurella agulhasae Clarke, 1961,
the type locality of which was erroneously believed to
be within the limits of Agulhas Basin (see Barnard, 1963:
296). A similar case of mistaken geographical coordinates
concerns the magellanic Marginella warrenii Marrat,
1876, described from 50°23'N, 64°04' W (Gulf of St. Law-
rence), an error for 50''23'S, 64°04'W (between the Falk-
lands and Cape Horn) (Bavay, 1913).
Dallivoluta, a synonym of Lyria: Okutani was struck
by the general resemblance of D surinamensis with the
NW Pacific genus Fulgoraria and regarded the non-
tilted protoconch, whitish columellar callus and color
pattern to differ from the character states observed in
Figure 2. Lyria surinamensis. Specimen collected on Sa\a de
Malha Bank by Soviet vessels (Bail collection). Height 87 mm.
Fulgoraria. However, we find D. surinamensis most
closely related to the E. African Lyria lyraeformis
(Swainson, 1821) and regard the generic characters enu-
merated by Okutani to be of specific importance only.
The shell characters of Dallivoluta surinamensis fall
within the range observed in the genus Lyria. as restrict-
ed by Bouchet (1990), and we regard Dallivoluta Oku-
tani, 1982 as a synonym of Lyria Gray, 1847.
Description of Lyria doutei: The second species of Lyr-
ia obtained from Soviet sources is known to us from a
single shell, originally in the Doute collection. At least
one other specimen was collected. It is in a private col-
lection in Europe or North America, and was available
for examination.
Lyria doutei n.sp.
(figures 4, 5)
Type material: Holot>pe in (Museum national d'His-
toirc iiaturclle (MNHN), leg. H. Doute.
Type locality: Saya de Malha Bank, 10°30'S, 62°25'E,
135 m.
p. Bouchet and P. Bail, 1991
Page 161
Description: Shell (figure 4) large, fusiform, solid, con-
sisting of 1.5 protoconch and 6 teleoconch whorls. Pro-
toconch (figure 5) bulbous, \\ith an indication of a cal-
carella on the initial part, remaining part with convex
smooth whorls. Protoconch/teleoconch transition sharp.
Teleoconch whorls convex, with appressed, but very
slightly channeled suture. Sculpture consisting of or-
thocline, gently sigmoid a.xial ribs; no spiral sculpture.
Fourteen axial ribs on first teleoconch whorl, gradually
increasing to 24 on body whorl. Last rib forming a thick
labial varix behind outer lip. ,\perture ovate, outer lip
smooth, inner lip with thin glaze over body whorl and
two groups of plaits; 8 plaits in columellar region, de-
creasing in strength from base to apex of shell; after a
smooth zone, 3 weaker plaits in parietal region. Siphonal
canal short, broad, open. Ground color creamy beige
with a more complex pattern of yellowish brown to brown
flamules and lines arranged in spiral bands. Spire whorls
with two darker bands, adapically and abapically, sep-
arated by lighter band on mid part of whorl. Adapical
band with 6 brown flamules usualh extending over one
rib and part of the interspaces adjacent to it; brown
flamules on abapical band with more indistinct contour
Medial light band with two interrupted brown spiral
lines. Body whorl with one light band, and one dark
basal band in addition to pattern described above. Ap-
erture creamy beige.
Dimensions: Height 83.5 mm, breadth 30.0 mm, ap-
erture 39 mm.
Remarks: Lyria doiitei is named after Mr. Harald Doute
(Bad Sackingen, Germany), a keen collector specializing
in volutes who first recognized it as a new species. Mr.
Doute generously provided the holotype, which he had
acquired from Mr. Igor Bondarev (Sebastopol, USSR).
Four Indian Ocean species have a general shell mor-
phology and/or color pattern that resemble that of L.
doutei. These are: L. lyraeformis, L. leslieboschae Em-
erson & Sage, 1986, L. cloveriana Weaver, 1963 and L.
surinamensis.
Lyria lyraeformis differs by having fewer axial ribs
on the body whorl: it has 16 ribs in southern populations
(Mombasa region, southern Kenya), and 20-23 ribs in
northern populations (northern Kenya and Somalia). The
color pattern of L. lyraeformis is extremely stable over
its entire range: there are three uninterrupted brown
bands on the body whorl, each lined with a dark brown
(sometimes interrupted) thread; these brown bands are
separated by two lighter spiral bands, each of them car-
rying 4-7 brown lines. Lyria lyraeformis also has a some-
what larger protoconch (height 4.9-5.4 mm vs 4.0 mm
in doutei), and when more specimens of L. doutei are
known it may appear that L. lyraeformis reaches a sig-
nificantly larger adult size: Weaver & DuPont (1970)
record specimens of 145 mm, but we have seen adults
of 180 mm.
Lyria leslieboschae is known only from the four shells
on which the original description was based (Emerson
& Sage, 1986). It is recognized by the distinct anal canal
Figure 3. Lyria surinamensis. Protoconch of specimen in fig-
ure 2 Scale line 2 mm.
formed by its outer lip. Its color is of the lyraeformis
type, but with a more diffused pattern. This species dif-
fers from both L. lyraeformis and L. doutei by having
dark axial flamules interrupting the lighter spiral bands.
Lyria cloveriana is more distantK related. It has an
even larger protoconch (height 6.4-7.1 mm); the axial
ribs become obsolete on the abapical part of the body
whorl; and the color pattern consists of evenly spaced
reddish brown spiral lines, without distinct darker spiral
bands as in the other species discussed abo\e.
Finall>, L. surinamensis differs from L. doutei by
having spiral cords in addition to the axial ribs. Lyria
surinamensis may also have fewer (19) ribs on body
whorl, and the color pattern consists only of poorly de-
fined orange-brown spiral bands on lighter beige back-
ground.
Mode of development and biogeographical remarks:
The bulbous protoconch of Lyria surinamensis (figure
3) and L. doutei (figure 5) are comparable in size with
those of L. lyraeformis and L. cloveriana Weaver, 1963.
As in all other modern volutes, it indicates non-planc-
totrophic development, and the large size further sug-
gests intracapsular metamorphosis and hatching as a
crawling juvenile (Bouchet & Poppe, 1988). This type of
development is incompatible with long distance passive
transport of larvae and/or juveniles and, considering the
isolation of Saya de Malha Bank (figure 6), suggests that
L. surinamensis and L. doutei are endemic to this bank.
Saya de Malha Bank is a large (ca. 4,000 km-) flat
plateau arising abruptly from the Indian Ocean sea-floor,
from depths in excess of 4,000 m. Its shallowest parts
reach as shallow as 10 m, with the large summital area
within the 200 m isobath. Together with the Seychelles
Page 162
THE NAUTILUS, Vol. 105, No. 4
Figure 4. Lijria doutei n.sp. Holotype MNHN. Height 83.5
Rank to the northwest and Nazareth and Cargados Ca-
rajos Banks to the south, it forms the Mascarene Plateau.
The Seychelles Bank is composed of late Precambrian
granitic rocks and is sometimes called the Seychelles
Microcontinent. On the other hand, Saya de Malha, Naz-
areth and Cargados C^arajos Banks have been formed by
the Reunion hotspot, Saya de Malha being the oldest in
the arc. It dates back to the Paleocene, while the car-
bonate rocks on Nazareth Bank are not older than Eocene
(Backman et ai, 1988). Saya de Malha and Nazareth are
separated by a sill shallower than 500 m and are only
200-250 km apart. Saya de Malha is separated from the
Figure 5. Lijria doutei n. sp. Protoconch of holotype. Scale
line 2 mm.
Seychelles Bank by depths greater than 2,000 m and a
distance of about 650 km. Distances to other Lyria-in-
habited continental shelves are: Saya de Vlalha-Mada-
gascar, 1,200 km; Saya de Malha-Kenya, 2,000 km; Saya
de Malha-Sri Lanka, 2,700 km.
For shelf-living volutes with large, crawl-away juve-
niles, this means that Saya de Malha and Nazareth Banks
are very isolated structures that are not easily colonized.
However, because they have never been connected to a
land mass since Lyria appeared in the Eocene, only the
dispersal of demersal lecithotrophic larvae can account
for colonization of such isolated oceanic banks. It is there-
fore remarkable that Lyria surinamensis has one of the
largest protoconchs in the genus, and that its concholog-
ical affinities appear to be with species geographically
remote: L. lyraeformis (Kenya), L. leslieboschae Em-
erson & Sage, 1986 (Arabian Peninsula), L. cloveriana
Weaver, 1963 (Sri Lanka). We hypothesize that the fol-
lowing scenario may have taken place: 1. Saya de Malha
is colonized by an ancestral Lyria species with smaller,
lecithotrophic, demersal larvae, having a broad Indian
Ocean distribution; 2. The species evolves intracapsular
metamorphosis with large crawl-away ju\eniles (this is
a general trend in volute evolution and has been noted,
e.g., by Hansen (1983) in American Paleogene volutes),
demersal dispersal is lost; 3. Geographical isolation en-
hances genetic isolation, and morphological divergence
accumulates through time into different species.
Lyria anna (Lesson, 1835) is another volute endemic
to this part of the Indian Ocean. The origin of this species
has long remained mysterious (Weaver & Dupont 1970),
but there are indications that it probabK originates from
Mauritius (Richards, 1990), nearb) areas NE of Mauritius
in 170-230 m (Anonymous, 1984) and/or Cinq Rrandons
p. Bouchet and P. Bail, 1991
Page 163
50«
I
SOMALI BASIN
io»
20'
60»
I
SEYCHELLES
\
SAYA DE MALHA
MASCARENE BASIN
CARGADOS CARAJOS
\r
Mauritius
Rodi'iguez
Reunion
Figure 6. Simplified bathymetric map of the southwestern Indian Ocean banks, showing the locaHties mentioned in the text
Stippled area, 0-100 m; thin intermediate line, 200 m isobath; thick line, 500 m isobath. Scale line 500 kilometers.
shoals on the Cargados Carajos Bank of the Mascarene
Plateau (unverified information from local Mauritius col-
lectors and dealers). Lijria anna is conchologically not
closely related to L. siirinamensis and indicates that the
colonization scenario above has occurred several times
in the family Volutidae. We predict that more species
of Volutidae will be found on banks of the Mascarene
Plateau, including the Seychelles Bank.
Finally, it is noteworthy that, beside Lyria siirina-
mensis and Murex surinamensis, Conus primus Rockel
& Korn, 1990 was very recently described from Saya de
Malha Bank, at depths of 80-98 m. It also has a paucis-
piral protoconch indicating non-planktotrophic larval
development.
LITERATURE CITED
Anonymous. 1984. Lyria (Harpeola) anna live dredged. La
Conchiglia 184/185:22.
Backman, J. et al. 1988. Introduction. Proceedings of the
Ocean Drilling Program, Initial Reports 115:5-15.
Barnard, K. H. 1963. Contributions to the knowledge of South
African marine Mollusca. Part IV. Ann. S. Afr. Mus. 47(2):
201-360.
Bavay, A, 1913. (no title). Journal of Conchologv 14(4):98-
99.
Bouchet, P. 1990. Systematics of Plicoliva with description
of a new subfamily (Gastropoda; Volutoidea). Archiv fiir
Molluskenkunde 120(1/3):1-10.
Bouchet, P. and G. Poppe. 1988. Deep water volutes from
the New Caledonian region, with a discussion on bioge-
ography. Venus 47(1): 15-32.
Hansen, T. 1983. Modes of larval development and rates of
speciation in early Tertiary Neogastropods. Science 204;
501-502.
Okutani, T. 1982. .\ new genus and five new species of gas-
tropods trawled from off Surinam. Venus 41(2):109-120.
Ponder, W. F. and E. H. Vokes. 1988. A revision of the Indo-
West Pacific fossil and Recent species of Murex s.s. and
Page 164
THE NAUTILUS, Vol. 105, No. 4
Haustelhtm (Mollusca: Gastropoda: Muricidae). Records
of the .Australian Museum, suppl. 8:1-160.
Poppe, G. 1987. New Volutes since 1970, part 1. La Con-
chiglia 218/219:8-22.
Richards, A. 1990. Mauritius, He de France. American Con-
chologist 18(4):8-10.
Rockel, D. and W. Korn. 1990. Conus species from the West-
ern Indian Ocean, dredged by Soviet biologists. Acta Con-
chyliorum 2:45-49.
Takeda, M. and T. Okutani. 1983. Crustaceans and moUusks
trawled off Suriname and French Guiana. JAMARC, To-
kyo. 356 p.
Vokes, E. H. 1990a. What ever happened to dear old Murex?
.American Conchologist 18(l):3-7.
Vokes, E H 1990b Cenozoic Muricidae of the Western At-
lantic region. Part VIII Murex s.s . Haustelhtm. Chico-
reus, and Hexaplex. .Additions and corrections. Tulane
Studies in Geology and Paleontology 23(1-3): 1-96.
Weaver, C. S. and J. E. DuPont. 1970. Living Volutes. A
monograph of the Recent Volutidae of the world. Dela-
ware Museum of Natural History, Greenville. 375 p.
THE NAUTILUS 105(4):165. 1991
Page 165
Mistaken Localities for Some Shells "from Surinam"
Takashi Okutani
Tok\o l'ni\ersit\ of Fisheries
4-5-7, Konan, Miiiato-ku
Tokvo 108. Japan
Dr. Philippe Bouchet, of the Museum national d'Histoire
naturelle, Paris, and Dr. William K. Emerson, of the
American Museum of Natural History, New York, co-
incidentulK alerted me that the type locality reported
for DallivoUita surinamensis was incorrect. They had
each recently obtained specimens from the Indian Ocean
that were identical to D. surinamensis. Both suggested
that the correct locality was not ir03'N, 61°01"W but
rather ITO.S'S, 61°0rE.
In publishing a survey report on shrimp trawl grounds
based on samples provided b\ the Japan Marine Fisheries
Research Center (JAMARC)' (Okutani, 19S3b), I discov-
ered several undescribed species of gastropods and bi-
valves in the material that had been sent to me for study.
A total of ten new species and one new genus were
described based on this material (Okutani, 1982, 1983a).
After being alerted b\ Drs. Bouchet and Emerson to the
problem with the locality data, I inquired of JAMARC
as to the origins of these samples.
JAMARC had conducted a bottom-fish survey on the
Saya de Malha Bank in the Indian Ocean in December
of 1978, and a survey of shrimp grounds off Surinam and
French Guiana from June of 1979 to March of 1983.
Although different vessels were used for each of these
surveys, the frozen samples were stored in the same re-
frigerated warehouse when the vessels returned to port.
Each frozen block was labeled with precise data as to
date, position, depth, haul number, etc. Among the large
series of samples, the designations N, S, E, W, were
sometimes omitted from the labels, although latitude and
longitude were always given. I was not aware of the
contamination of the samples as the numerical values of
the latitudes and longitudes were so close to each other.
In reviewing the records, I found that species described
on the basis of material collected prior to 1979 were all
from Indian Ocean samples. These ta.xa, together with
their correct type localities, are:
Murex (Murex) surinamensis Okutani, 1982
Type locality: I0°46'S, 6r32'E, in 94 m.
Dallivohita surinamensis Okutani, 1982 (now Lyria suri-
namensis, see Bouchet and Bail, 1991)
Type locality: 11°03'S, 6I°01'E, in 128 m.
Pteria pictura Okutani, 1983a
Type locality: 10°34'S, 61°35'E, in 97 m.
Spondylus jamarci Okutani, 1983a
Type locality: 11°03'S, 6r01'E, in 128 m.
The following ta.xa are from the continental shelf or
slope off Surinam or French Guiana as originally pub-
lished:
Plicifusus jamarci Okutani, 1982
Buccinofusus surinamensis Okutani, 1982
Volutomitra bayeri Okutani, 1982
Barbarofusus chocolatus Okutani, 1983a
Sarepta surinamensis Okutani, 1983a
Portlandia nigromaculata Okutani, 1983a
It is hoped that this note will clarify the doubts posed
by Ponder and Yokes (1988), Yokes (1990) and Bouchet
and Bail (1991).
LITERATURE CITED
Bouchet, P. and P. Bail. 1991 Mistaken geographical coor-
dinates: the saga of Lyria surinamensis and a new species.
The Nautilus 105(4):159-164.
Okutani, T. 1982. A new genus and five new species of gas-
tropods trawled from off Surinam. Venus 41(2)109-120,
Okutani, T 1983a. Four new species of bivalves and a new-
gastropod trawled from off Surinam. Venus 42(1 );17-2.5.
Okutani, T. 1983b, Moliusks, In. Takeda, T, and T, Okutani,
Crustaceans and Moliusks traw led off Suriname and French
Guiana. Japan Marine Fisher\ Resources Center, Tokyo,
p. 189-343.
Ponder, W. F. and E. H. Yokes. 1988. A revision of the Indo-
Paciflc fossil and Recent species of Murex ss. and Haus-
tellum (Mollusca: Gastropoda: Muricidae). Records of the
Australian Museum. Supplement 8: 1-160.
\dkes, E. H. 1990 What ever happened to dear old Murex?
American C:onchoiogist 18(l):3-7.
THE NAUTILUS 105(4):166-172, 1991
Page 166
New Species of Gaza, Mirachelus, Calliotropis, and
Echinogurges (Gastropoda: Trochidae) from the
Northwestern Atlantic Ocean
James F. Quinn, Jr.
Florida Marine Research Institute
Department of Natural Resources
100 Fighlh Avenue, SE
St. Petersburg, FL 33701-5095, USA
ABSTRACT
Five new species of Trochidae from the northwestern Atlantic
Ocean are described. Gaza olivacea, from the northern coast
of South America, is similar to C. superha (Dall, 1S81) but
differs in shell shape and characters of the outer lip and um-
bilical septum. Mirachelus acanlhus. known only from Ber-
muda, most resembles A/, clinocnemus Quinn, 1979, but differs
in shell shape and sculpture. Calliotropis globosa, from off the
Yucatan Peninsula, Mexico, Jamaica, and the northeastern coast
of South ,\merica from Venezuela to Suriname, is most similar
to C. actinophora (Dall, 1890) but differs in relative shell height
and umbilical diameter and in sculpture. Calliotropis dentata,
known only by the holotype from off Venezuela, is most similar
to C. persculpta (Sowerby, 1903) from off South Africa but has
a proportionately higher shell and stronger, coarser sculpture.
Echinogurges tuherciilatus. from off eastern Florida, the Straits
of Florida, and the northern Bahamas, is most similar to £.
clavatus (Watson, 1879) but is distinguished by sculptural de-
tails.
Key words: Trochidae; Gaza. Mirachelus; Calliotropis; Echi-
nogurges; systematics; new species.
INTRODUCTION
Many undescribed species were discovered during the
preparation of two geographically limited monographs
of western Atlantic Trochidae (Quinn, 1979, in press a).
This paper presents accounts of five new species of the
subfamilies Margaritinae and Eucyclinae (following the
cla.ssification of Hickman & McLean, 1990). New species
of the subfamilies Solariellinae and Calliostomatinae are
presented elsewhere (Quinn, 1991; in press a,b; in prep-
aration).
Institutional abbreviations used in this paper are as
follows: AMNH — American Museum of Natural History;
ANSP — Academy of Natural Sciences of Philadelphia;
DMNH — Delaware Museum of Natural History, Wil-
mington; UF — Florida Museum of Natural History, Uni-
versity of Florida, Gainesville; FSBC I — Florida Marine
Research Institute, Department of Natural Resources, St.
Petersburg; LACM — Natural History Museum of Los
Angeles County, Los Angeles; MCZ — Museum of Com-
parative Zoology, Harvard University, Cambridge;
TAMU— Texas A&M University, College Station;
UMML — Rosenstiel School of Marine and Atmospheric
Science, University of Miami, Miami; USNM — National
Museum of Natural History, Smithsonian Institution,
Washington.
SYSTEMATICS
Family Trochidae Rafinesque, 1815
Subfamily Margaritinae Stoliczka, 1868
Tribe Gazini Hickman and McLean, 1990
Genus Gaza Watson, 1879
Type species (monotypy): Gaza daedala Watson, 1879.
Gaza olivacea new species
(figures 1-3)
Gaza superha: Okutani, 1983:240.
Gaza sp.: Hickman and McLean, 1990: fig. 51 A.
Description: Shell large, attaining 38. 1 mm height, 44.9
mm width, broadly conical; whorl profiles weakly con-
vex, with broadly rounded peripheries; umbilicate with
umbilical septum; drab-olive to light brown, nacreous
under thin outer porcelaneous layer. Protoconch usually
missing. Teleoconch whorls about 6, weakly shouldered
subsuturally, weakly convex with broadly rounded pe-
ripheries, smooth except for microscopic spiral striae.
Base almost flat, smooth except for striae like those above.
Umbilicus wide, about 25% maximum whorl diameter;
wall smooth, parallel to shell axis. Aperture ovate; outer
lip oblique and reflected into slightly thickened, nacreous
rim, tangential to periphery of previous whorl; inner lip
reflected and expanded to form nacreous septum cov-
ering umbilicus; columella weakh' concave in immature
specimens, sigmoid in mature specimens.
J. F. Quinii, 1991
Page 167
^
^.
Figures 1-3. Caza olivacea new species, Hoiotype, I'SNM 752369, height 32.9 mm, width 39,8 mm, from N of Cabo de la Vela,
Peninsula de la Guajira, Colombia, Oregon Station 5690, 12°30'N, 72°08'W, 470 m.
Type material: Holotvpe, USNM 752369; 22 paratypes,
USNM 859424; Oregon Station 5690, 12°30'N, 72°08'W,
470 m; 40-foot otter trawl; 10 October 1965.— 17 para-
types, USNM 752390; 1 paratvpe, UMML 30.8365; 1
paratype, FSBC I 39513; Oregon Station 4301, 7°35'N,
54°13'W, 366 m; 65-foot otter trawl; 24 March 1963.
Type locality: North of Cabo de la Vela, Peninsula de
la Guajira, Colombia, Oregon Station 5690, 12°30'N,
72°08'W, 470 m.
Other material examined: 14 specimens, USNM 752391
(+ 61 uncatalogued USNM, and duplicates in other mu-
seums); Oregon Station 2026, 7°10'N, 53°07'W, 366 m;
40-foot otter trawl; 9 November 1957. — 15 specimens,
USNM 752389 (-1- 200-300 uncatalogued USNM, and
duplicates in other museums); Oregon Station 2005,
7°34'N, 54°50'W, 366 m; 40-foot otter trawl; 6 November
1957.— 1 specimen, USNM 752383; Oregon Station 1982,
10°00'N, 59°59'W, 457 m; 40-foot otter trawl; 3 Novem-
ber 1957—8 specimens, USNM 752388; Oregon Station
5028, ir.30'N, 60°46'W, 366-439 m; 40-foot otter trawl;
22 September 1964—2 specimens, USNM 752393; Or-
egon Station 5930, 15°38'N, 61°07'W, 808 m; otter trawl;
5 March 1966—19 specimens, USNM 752384 (-1- 54
uncatalogued USNM, and duplicates in other museums);
Oregon Station 2351, ir31'N, 62°24'W, 338-366 m; 40-
foot otter trawl; 23 September 1958. — 17 specimens,
USNM 752377; Oregon Station 2780, ll''36'N, 62°52'W,
393-421 m; 40-foot otter trawl; 20 April 1960—15 spec-
imens, USNM 752366; Oregon Station 4420, 11°46'N,
eg'n'W, 421 m; 40-foot otter trawl; 4 October 1963 —
7 specimens, USNM 752372; Oregon Station 5692,
12°31'N, 71°58'W, 375 m; 40-foot otter trawl; 10 October
1965. — 4 specimens, TAMU 4-1737; 2 specimens, TAMU
4-0388; Alaminos Station 70A10-40, 12°40'N, 72°00'W,
622-658 m; 57-foot trawl; 18 Julv 1970—9 specimens,
USNM 752373; Oregon Station 4923, 12°21'N, 72°40'W,
439-448 m; 40-foot otter trawl; 2 June 1964—13 spec-
imens, UMML 30.8364; John Elliott Pilkburij Station P-
781, 11°30.1'N,73°26.5'W, 567-531 m; 10-foot otter trawl;
30 July 1968—16 specimens, USNM 752376; Oregon U
Station 11252, 11°25'N, 73°56'W, 435 m; 71-foot otter
trawl; 10 November 1970—3 specimens, USNM 752374;
Oregon II Station 11251, ir23'N, 74°16'W, 457 m; 71-
foot otter trawl; 10 November 1970. — 7 specimens, USNM
752364; Oregon II Station 10266, 11°04'N, 74°25'W, 27
m; 71-foot otter trawl; 2 December 1968. — 12 specimens,
USNM 752365; Oregon II Station 11248, 11°18'N,
74°44' W, 600 m; 71-foot otter trawl; 9 November 1970.—
9 specimens, USNM 752363 (-f 100 uncatalogued USNM,
and duplicates in other museums); Oregon II Station
10260, 11°03'N, 7.5°18'W, 366 m; 71-foot otter trawl; 2
December 1970. — 5 specimens, USNM 752375; Oregon
Station 5722, 9°.36'N, 76°22' W, 512 m; 65-foot otter trawl;
16 October 1965—1 specimen, USNM 94991; Albatross
Station 2143, 9°30'45"N, 76°25'30"W, 283 m; small beam
trawl; 23 March 1884—4 specimens, USNM 752.392;
Oregon Station .3584, 9°13'N, 81°30'W, 366 m; 40-foot
otter trawl; 25 May 1962—20 USNM lots with localities
within range indicated above.
Remarks: Shells of Gaza olivacea are most similar to
those of Gaza superba (Dall, 1881) but differ in being
more broadly conical and in having more flattened whorl
profiles and much weaker spiral striae; the outer lip is
not angulate at the junction with previous whorl (figures
2, 3); and the nacreous septum covers the entire umbili-
cus. With the exception of the spiral striae and umbilical
septum, shells of G. olivacea differ from those of the
Galapagan G. rathbuni Dall, 1890, by the same char-
acters that differentiate G. olivacea and G. superba. Gaza
olivacea is known from Golfo de los Mosquitos, Panama,
eastward along the northern coast of South America to
Suriname and French Guiana. Although the depth range
for this species is 283-808 m (x = 433 m, n = 38), 87%
of all collections were made in depths of 350-550 m;
these analyses ignored records of 15 fms (27 m; Oregon
II Station 10266) and 25 fms (46 m; Oregon II Station
11275) because they are most likeK' errors for 150 fms
and 250 fms, respectively. These depths are shallower
than those for G. superba (range = 360-1,006 m; x =
Page 168
THE NAUTILUS, Vol. 105, No. 4
Figures 4-6. Mirachelus acanthus new species. 4. Paratype, DMNH 187590, height 3.45 mm, width 2.95 mm, trom oH Bermud.i.
.3. Same specimen, aperture, 27 x . 6. Same specimen, obHque view of protoconch and first whorl, 108 x.
556 m, n = 78), for which 86% of all collections have
been made between 440 m and 680 m (Quinn, in press
a). The geographic ranges of G. olivacea and G. superba
overlap slightly only in Panama in the west, and off
Grenada and Suriname in the east. A third western At-
lantic species, G. fischeri Dall, 1889, has a geographic
range that encompasses the ranges of both G. olivacea
and G. superba, but occurs in depths deeper than either
of those species (80% of lots from 550-825 m, x = 692
m; n = 33), although G. fischeri and G. superba have
sometimes been collected together (Quinn, in press a).
Like shells of G. superba, almost all shells of G. olivacea
have a small, neat hole replacing the protoconch and
leading into the umbilicus, and a narrow channel at the
junction of the umbilical rim and outer lip (figure 3),
both caused by a commensal polychaete worm that lived
in a mud tube within the umbilicus (Quinn, in press a).
Subfamily Eucyclinae Koken, 1897
Tribe Ch'ilodontini Wenz, 1938
Genus Mirachelus Woodring, 1928
Type species (original designation): Calliostoma corbis
Dall, 1889.
Mirachelus acanthus new species
(figures 4-6)
Description: Shell small for genus, attaining 3.70 mm
height, 2.95 mm width, conical, nonumbilicate, white,
nacreous under thin outer porcelaneous layer. Proto-
conch about 325 /irn maximum diameter, about one whorl.
Teleoconch whorls about 5.4; first 1.5-2 whorls with strong
axial lamellae, remaining strong but becoming more
rounded on subsequent whorls, spaced about 0.5 mm
apart on fifth whorl; very fine, oblique threads covering
whorl surface; 2 spiral cords appearing near end of sec-
ond whorl, abapical cord stronger and forming periph-
ery; weak angulation or spiral cord sometimes present
at suture; strong, conical tubercles at intersections of axial
and spiral sculpture. Rase convex, with 4-5 strong spiral
cords; spiral cords undulate or weakly beaded abaxially,
more distinctly beaded adaxially. Aperture oblique, ovate,
thickened within, with 8-9 weak ridges on thickening;
columella short, thickened, weakly inflated medially.
Type material: Holotvpe, DMNH 187589; 3 paratypes.
DMNH 187590; 1 paratype, USNM 860246; Lightbourn-
Guest Northstar Expedition Station 582, off Castle Har-
bour, Bermuda, 100 m.
Type locality: Off Castle Harbour, Bermuda, in 100 m.
Remarks: Shells of this species are most similar to those
of Mirachelus clinocnemus Quinn, 1979, but differ in
having weaker but sharper spiral cords, the peripheral
one stronger and more projecting than the one just adapi-
cal; in having the axial lamellae sharper and more wideK
spaced (0.5 mm apart in M. acanthus, 0.25 mm apart
in M. clinocnemus); and in having more prominent,
sharply conical tubercles on the peripheral and supra-
peripheral spiral cords. The shells of M. acanthus are
also slightly broader than those of M. clinocnemus.
Tribe Calliotropini Hickman and McLean, 1990
Genus Calliotropis Seguenza, 1903
Type species (original designation): Trochus ottoi Phi-
lippi, 1844.
Calliotropis glabosa new species
(figures 7, 8)
Description: Shell of medium size for genus, attaining
9.7 mm height, 8.7 mm width, turbinate with inflated
whorls, umbilicate, white, nacreous under thin outer por-
celaneous layer. Protoconch large, about 525-550 ^m
maximum diameter, 1 whorl. Teleoconch whorls 6.25;
first 1.5 whorls with thin axial riblets, rapidly becoming
weaker on next whorl, finally becoming restricted to
subsutural area on last w horl; pair of spiral threads ap-
pearing on first whorl, one subsutural, one just below
midwhorl and forming whorl periphery, both becoming
slightly stronger on sub.sequent whorls; third spiral thread
appearing on third v\ horl between suture and primary
subsutural spiral thread, gradually becoming slightly
J. F. Quinn, 1991
Page 169
/
r-
>
r
V
y
\
>
m
Figures 7, 8. Calliotropis globosa new species. Holot) pe, USNM 859419, height 9.45 mm, width S.70 mm, from S of Jamaica,
17°21.4'N, 77°.34.S'W, 805-1,089 m.
stronger than primary subsutural spiral thread on sub-
sequent whorls; fourth spiral thread similar to peripheral
thread, appearing on last whorl, coincident with suture
on spire whorls. Base convex, with 4 spiral cords; out-
ermost cord almost smooth, cords becoming progressively
more strongly tuberculate adaxially; innermost 2 cords
most closely spaced; transverse rugae becoming pro-
gressively stronger adaxially. Umbilicus deep, walls ax-
ialK rugose and almost vertical Aperture subovate, lips
thin; columella sigmoid, thin, with rather strong, rounded
tooth medially in some specimens.
Type material: Holotype, USNM 859419; 2 paratypes,
USNM 859420; 2 paratypes, ANSP 383289; 2 paratypes,
FSBC I 39515; 2 paratypes, UP 169956; 2 paratypes,
MCZ 302452; 2 paratypes, AMNH 232160; 21 paratypes,
UMML 30.8358; John Elliott Pillsbury Station P-i262,
17°21.4'N, 77°34.8'W, 805-1,089 m; 10-foot otter trawl;
15 July 1970.
Type locality: South of Jamaica, John Elliott Pillsbury
Station P-1262, 17°21.4'N, 77°34.8'W, 805-1,089 m.
Other material examined: 17 specimens, UMML
30.8355; John Elliott Pillsbury Station P-604, 18°58'N,
87°28'W, 970-988 m; box dredge; 17 March 1968.— 27
specimens, UMML 30.8356; John Elliott Pillsbury Sta-
tion P-605; 18°50.1'N, 87°31.5'W, 695-773 m; 10-foot
otter trawl; 17 March 1968. — 10 specimens, UMML
30.5703; 11 specimens, UMML 30.8361; John Elliott
Pillsbury Station P-607, 18°30'N, 87°37'W, 715-787 m;
10-foot otter trawl; 17-18 March 1968.— 3 specimens,
MCZ 294845; Blake Station XVIII, 18°20'30"N,
87°16'40"W, 1,097 m; 1880.— 4 specimens, UMML
30.8360; John Elliott Pillsbury Station P-1255, 17°18'N,
78°32'W, 622-823 m; 10-foot otter trawl; 14 July 1970.—
2 specimens, UMML 30.8359; John Elliott Pillsbury Sta-
tion P-1261, 17''13'N, 77°50'W, 595-824 m; 10-foot otter
trawl; 15 July 1970—1 specimen, MCZ 135174; A</an<ts
Station 3454, 23°24'N, 80°36'W, 1,097 m; 4 May 1939 —
4 specimens, MCZ 135175; Atlantis Station 3457, 23°23'N,
80°36'W, 1,006 m; 4 May 1939—3 specimens, MCZ
135179; Atlantis Station 3345, 2r08'N, 79°56'30"W,
1,280 m; 8 April 1939—3 specimens, MCZ 135177; 6
specimens, MCZ 135178; Atlantis Station 3366, 20°46'N,
74°59'W, 1,097 m; 19 April 1939—5 specimens, MCZ
135183; Atlantis Station 3359, 20°38'N, 74°32'W, 1,829
m; 18 April 1939—1 specimen, UMML 30.8357; John
Elliott Pillsbury Station P-919, 16°05.3'N, 61°19.3'W,
704-732 m; 5-foot Blake trawl; 12 July 1969—6 speci-
mens, UMML 30.8362; John Elliott Pillsbury Station P-
850, ir45.5'N, 61°29.5'W, 896-923 m; 10-foot otter trawl;
3 July 1969—9 specimens, UMML 30.7337; John Elliott
Pillsbury Station P-846, 11°37.8'N, 60°37.4'W, 878-942
m; 10-foot otter trawl; 2 July 1969. — 1 specimen, UMML
30.7347; John Elliott Pillsbury Station P-847, ir37.3'N,
0°59.4'W, 920-1,244 m; 41-foot otter trawl; 2 July 1969 —
13 specimens, UMML 30.6912; John Elliott Pillsbury
Station P-754, irSe.O'N, 68°42.0'W, 684-1,574 m; 10-
foot otter trawl; 26 July 1968.
Remarks: Shells of this species are most similar to those
of Calliotropis actinophora (Dall, 1890) in size, general
shape, and sculpture but differ in being proportionately
narrower; in having narrower umbilici; in having stron-
ger, more widely spaced subsutural axial rugae; in having
the adaxial two spiral rows of tubercles more closely
spaced; and in having four rather than three basal spiral
cords. Calliotropis globosa is known from the Yucatan
Page 170
THE NAUTILUS, Vol. 105, No. 4
/
f %\-'^
Figures 9-1 1. Calliotropis dentata new species. Holot>pe, USNM 859419, height 9.4 mm, width 9.2 mm, from off Isla Orchilla,
Venezuela, ll''47.8'N, 66°06.8'W, 1,052-1,067 m.
coast near Banco Chinchorro, from Cay Sal Bank and
Cuba, from south of Jamaica, and along the northern
South American coast from Venezuela to Suriname,
mostly in depths of about 700-1,100 m (range = 595-
1,829m).
Calliotropis dentata new species
(figures 9-11)
Description: Shell of medium size for genus, attaining
at least 9.4 mm height, 9.2 mm width, conical with
shouldered whorls, peripherally carinate, strongly sculp-
tured, white, nacreous under outer porcelaneous layer.
Protoconch and much of first whorl missing. First re-
maining whorl with narrow axial riblets and weak sub-
sutural and peripheral spiral angulations. Second whorl
largely decorticated but showing strengthening angula-
tions. Subsutural angulation becoming spiral row of strong,
conical, axially elongate tubercles rising above level of
suture; peripheral angulation forming carina set with
closely set, cog-like tubercles connected and crossed by
fine spiral thread. Final whorl with second carina, similar
to peripheral carina but slightly weaker, abapical to pe-
ripheral carina and coincident with suture line. Whorl
surface between spiral carinae flat, with fine, collabral
growth lines. Base convex, with 3 strong, tuberculate
spiral cords; whorl surface between innermost 2 spiral
cords with weak collabral rugae that generally connect
tubercles of cords. I'mbilicus about .30% maximum shell
diameter, furuiel-shaped, walls with axial rugae and sin-
gle weak spiral thread. Lips of aperture and columella
broken.
Type material: Holotype, USNM 859422; John Elliott
Pillslmrij Station P-741, 11°47.8'N, 66°06.8'W, 1,052-
1,067 m; 40-foot otter trawl; 23 July 1968.
Type locality: Off Isla Orchilla, Venezuela, John Elliott
Pilhbury Station P-741, ir47.8'N, 66°06.8'W, 1,052-
1,067 m.
Remarks: The shell of Calliotropis dentata is easily
distinguished from those of all other western Atlantic
Calliotropis by having the peripheral and subperipheral
spiral cords with closely set, cog-like beads instead of
conical tubercles. The shell of C. dentata is very similar
in sculpture to that of C. persculpta (Sowerby, 1903) (see
Kaicher, 1990: card 5695) from South .Africa but differs
in being higher spired and in having stronger basal spiral
cords and a narrower umbilicus.
Genus Echinogurges Quinn, 1979
Type species (original designation): Trochus (Mar-
garita) clavatus Watson, 1879.
Echinogurges tubercidatus new species
(figures 12-14)
Echinogurges rhysus: Quinn, 1979:21, 22, figs. 39, 40.
[Misidentification; not Trochus (Margarita) rhysus Wat-
son, 1879.]
Description: Shell small, attaining 4.6 mm height, 3.9
mm width, conical with shouldered whorls, strongly
sculptured, umbilicate, white, nacreous under thin outer
porcelaneous layer. Protoconch about 375 fim maximum
diameter, bulbous, 1 whorl. Teleoconch whorls 5.5, tu-
bular, carinate, first whorl with strong, thin axial riblets
that become weak collabral ridges on subsequent whorls;
pair of spiral ridges appearing on second whorl, one
subsutural, one at midwhorl forming carinate whorl pe-
riphery; third tuberculate spiral ridge present on body
whorl, coincident with suture on spire whorls; strong,
conical tubercles at intersections of axial and spiral sculp-
ture. Base convex, rounding into umbilicus, with 4
smoothish to finely tuberculate spiral cords. L'mbilicus
narrow, pore-like, wall with axial rugae. Aperture sub-
circular, lips thin, inner lip weaklv reflected into um-
bilicus; columella concave, thin, rounding smoothly into
outer lip.
Type material: Holotvpe, USNM 330606; 4 paratypes,
USNM 859426; Albatross Station 2654, 27°57'3'0"N,
77°27'30"W, 1,207 m; large beam trawl; 2 May 1886.—
13 paratypes, MCZ 302536; Atlantis Station 2993,
J. F. Quinn, 1991
Page 171
v^ ^
Figures 12-14. Echinogurges tuberculatus new species. 12, 13. Paratype, UMML 30 8095, height 4,0 mm, width 3.1 mm, from
southern Straits of Florida, Cerda Station G-1106, 24°02'N, 81°30'W, 1,706-1,723 m. 14. Protoconch of paratype (USNM 859426)
from NE of Little Bahama Bank, Albatross Station 2654, 27°57'30"N, 77°27'30"W, 1,207 m, 97 x ,
33°19'N, 80°44'W, 1,061 m; 15 March 1938.— 4 para-
types, USNM 859426; Albatross Station 2415, 30°44'N,
79°26'W, 805 m; large beam trawl; 1 April 1885.— 2
paratvpes, UMML 30.8054; Gerda Station G-368,
24''03'N, 8riO'W, 961-1,016 m; 16-foot otter trawl; 15
September 1964.— 1 paratype, UMML 30.8041; Gerda
Station G-130, 23°59'N, 81°10'W. 1,021 m; 10-foot otter
trawl; 21 June 1963—1 paratype, UMML 30.8163; Ger-
da Station G-859, 23°54'N, 8r57'W, 1,161-1,200 m; 10-
foot otter trawl; 29 August 1967. — 2 paratypes, USNM
438284; 10 miles N of Cuba, 1,427 m; J. B. Henderson
coll. — 2 paratypes, MCZ 7584; Blake Station, Yucatan
Channel, 1,171 m.
Type locality: Northeast of Little Bahama Bank, Al-
batross Station 2654, 27°57'30"N, 77°27'30"W, 1,207 m.
Other material examined: 1 specimen, UMML 30.8095;
Gerda Station G-1106, 24°02'N, 81°30'W, 1,706-1.723
m; 10-foot otter trawl; 29 April 1969. — 4 specimens, MCZ
135026; Atlantis Station 2988, 23°15'N, 79°57'W, 695 m;
14 March 1938.
Remarks: Shells of Echinogurges tuberculatus are most
similar to those of E. clavatus (Watson, 1879) but differ
in having only a single peripheral spiral ridge; weaker,
more widely spaced axial sculpture; and fewer basal spi-
ral cords. Specimens of E. tuberculatus were misiden-
tified as £. rhysus (Watson, 1879) by Quinn (1979). Sub-
sequent examination of the types of Trochus (Margarita)
rhysus Watson, 1879, revealed that T. (M.) rhysus is
actually a species of Calliotropis Seguenza, 1903 (Quinn,
in press a). Echinogurges tuberculatus is known only
from off the east coast of Florida, the Bahama Islands,
and the Straits of Florida in depths of 805-1,723 m.
ACKNOWLEDGMENTS
I thank the late Gilbert L. Voss (Rosenstiel School of
Marine and Atmospheric Science, University of Miami)
for access to the collections under his care and for per-
mission to describe the new species in that material.
Richard S. Houbrick and the late Joseph Rosewater (both
USNM), and Riidiger Bieler (formerly DMNH) kindly
allowed me to examine types and other material. Russell
Jensen (DMNH) brought the Bermuda specimens to my
attention. Marjorie Myers, Llyn French, and Judy Leiby
provided valuable word-processing and editorial assis-
tance. Thomas H. Perkins and William G. Lyons (both
Florida Marine Research Institute, Department of Nat-
ural Resources), and two anonymous reviewers provided
valuable critiques of previous drafts of this paper.
LITERATURE CITED
Dall, W. H. 1881. Reports on the results of dredging, under
the supervision of Alexander Agassiz, in the Gulf of Mex-
ico, and in the Caribbean Sea, 1877-79, by the United
States Coast Survey steamer "Blake," Lieutenant-Com-
mander C. D Sigsbee, L'.S.N., and Commander J. R Bart-
lett, U.S.N., commanding. W. Preliminars report on the
Mollusca. Bulletin of the Museum of Comparative Zool-
ogy, Harvard University 9(2):33-144.
Dall, W H. 1889. Reports on the results of dredging, under
the supervision of Alexander Agassiz, in the Gulf of Mexico
(1877-78) and in the Caribbean Sea (1879-80), by the U.
S. Coast Survey steamer "Blake , Lieut -Commander C
D Sigsbee, U S N , and Commander J R Bartlett, US N ,
commanding. XXI.X. Report on the Mollusca Part 2, Gas-
tropoda and Scaphopoda Bulletin of the Museum of Com-
parative Zoology, Harvard I'niversity 18:1-492.
Dall, W. H. 1890. Scientific results of explorations by the U.
S. Fish Commission steamer "Albatross '. VII. Preliminary
report on the collection of Mollusca and Brachiopoda ob-
tained in 1887-88. Proceedings of the United States Na-
tional Museum 12(773):219-362.
Hickman, C. S. and J. H. McLean. 1990. Systematic revision
and suprageneric classification of trochacean gastropods.
Natural History Museum of Los .Angeles Count)-, Science
Series, No. 35, vi -I- 169 p.
Kaicher, S. D. 1990. Card catalogue of world-wide shells
Pack #56 — Trochidae \'. PrivateK published, St. Peters-
burg, FL.
Okutani,"T. 1983. Mollusks In: Takeida, M and T Okutani
Crustaceans and mollusks trawled off Suriname and French
Guiana. Japan Marine Fisherv Resource Research Outer,
Tokyo, p. 187-354.
Page 172
THE NAUTILUS, Vol. 105, No. 4
Philippi, H. .\. 1844. Enunicratio molhist-orum Siciliae cum
viventium turn tellure tertiaria fossiliuni. quae in itinirt-
suo observavit, Vol 2 Halis Sa.\oiuini, iv + 303 p.
Quinn, J. F., Jr. 1979. Hinlogical rt-sults of the University of
Miami deep-.sea e.xpcditioiis 130 The systematics and zo-
ogeograph\ of tlie gastropod family Trochidae collected
in the Straits of Florida and its approaches. Malacologia
19(1); 1-62.
Quinn, J. F., Jr. 1991. Larucllilwchus. a new genus of So-
lariellinae (Gastropoda: Trochidae), with descriptions of
six new species from the western .Atlantic Ocean The
Nautilus 105(3):81-91.
Quinn, J. F., Jr. In press a The Trochidae of the Gulf of
Mexico (Prosobranchia: .'Vrcliaeogastropoda), Memoirs of
the Hourglass CruLses.
Quinn. J. !■' , Jr In press b. New species of Soliiriclla (Gas-
tropoda: Trochidae) from the uestern .\tlantic Ocean The
Nautilus.
Seguenza, G. 1903. Molluschi poco noti dei terreni terziarii
di Messina Societa Geologica Italiana, Bolletino 21:455-
464.
Sowerby, (i. H , 111. 1903 Mollusca of South .Africa Marine
Investigations in South .\frica 2:213-232.
Watson, R. B 1879. Mollusca of H.M.S. Challenger Expe-
dition. III. Trochidae, viz. the genera Seguenzia. Basilissa,
Gaza, and Bernhix. Journal of the Linnean Society of Lon-
don, Zoology 14:586-605.
Woodring, W. P. 1928. Miocene moUusks from Bowden, Ja-
maica, II. Gastropods. Carnegie Institution of Washington,
Publication 385:1-564,
THE NAUTILUS 105(4):173-174, 1991
Page 173
Acroloxus coloraderisis (Henderson), a Rare
North American Freshwater Limpet
Andrew J. Paul
Hugh F. Clifford
Deparlment of Zoology
Universit) of Alberta,
Edmonton, .'Alberta,
Canada T6G 2E9
The freshwater limpet family Acroloxidae (Basomma-
tophora) contains seven species, all in the genus Acro-
loxus. One, A. laciistris (Linnaeus), is common in lentic
habitats of much of Europe and parts of Asia. Five spe-
cies, A. improvistts (Polinski), A. macedonicus (Had-
zisce), A. kobelti (Dybowski), A. sibiricum (Gerstfeldt),
and A. froscheli (Dybowski), appear to be endemic to a
few ancient lakes (e.g.. Lake Ochrid and Lake Baikal)
of Yugoslavia and U.S.S.R. (Clarke, 1973). Only one spe-
cies, A. coloradensis (Henderson), is found in North
America. It is readily distinguished from other fresh-
water limpets by its spine-like apex, which is directed
posteriorly towards the left margin (figure 1). Acroloxus
coloradensis has hitherto been reported from only seven
North American localities. Four of the records are from
Rocky Mountain lakes: Colorado (Walker, 1925), Alberta
(Moziey, 1930), Montana (Russell and Brunson, 1967),
and British Columbia (Clarke, 1981). The other three
records are from ponds in Ontario and Quebec (Clarke,
1970).
We recently collected Acroloxus coloradensis speci-
mens from one location of the Beaver River in Alberta,
Canada. Voucher specimens are deposited in the Ca-
nadian Museum of Nature, Ottawa, Ontario, Canada,
catalog number #92900. This is the first report of A.
coloradensis collected from a stream and apparently the
first report of any of the seven Acroloxus species from a
lotic habitat. The Beaver River runs through the southern
limit of mixed boreal forest and the northern limit of
agricultural activity in northeastern Alberta and is a trib-
utary of the Churchill River, which empties into Hudson
Bay.'
We sampled various sites of the Beaver River, but
Acroloxus coloradensis was collected at only one loca-
tion: where the Beaver River is crossed bv a small road
bridge south of Highway 55 (54°32'N, lli°52'W). Spec-
imens were collected to a depth of about 0.8 meters.
Samples were collected between 30 May and 2 August
1989 by kick samples taken with a dip net (mesh size:
500 fim) and by inspecting substratum taken from the
river. The width of the Beaver River at the collecting
site was 17.5 m; water temperatures ranged from 17 °C
on 30 May 1989 to 25 °C on 20 July 1989. There were
no riffle regions in the vicinity, and current velocity was
undetectable using a Price AA current meter. Free-float-
ing and submergent macrophytes were common at the
sampling site. The river's substratum immediateK up-
stream and downstream of the collecting site (the bridge)
consisted of mud and plant detritus with rooted mac-
rophytes near the banks of the river. However, A. colora-
densis was collected only where the bridge crossed the
stream, and here the substratum consisted of gravel, peb-
Figure 1 . Scanning electron micrographs of Acroloxus colora-
densis from the Beaver River, .'\lberta, Canada, 1. Side view.
2. Top view.
Page 174
THE NAUTILUS, Vol. 105, No. 4
bles and large cobbles; presuinabK the large substrata
was introduced when the bridge was constructed. Other
gastropods collected from this location were Valvata sin-
cera helicoidea Dall; Menetiis cooperi Raker, Prome-
netus exacuous exacuous (Say), Armiger crista (Linnae-
us), Lijmnaea stagnaiis jugularis (Say), Stagnicola spp.,
Helisoma spp., and Gtjraulus spp.
Bryce (1970) postulated that Acroloxits coloradensis,
based on its known distribution at tliat time, was a cold-
water stenotherm, limited to Rocky Mountain lakes. More
recent records of the limpet in Ontario and Quebec
(Clarke, 1970), Colorado (VVu 1989), and our finding
indicate that A. coloradensis is not restricted to Rocky
Mountain lakes. The large temperature variations (17°-
25 °C) experienced over the summer by A. coloradensis
in the Beaver River indicate that it is not a cold-water
stenothermal species.
Clarke (1973) and Burch (1982) suggest A. coloraden-
sis might be a rare species with a relic distribution. Its
discovery in the Beaver River is not inconsistent with
this idea, and we can only speculate on its occurrence in
the Beaver River of Alberta, Canada. Possibly A. colora-
densis will eventually be recorded from Saskatchewan
and Manitoba, indicating a wide but rare distribution
across Canada on both sides of and in the Cordillera.
The southerly populations of A. coloradensis, e.g., in
Montana and Colorado, might now be restricted to in
and near the Cordillera.
ACKNOWLEDGEMENTS
We would like to thank Dr. Arthur H. Clarke, Eco-
search, Inc., Portland, Texas, for verifying the identifi-
cation of A. coloradensis. Karen A. Saffran (University
of Alberta) provided technical assistance; Ronald Koss
(University of .Alberta) took the scanning electron mi-
crographs. Financial assistance was provided to .^JP and
HFC by the Natural Sciences and Engineering Research
Council of Canada.
LITERATURE CITED
Bryce, G. W. 1970. Rediscovery of the limpet Acroloxus
coloradensis (Basommatophora: .\croloxidae) in Colorado.
The Nautilus 83:105-108.
Burch, J B 1982. Freshwater snails (MoUusca: Gastropoda)
of North America. United States Environmental Protection
Agency, Cincinnati, OH, 294 p.
Clarke, A, H. 1970. On Acroloxus coloradensis (Henderson)
(Gastropoda, Basommatophora) in Eastern Canada. Pub-
lications in Zoology, No, 2, National Museums of Canada,
13 p.
Clarke, A. H. 1973. The freshwater molluscs of the Canadian
Interior Basin. Malacologia 13; 1-509,
Clarke, A, H, 1981, The freshwater molluscs of Canada. Na-
tional Museum of Natural Sciences/National Museums of
Canada, Ottawa, Ontario, 446 p.
Mozley, A, 1930. Reports of the Jasper Park Lakes investi-
gations, 1925-26. The Mollusca of Jasper Park. Transac-
tions of the Royal Society of Edinburgh 56:647-669.
Russell R, H, and R. B. Brunson, 1967. Acroloxus coloradensis
from Montana, The Nautilus 81:33,
Walker, B, 1925. New species of North American .\ncylidae
and Lancidae, Occasional Papers of the Museum of Zo-
ology, University of Michigan 165:1-13,
Wu, S.-K. 1989. Colorado freshwater mollusks. Natural His-
tory Inventory of Colorado, No. 11:1-117.
w, t ■J
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