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VOL. 36, NO. 1-2 . MALACOLOGIA | 1995 |
| | Aue | Se TUE

CONTENTS о

MARÍA VILLARROEL Y JOSÉ STUARDO i ey
Morfologia del Estomago y Partes Blandas en Mytella strigata (Hanley 1849 Mey

(Bivalvia: Mytilidae) ia La 222 2a ns о a A Pet ERBE fi |
JOST BORCHERDING К 3 п
Laboratory Experiments on the Influence of Food Availability, Temperature and FX }
- Photoperiod on Gonad Development in the Freshwater Mussel Dreissena EN
„NPalymearpha 2/72 So oe) O A er ee RSR АЕ Е i 15 4
R. ‚ ARAUJO, J. М. ВЕМОМ, D. МОВЕМО & M. А. RAMOS HER - 39
Relaxing Techniques for a! Molluscs: Trials for Evaluation of Different à
Méthode Ch Den И O a ee
KENNETH C. EMBERTON qlo TEN

Land-Snail Community Morphologies of the Highest-Diversity Sites of Ман
gascar, North America, and New Zealand, with RECO Alternatives to”.
HelghteDiameter Plots N Rune ac eben ben eee eee See >
KENNETH C. EMBERTON \ ENT FO | dino PE
Distributional Differences Among Acavid Land Snails Around Antalahdy Mada- eee

gascar: Inferred Causes and Dangers Of Extinction N RTS ESS ES HA. 4
Ir KATHERINE COSTIL & JACQUES DAGUZAN | ne iR
+ Effect of Temperature on Reproduction in Planorbañus corneus (L.) and Plan- . I
orbis planorbis (L.) Throughout the Life “ad О OS + eats See ИТ
L М. COOK 4 J. BRIDLE ewe ye! 7
1 Colour Polymorphism in the a Snail Littoraria intermedia т Sinai..... 91
> MICHAEL G. GARDNER, PETER В. MATHER, IAN WILLIAMSON & JANE M. HUGHES : via
р The Relationship Between Shell-Pattern Frequency and Microhabitat Variation nS
- in the Intertidal Prosobranch, ‚Clithon oualaniensis (Lesson) .......... Jesse ta

MIGUEL, IBÁÑEZ, ELENA PONTE-LIRA 8 MARÍA R. ALONSO < |
El Género Canariella Hesse, 1918, y su Posición: en Ja Familie Hygromidae e

| | Gastropoda, Pulmonata, Helicoidea) .......... rn IA 2 «le wu; EN
"N. ELEUTHERIADIS & М, LAZARIDOU-DIMITRIADOU

~~ Age-Related Differential Catabolism in the Snail Bithynia graeca (Westárluñd, À ay
1879) and its Significance in the Bioenergetics of Sexual Dimorphism “4 №, ee

HEINZ BRENDELBERGER | ' Y # Kar
Dietary Preference of Three Freshwater Gastropods for Eight Natural Foods of |
Different Engraetic ‘Content Lil. a de ee PE VE SEE a

ROBERT H. COWIE, GORDON M. NISHIDA, YVES BASSET & SAMUEL M. GON, mo
Patterns of Land Snail Distribution 16, a Montane Habitat on the ‘sland oft

> Ва [a dis aye a ele ase mia A O AA LER e 15
ALAN Е. STIVEN a ; O
E Genetic Heterozygosity and Growth Rate in the Southern Appalachian Land A
4 y Snail Mesodon normalis (Pilsbry 1900): The Effects of реа Stress . Tey
| DAVID В. LAWRENCE / IR N N Rr A sats.
| Diagnosis of the Genus Crassostrea (Bivalvia, Ostreidae) . ón ot tk an, |:
KENNETH С. EMBERTON & SIMON TILLIER | i р 7 o! 14
Clarification and Evaluation of Tillier’s a0 989) Siylommatophora Mono-
1 EA A A A E AS AU RS
grap RA Bu ae
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} VOL. 37, NO. 1 LIBRARY 1995
À | 210. 1995
y JER ds | TY
MALACOLOGIA
\
À
International Journal of Malacology
Revista Internacional de Malacologia

Journal International de Malacologie
Международный Журнал. Малакологии

Internationale Malakologische Zeitschrift

MALACOLOGIA | he
Editor-in-Chief:
GEORGE M. DAVIS

t

Editorial and Subscription Offices:

Department of Malacology
The Academy of Natural Sciences of Philadelphia т
1900 Benjamin Franklin Parkway D Sp I
Philadelphia, Pennsylvania 19103-1195, U.S.A. ь

Co-Editors: ^ not TIER:

EUGENE COAN CAROL JONES KR
California Academy of Sciences ~ Denver, CO - A
San Francisco; CA ;

` Assistant Managing Editor: Е
CARYL HESTERMAN er
` Associate Editors: ” ieee
и ANNE GISMANN
University of Michigan Ма

: ) | | €
Ann Arbor ' ‚Egypt, : Er y

MALACOLOGIA is published by the INSFITUTE OF MALACOLOGY, the Sponsor Members OF
which (also serving as editors) are: | | 4

- RUDIGER BIELER ^^ ALAN KOHN
Field Museum, Chicago University, of Washington, Seattle à
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President Elect | California |
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KENNETH J. BOSS. | y
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| "Copyright © 1995 by the Institute of Mall ; Y

J. A. ALLEN
Marine Biological Station
Millport, United Kingdom

E. E. BINDER
Muséum d'Histoire Naturelle
Genève, Switzerland

А. J. САМ
University of Liverpool
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P. CALOW
University of Sheffield
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University of North Carolina
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R. COWIE
Bishop Museum
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Е. GIUSTI
Universita di Siena, Italy

А. М. GOLIKOV
Zoological Institute
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1995
EDITORIAL BOARD

5. J. GOULD
Harvard University
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A. V. GROSSU
Universitatea Bucuresti
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T. HABE
Tokai University
Shimizu, Japan

R. HANLON
Marine Biomedical Institute
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J. А. HENDRICKSON, уг.
Academy of Natural Sciences
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Forschungsinstitut Senckenberg,
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Natal Museum
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Museo Nacional de Historia Natural
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A. LUCAS
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Tropenmedizinisches Institut
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Н. К. MIENIS
Hebrew University of Jerusalem
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J. E. MORTON
The University
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J. J. MURRAY, Jr.
University of Virginia
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R. NATARAJAN
Marine Biological Station
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J. OKLAND
University of Oslo
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Т. OKUTANI
University of Fisheries
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Instituto Oswaldo Cruz, Rio de Janeiro

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J. J. PARODIZ
Carnegie Museum
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J. P. POINTER
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Australian Museum
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QUIZE
Academia Sinica
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D. G. REID
The Natural History Museum
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University College of North Wales
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S. G. SEGERSTRLE
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А. STANCZYKOWSKA
Siedlce, Poland

Е. STARMÜHLNER
Zoologisches Institut der Universitát
Wien, Austria

У. |. STAROBOGATOV
Zoological Institute
St. Petersburg, Russia

W. STREIFF
Université de Caen
France

J. STUARDO
Universidad de Chile
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S. TILLIER
Muséum National d'Histoire Naturelle
Paris, France

R. D. TURNER
Harvard University
Cambridge, Mass., U.S.A.

J.A.M. VAN DEN BIGGELAAR
University of Utrecht
The Netherlands

J. А. VAN EEDEN
Potchefstroom University
South Africa

N. H. VERDONK
Rijksuniversiteit
Utrecht, Netherlands

B. R. WILSON

Dept. Conservation and Land Management

Kallaroo, Western Australia

Н. ZEISSLER
Leipzig, Germany

A. ZILCH
Forschungsinstitut Senckenberg
Frankfurt am Main, Germany

MALACOLOGIA, 1995, 37(1): 1-11

THE LIFE CYCLE, DEMOGRAPHIC ANALYSIS, GROWTH AND
SECONDARY PRODUCTION OF THE SNAIL HELICELLA (XEROTHRACIA)
PAPPI (SCHÜTT, 1962) (GASTROPODA PULMONATA)

IN E. MACEDONIA (GREECE).

M. Lazaridou-Dimitriadou

Laboratory of Zoology, School of Biology, Faculty of Sciences, Aristotle University of
Thessaloniki, 54006 Thessaloniki, Macedonia, Greece

ABSTRACT

The life cycle, population dynamics, and growth of the pulmonate snail Helicella (Xerothracia)
pappi were studied in northern Greece. The spatial distribution of H. рарр! was found to be
contagious. Demographic analysis of its population revealed that (a) three cohorts exist
throughout the year, (b) increased growth rate is observed during spring, (c) snails attain their
maturity 21 months after hatching when their greatest shell diameter reaches 15 mm, and (d)
it is an iteroparous species, with egg-laying in autumn. Von Bertalanffy analysis showed that H.
рарр! may reach its maximum size (dead shells collected in the field had 25.5 mm maximum
shell diameter) in four years. Shell morphology changes when spermatozoa are fully formed in
the gonad for the first time. Mortality rate is high just after hatching and also after winter, when
tissue degrowth occurs, at which time the snails lose 33% of their biomass. Life expectancy
decreases with increasing age. Net reproductive rate (Ro) was 3.025, per capita rate of increase
(г) is 0.04, annual production (P) was 5.82 + 0.45 g/(m?-yr) in 1987 and 3.73 + 0.31 g/(m?-yr) in
1988, mean standing crop (В) was 2.89 g/m? in 1987 and 1.81 g/m? т 1988, and annual
productivity rate constant (P/B) was 0.2 per year in both 1987 and 1988.

Key words: Biology, ecology, growth, production, snail, Helicella pappi, Xerolenta obvia.

INTRODUCTION

Little is known about the species Helicella
(Xerothracia) pappi Schútt, 1962. Wagner
(1927) referred population of this species
from near our study area in Xanthi, southern
Thrace, to Martha filimargo Krynicki, placing
it in the genus Helicopsis Fitzinger, 1833. Ur-
banski (1960) said that samples from south-
ern Thrace of Helicella (Helicopsis) filimargo
(Rossmássler) did not differ from samples of
Krimea and Odesco. Schútt (1962) described
the shell and genital apparatus of samples of
Helicella (Xerothracia) pappi from a type lo-
cality in Philippi, Kavala, eastern Macedonia;
he claimed that this species could not be
placed in the genus Helicopsis, as Wagner
(1927) had done, because of such shell fea-
tures as curvature and the presence of a keel.
Hausdorf (1988) claimed that H. рарр! could
not be placed in a subgenus of Helicella be-
cause it differs in the dart sac and in the
nerves coming from the cerebral ganglia. He
concluded that it is a subspecies of Xerolenta
obvia.

In the present study we are using the name
Helicella (Xerothracia) pappi following Schütt

(1962), because the samples come from the
type locality. We have studied the biology
and ecology of this species, which 1$ re-
stricted to Philippi, eastern Macedonia, in or-
der to find out why this species does not
have a broader and continuous distribution
from Krimea to Greece.

METHODS AND MATERIALS

Helicella рарр! was studied in the archae-
ological site of Philippi, 25 km from Kavala,
eastern Macedonia, Greece. Philippi is a
limestone area with limited vegetation, т-
cluding mosses, lichens, several Taraxacum
species, and succulent plants. Several grass
species are dominant. Apart from Н. pappi,
there are small, sparse populations of Lindol-
holmia lens and Helix figulina. Weather con-
ditions during the investigation period are
shown in Figure 1.

Random samples (Lewis 4 Taylor, 1972)
were taken every month for three consecutive
years between April 1986 and April 1989. No
samples were taken in winter (December
through February), when the snails are in di-

2 LAZARIDOU-DIMITRIADOU

Mean monthly
90 temperature (°С)
==_==

Total precipitation 50
(mm)
ern 45

A86M J J AS О М ОМ87А M J J AS О NM88A MJ JAS ON DM89A М

Months

FIG. 1. Ombrothermic curve showing mean monthly temperatures (°С) and total monthly precipitation (mm)
at Philippi from April 1986 to Мау 1989 (striated areas represent arid periods of the year).

apause and hidden under vegetation. Quadrat
sample-size (25 x 25 cm?) was determined by
Healy's method (Cancela da Fonseca, 1965).
Elliot's (1971) method was used to determine
the necessary total number of sampling units
(sampling error less than 20%). Sampling was
carried out during morning hours in the ab-
sence of rain. All snails found in a quadrat
were collected, measured and then replaced.
The largest diameter of the shell (D) and the
peristome diameter (d) were measured with
vernier calipers to the nearest 0.1 mm. D was
used for size-frequency histograms, using
3-mm class intervals (Cancela da Fonseca,
1965). The cohorts were discriminated using
probability paper (Harding, 1949). This
method was valid because the modes of the
age-classes were separated by at least 2.5
standard deviations (Grant, 1989), except for
two cases, one in March 1987 and one in June
1988; although many age classes had less
than 50 individuals, the modal values were
consistent from month to month. This con-
firms that the modes were real and not the
result of sampling variation. The same method
has been used before for demographic anal-
yses of other populations of molluscs
(Hughes, 1970; Lévéque, 1972; Daguzan,
1975; Lazaridou-Dimitriadou et Kattoulas,
1991).

There were no statistically significant vari-
ations in field density over the three years.
Life data and rate estimates were based on
successive samples over 1987 and 1988.

Spatial distribution of the snails in the hab-

itat was examined by using Taylor's (1961)
power law 6° = ax”, where 6° = variance and
x = mean number of snails/0.25 m“.

For the study of relative growth, the mor-
phometric criteria of shell diameter (D) in re-
lation to the peristome diameter (d) were
used from all the animals sampled during
1987 (М = 2567). Mayrat's method (1965а, b)
was used to compare the growth of D in re-
lation to d between immature and mature
snails. A logarithmic transformation was ар-
plied to the data.

Bertalanffy’s (1938) method was used to
calculate the theoretical growth curve and life
span; this method is widely used (Moreteau,
1987).

Life-table and fertility data, as well as an-
nual secondary production, were estimated
as described for Helix lucorum (Staikou et al.,
1988) and Eobania vermiculata (Lazaridou-
Dimitriadou 4 Kattoulas, 1991). Annual pro-
duction is calculated by the size-frequency
method because single cohorts need not be
identified.

For dry body-weight (Wb) analysis, 100 an-
imals comprising five from each size class
were individually marked and their greatest
shell diameter (D) was measured; they were
then dried to constant weight over a period of
36 h in a vacuum at room temperature. Shell
organic matter was calculated as the residual
weight of dry shell after treatment with 5 m
mol/ml HCl, successive washing over a fine
filter; the residue was dried in a vacuum. Two
size classes were used; one comprised im-

BIOLOGY AND ECOLOGY OF HELICELLA PAPPI 3

2
N/2500 ст

30

egglaying

0
A86M J J O М DM87A M JJ

ег аут?
egglaying

О NM88A M JJ N DM89M

Months

FIG. 2. Density of Helicella (Xerothracia) pappi (number of snails/2500 ст? (mean + SD)) at Philippi from
April 1986 to May 1989. Double lines on the x-axis indicate periods during which no samples were taken.

mature snails (4 < D < 13 mm) with shell or-
ganic matter containing 1.4% of the total
shell weight, and the other comprised mature
snails (14 < D < 21 mm) with shell organic
matter containing 0.19% of the total shell
weight. These size classes were chosen as a
result of study of the maturation of the geni-
talia and gonad. The shell organic matter was
then added to the dry body weight of the
different size classes and the sum was used
(Table 3) in estimating annual secondary pro-
duction.

RESULTS
Aspects of the Biology of Helicella pappi

Helicella pappi is an iteroparous species
with overlapping generations throughout the
year. Rate of growth and reproduction were
not constant during the three study years
(Figs. 2, 4). The greatest shell diameter of
sexually mature snails was 22 mm, and ma-
turity was attained 21 months after hatching
(Figs. 3, 4) for 98% of the population. The
aperture lip started to form 23 to 24 months
after hatching. In the third year, the other 2%
of the population matured. Egg laying took

place at the end of September, October or
November (Fig. 2), depending on the prevail-
ing climatic conditions, especially precipita-
tion (Fig. 1) Measurements of ten egg
clutches in the field showed that the mean
number of eggs laid was 69 + 4.3 (т + SE),
with a range of 42 to 85 eggs, and the mean
weight of 70 eggs was 0.03 + 0.004 g. The
mean number of eggs laid by older adults,
which had already laid eggs in the previous
year, was 22 + 1.9 (m + SE) (N = 5), with a
range of 18 to 28 eggs. Hatching took place
25 days to one month later, but hatchlings
remained in the soil. The greatest shell diam-
eter of the newly hatched snails was 1.5 +
0.09 mm. By marking ten different egg
clutches, hatching and after-hatching losses
were estimated at 55%. After-hatching
losses were estimated in the laboratory to be
20%. During winter, from the end of Novem-
ber to the end of February, the snails did not
really hibernate. Juveniles formed a thin,
transparent epiphragm, whereas 30% of the
snails with 14 > D > 7 formed a thick epi-
phragm. Snails with D > 14 mm did not form
an epiphragm, but they diapaused under
creeping plants occurring on the soil or
stones. No real aestivation took place. During
July and August, all snails diapaused during

4 LAZARIDOU-DIMITRIADOU

TABLE 1. Estimation of statistical parameters of the population of Helicella pappi (where a, b
= constants, г = correlation coefficient, N = number of snails examined, logd + 6,4 and
logD + O\go = Means of the greatest shell diameter (mm)(D) and the peristome diameter

(mm)(d) + SD)).

Entire sample
ato, 1.189 + 0.005
DEBIGE 0.302 + 0.003
e 0.954
1099 + Goga 0.603 + 0.153
logD + 6,40 1.019 + 0.182
М 2569

dry weather, as in summer 1986 (Fig. 1), but
only the juveniles formed a thin, transparent
epiphragm. Snails were active during spring
and on humid weather in summer (Fig. 1).
Helicella pappi became mature and the
genitalia became well formed when D > 14
mm. Histologically, the gonad of these snails
showed fully formed spermatozoa during Oc-
tober (the reproductive period), although
oocytes were not fully grown. There was a
positive correlation (r? = 0.918, М = 100, P<
0.001) between (а) the greatest shell diameter
(D) and the corresponding dry body weight
and (b) the dry body weight and the dry shell
weight (r° = 0.742, N = 100, Р < 0.001).

Population Dynamics and Spatial
Distribution

The population fluctuated during the study
period (Fig. 2). The mean population density
was 18.1 + 3.3 snails/0.25 m° (mean + SE) in
1986, 14.4 + 6.6 in 1987, and 12.7 + 1.5 in
1988. An ANOVA test among the three con-
secutive years did not show any statistical
differences (F = 1.646, P = 0.2077) in the
population densities. Densities of H. рарр!
peaked in early spring. The population den-
sity was above the mean density for 5-6
months (Fig. 2). The spatial distribution of H.
рарр! was found to be contagious because
parameter b of Taylor's power law was equal
fo) 21043 [© ОТ

Demographic Analysis of the Population of
Helicella рарр!

The analysis of size frequency histograms
(Fig. 3, 4) with probability paper showed that:
(a) three cohorts existed in the habitat
throughout the year; a fourth was added after
the reproductive period, though the third co-
hort contained adults of different ages (Fig.

Juveniles 2<D<14

Adults 14<D<22

1.254 + 0.088 0.859 + 0.012
0.270 + 0.005 0.545 + 0.009
0.922 0.801
0.515 + 0.153 0.75 + 0.057
0.916 + 0.122 1.19 + 0.049
1600 969

4); (b) increased growth rate was observed
during spring. An ANOVA test showed no
statistical differences in the daily rate of
spring growth among the three consecutive
years of study (Fig. 4); (c) mature snails of 15
mm attained their maximum size 21 months
after hatching; (d) when egg laying took place
in October (Fig. 2: 1987), depending on the
degree of precipitation (Fig. 1), the hatchlings
appeared in November (Fig. 3: 27-11-87).
When egg laying took place in November, the
hatchlings appeared in early spring (Fig. 3:
20-3-88 and 31-3-89), possibly because it 1$
safer for thin-shelled juveniles to stay buried
in the soil.

Relative Growth of D in Relation to d

In the field, growth rate was high during
periods of suitable weather, but there was no
significant growth during winter and summer.
Growth was most rapid in spring, from March
till May (Fig. 4). There was a positive correla-
tion between greatest shell diameter and peri-
stome diameter for the whole population of
Н. рарр! (Table 1). Growth rate was faster in
juveniles (a = 1.254) than in adults (a = 0.859),
and growth was more heterogenous in juve-
niles than in adults, because their standard
deviations of the mean were greater (Table 1).
According to Mayrat's (1965a, b) method, the
intersection point between the rate growth of
these two subpopulations (juveniles + adults)
occurred at D = 13.9 mm. This, according to
histological examination of gonads in relation
to the age of the snails, was the diameter at
which spermatozoa were fully grown.

Absolute Growth
The growth pattern of H. рарр! seems to

conform to the equation D, = D,
[1-е ^“ °], which was given by D... =

BIOLOGY AND ECOLOGY OF HELICELLA PAPPI 5

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0 0 0 0
1 10 20 1 7 14 21 1 9 19 1 9 19
207 41286 = 30/9/87 1 26/7/88
8 10
10 6
4 5
2
0 0 0
1 10 20 1 7 14 21 1 9 19
Dmm

FIG. 3. Size-frequency histograms of the population of Helicella pappi at Philippi from April 1986 to May

1989.

28.8[1—е %-0*8+0,313]. According to this
method, the greatest shell diameter mea-
sured in the field (inferred from dead shells (D
= 25.6 mm)) may be attained in 48 months.
The above equation was calculated by using

the known greatest shell diameter in the field
(26 mm), the mean D value of newly hatched
snails, which was 1.2 mm, and the mean size
of young snails first sampled in the field,
which was 1.5 mm (Fig. 3: November 1987).

6 LAZARIDOU-DIMITRIADOU

АЗ © М D MESA М т

Аз SOsM Ти

$ О ММ: 8 А МО Л ВА $

Months

FIG. 4. Мода! distribution of Helicella рарр! at Philippi from April 1986 to May 1989. Broken lines indicate
periods during which no samples were taken. G,-G, indicate the different generations during the study
period. Time breaks denote winter time, during which diapause took place and no samples were taken.

Dt is the shell diameter at time t; D,,,., is the
diameter at the upper growth asymptote cal-
culated according to Ford-Walford equation
(Walford, 1946); t is time in months, t, is the
hypothetical time when D is equal to zero (mi-
nus the egg stage for this paper), and k is the
growth rate coefficient.

Life and Fertility Table

For the construction of the fertility table,
we used (a) the numbers of eggs laid by two
or three-year old adults, and by adults of
more than three years old, known from field
observations and (b) egg-hatching and after-
hatching losses calculated in the field and the
laboratory respectively.

From Table 2, the following may be con-
cluded: (a) mortality rate (Kx) was high after
hatching and then stabilized. It increased af-
ter the first winter, before the second winter
(in 3-year-old adults snails) and stayed high
after the third winter, (b) values of expecta-
tion of life (ex) decreased with increasing age,
(c) the value of net reproductive rate (Ro) was
high (Ro = 3.024), and (d) the per capita rate
of increase was greater than zero, with a rate
of 0.04 per unit of time.

Annual Secondary Production

The calculations for Hynes’ size-frequency
method are listed in Table 3. The mean bio-
mass of each size class was expressed as
dry-weight of body plus organic material of
the shell. After conversion, using Benke's
(1979) correction, annual production (P) was
found to be 5.82 + 0.45 g/(m?-yr) in 1987 and
3.73 + 0.31 g/(m*-yr) in 1988. Biomass (В)
was 2.89 g/m” т 1987 and 1.81g/m° in 1988.
The annual productivity rate constant (P/B)
was 0.21 in both 1987 and in 1988. Turnover
time (B/P x 365 days) was 1765 days in 1987
and 1775 days in 1988.

DISCUSSION

The contagious spatial distribution of H.
рарр! is similar to the xerothermophilic spe-
cies living in similar habitats in Greece (e.g.,
Xeropicta arenosa and Cernuella virgata (Laz-
aridou-Dimitriadou 8 Kattoulas, 1985).

Recruitment of newly hatched snails of H.
pappi seems to be the main reason for the
rise in population density after winter. The
low values of population density in November
and December are due to the fact that some

BIOLOGY AND ECOLOGY OF HELICELLA РАРР!

TABLE 2. Life and fertility table of a cohort of Helicella (Xerothracia) pappi starting in April
1986 (Figure 3-G:).

Age
(months) ly Q, К, e, m, Em;
1 1000 0.27 0.31 12.29 0.00 0.00
2 730 0.00 0.00 15.65 0.00 0.00
3 730 0.00 0.00 14.65 0.00 0.00
4 730 0.00 0.00 13.65 0.00 0.00
5 730 0.18 0.20 12.65 0.00 0.00
6 600 0.00 0.00 14.28 0.00 0.00
ih 600 0.06 0.06 13.28 0.00 0.00
8 565 0.00 0.00 13.07 0.00 0.00
9 565 0.00 0.00 12.07 0.00 0.00
10 565 0.01 0.01 11.07 0.00 0.00
11 560 0.00 0.00 10.14 0.00 0.00
12 560 0.39 0.47 9.14 0.00 0.00
13 350 0.00 0.00 13.32 0.00 0.00
14 350 0.00 0.00 12:32 0.00 0.00
15 350 0.00 0.00 11.32 0.00 0.00
16 350 0.00 0.00 10.32 0.00 0.00
17 350 0.00 0.00 9.32 0.00 0.00
18 350 0.00 0.00 8.32 0.00 0.00
19 350 0.01 0.01 7.32 0.00 0.00
20 347 0.63 0.98 6.39 0.00 0.00
2] 130 0.00 0.00 15.25 0.00 0.00
22 130 0.00 0.00 14.25 0.00 0.00
23 130 0.00 0.00 13.25 18.00 53.75
24 130 0.14 0.15 12.25 0.00 0.00
25 112 0.00 0.00 13.09 0.00 0.00
26 112 0.00 0.00 12.09 0.00 0.00
27 112 0.00 0.00 11.09 0.00 0.00
28 112 0.03 0.03 10.08 0.00 0.00
29 110 0.00 0.00 9.40 0.00 0.00
30 110 0.00 0.00 8.40 0.00 0.00
31 110 0.00 0.00 7.40 0.00 0.00
32 110 0.10 0.10 6.40 0.00 0.00
33 101 0.00 0.00 6.03 0.00 0.00
34 101 0.00 0.00 5.03 0.00 0.00
35 101 0.07 0.07 4.03 7.00 24.07
36 90 0.15 0.17 3.30 0.00 0.00
37. 77 0.27 0.32 2.81 0.00 0.00
38 56 0.19 0.21 2.67 0.00 0.00
39 46 0.38 0.49 2.18 0.00 0.00
40 28 0.38 0.47 2:22 0.00 0.00
41 18 0.40 0.51 2.25 0.00 0.00
42 10 0.33 0.41 2.43 0.00 0.00
43 7 0.00 0.00 2.40 0.00 0.00
44 7 0.50 0.69 1.40 0.00 0.00
45 4 0.50 0.69 1.30 0.00 0.00
46 2 0.60 0.92 1.10 0.00 0.00
47 1 0.50 0.69 1.00 0.00 0.00
48 0 1.00 0.00 0.50 0.00 0.00

Во = УЁ т, = 3.025 г = InRo / Тс = 0.04

|, : Number of animals surviving at the beginning of age-class x (months) out of 1,000 originally hatched.
а, : Mortality rate during age interval x (d,/l,, where d, is the number of animals during age interval x).
K, : Intensity or rate of mortality: loga, — loga
e, : Expectation of life: T,/l, where T, =L, + Ly; ...... L, (L, : is the number of animals alive between age
x and х+1: (I, + 1, 1)/2; L, is the total number of animals x age units beyond age x).

L,m, : Total number of hatchlings in each age interval (m, : Number of living animals hatched per adult
snail).

where Ro is net reproductive rate, г is per capita rate of increase, and Tc is generation time (25,7 months).

х— 1

8

LAZARIDOU-DIMITRIADOU

TABLE 3. Calculation of production of Helicella pappi by the size-frequency method. Annual production
based on sets of samples from April 1988 to April 1989 (where п, = number of snails at the size class j
in number; Un, = variance of п; W, = mean individual dry body weight + mean dry shell of organic
matter (in mg); G, = geometric mean of weight of pairs of successive size classes; В = mean standing
crop or population biomass in mg; P = annual production in mg; P/B = annual turnover ratio; a =
number of size classes; CPI = cohort production interval: 730 days).

Class п, W, (mg)
range п, /0.25т? Un; Di + st. error
1-2 0.01 0.0000 —0.13 0.410 + 0.01
2-3 0.15 0.0020 —0.35 0.410 + 0.02
3— 0.50 0.0554 —0.22 1.000 + 0.02
4-5 0.72 0.1764 —0.05 1.000 + 0.03
5-6 0.77 0.0806 —0.25 2.000 + 0.14
6-7 1.02 0.0960 —0.46 5.000 + 0.27
7-8 1.48 0.2454 012 7.000 + 1.00
8-9 1.36 0.2364 0.37 8.000 + 0.44
9-10 0.99 0.0892 0.04 11.000 + 1.00
10-11 0.95 0.0828 0.39 15.000 + 1.00
11-12 0.56 0.0493 —0.06 22.000 + 3.00
12-13 0.61 0.0436 0.07 26.000 + 2.00
13-14 0.54 0.0483 —0.15 27.000 + 4.00
14-15 0.70 0.0616 —0.23 36.000 + 2.00
15-16 0.93 0.0980 0.10 49.000 + 2.00
16-17 0.83 0.0677 —0.07 51.000 + 2.00
17-18 0.90 0.0736 0.37 65.000 + 4.00
18-19 0.53 0.0427 0.37 71.000 + 5.00
19-20 0.16 0.0058 0.15 85.000 + 5.50
20-21 0.01 0.0003 0.01 134.000 + 6.00
13.73 x 4 =
54.92/m? (259 days)
77.39/m* (365 days)

(B) Р’
AC, [njWj] (пп. 1)(@,)
(WW.,,)°° (mg/0.25 m°) (mg/0.25 m°)
0.41 0.0051 —0.0549
0.64 0.0600 —0.2258
1.00 0.4990 = 0.2233
1.41 0.7223 —0.0738
3.16 1.5489 —0.7781
5.92 5.1025 —2.7369
7.48 10.3819 0.9115
9.38 10.8906 3.5145
12.85 10.8534 0.5313
181 14.1796 7.0363
23.92 12.2754 —1.3548
26.50 15.9802 1.9403
31.18 14.6175 —4.7975
42.00 25.0296 —9.7470
49.99 45.4396 4.9246
57.58 42.2702 —4.2654
67.93 58.6892 25.1060
77.69 37.8676 28.9068
106.72 13.7058 16.0344
134.00 1.4744 1.4744
321.593 x 4 = 66.1227 x 4 =

1286.36 mg/m?” (259d) 264.49 mg/m? (259 а)
1812.82 (365 а) 372.74 (365 d)

P = 20 x 365/730 x 372.74 = 3727.4 mg/(m?.yr) or ог 3,727 g/(m?.yr)

U(P) = Uñ(G,_G,_,) x a? x (365/730) = 24371.67
¿UPS = (22371.67)? = 312.22 = 0:31

P = 3.73 + 0.31 g/(m°.yr)

Р/В = 372.74/1812.82 = 0.206

Turnover time = B/P x 365 = 1775.2 days

snails are already dormant because of the
prevailing weather conditions and the fact
that old adults and some new adults (D > 15
mm) die after egg laying. The population dy-
namics suggests that there is a characteristic
annual periodicity, with synchronization of
population and life-cycle development. This
appears to be a species adapted to recover
slowly after an adverse period. The slow re-
covery of the population results from the low
hatch-rate of eggs deposited just before the
adverse period. The low rate of juvenile de-
velopment into adult stage during spring, and
the rapid decline in population size after the
density peak, indicate a reduced reproduc-
tive effort of later adult stages.

Increased growth took place during spring
because temperatures were not exceedingly
high (20°C), and total monthly precipitation

did not fall below 20-30 mm (Fig. 1). During
autumn, growth took place only in juveniles
(Fig. 4) when temperatures were around 15°С
(in October and November) and only if there
was precipitation. Newly hatched snails dur-
ing that period of the year remained dormant.
The rate of growth, however, was not the
same for newly hatched snails and juveniles
(Fig. 4). This is also related to differences in
temperature [e.g., the 1987 March (5.8°) and
April (12.5%) temperatures were lower than in
1986 (8.6 — 15.4”C respectively)]. Addition-
ally, the growth rate was not the same for
juveniles and mature snails. This was evident
from the study of the population analysis of
H. pappi and from the comparison of the rate
of relative growth of D in relation to d be-
tween juveniles and adults. This is a general
phenomenon in many Helicidae and it is usu-

BIOLOGY AND ECOLOGY OF HELICELLA PAPPI 9

ally due to internal changes in genitalia and
gonad maturation (Yom-Tov, 1971; Bonavita,
1972; Williamson, 1976; Lazaridou-Dimitria-
dou, 1986; Staikou et al., 1988). Seasonal
variation in growth and, more specifically, in-
creased growth rate in spring, have been re-
ported for other snails in Greece, including Е.
vermiculata (Lazaridou-Dimitriadou & Kattou-
las, 1985), X. arenosa and C. virgata (Lazari-
dou-Dimitriadou, 1986), H. lucorum (Staikou
et al., 1988), B. fruticum (Staikou et al., 1990),
and М. cartusiana (Staikou & Lazaridou-Dim-
itriadou, 1990). Baba (1985) also reports that
growth is related to climatic factors and that
it increases only before sexual maturity. As
for H. lucorum (Staikou et al., 1988), B. fruti-
cum (Staikou et al., 1990) and other terrestrial
snails, H. pappi continues to increase D even
after maturation, because it is heterothermic.
There were adverse periods during which ac-
tivity, and consequently growth and repro-
duction, stopped. These periods coincided
mainly with winter and summer drought.
Moreover, the values of k showed that winter
constitutes the most important environmen-
tal stress. Helicella pappi matures only in the
second year of its life, with a rate of increase
equal to 0.041. Being iteroparous, this snail
reproduces again after its first egglaying.
However, it seems that high reproductive
output on one occasion influences future re-
productive output. Although Ro and the turn-
over time was high (1,775 days), productivity
rate (0.2/year) was low, which may be related
to the long life span (4 years) and high mor-
tality of this species at subadult and adult
stages, especially after winter. Moreover,
maintenance costs during dormant periods
are increased; 33% of the previously ac-
quired biomass 1$ lost. These snails try to ex-
ploit favourable conditions, but the absence
of a true hibernation or aestivation and a var-
ied capacity for dormancy cause the death of
much of the population. The annual adult
mortality rate was similar to that reported by
Osterhoff (1977) and Williamson et al. (1977)
for С. nemoralis and by Shachak et al. (1975)
for Sphincterochila zonata, which also has a
life-span of 4-6 years. Turnover times are ob-
viously related to length of life; long turnover
times have only been reported for such bi-
valves as Anodonta (1,789 days/4.9 years)
(Russell-Hunter & Buckley, 1983) and for the
terrestrial snail Monacha (1,177 days/2-3
years) (Staikou 8 Lazaridou-Dimitriadou,
1990). Otherwise, terrestrial snails seem to
have short turnover times from 50.7 days

(Vallonia, Russell-Hunter 8 Buckley, 1983) to
293 days/3 years (Helix lucorum, Staikou et
al., 1988). However, the possibility of long
turnover times in age-structured populations,
especially in relation to interspecific compar-
isons, must be treated with caution, because
some average standing-crop values in the lit-
erature are estimated from an entire popula-
tion and others from a model cohort. Gener-
ally, turnover times of more than two years
appear to be associated with life spans of
four years or more. The fact that H. pappi has
long turnover times and a low productivity
rate may be related to its small size, its long
life span, and the fact that it is iteroparous.

Published distribution data seem to sug-
gest that H. pappi comprises a species with a
patchy distribution (Schútt, 1962). Because of
its evolutionary origin in colder climates (Ur-
banski, 1960), this species manages barely
to survive in this region of Greece, with wide
daily and seasonal temperature variations.
Actually, it is the climatic factors that play an
important role in controlling energy flux in H.
рарр!. Low rates and efficiencies of growth
place severe restraints on the snail's ability to
meet the demands of over-winter mainte-
nance and reproduction. As a result, repro-
duction is delayed until the second autumn.
Energy investment is not concentrated on egg
production, and there 1$ variation in rates of
growth and fecundity according to age. Mor-
tality takes place at all stages, but mainly after
winter; high mortality after adverse periods of
the year is a common characteristic of many
helicid snails. Consequently, the demograph-
ic characteristics of Helicella conform to an
A-selectionist's strategy, as defined by Green-
slade (1983); that is, suitable conditions for
breeding last for only a short period but occur
regularly and predictably, such that the pop-
ulation synchronizes with those conditions
(Figs. 1, 2). Moreover, interspecific competi-
tion is rare, because only very small, sparse
populations of Lindolhomia lens and Helix
figulina were observed.

ACKNOWLEDGEMENTS

| would like to thank Dr. E. Gittenberger of
The Natural Museum of Leiden for informa-
tion on the distribution and systematic posi-
tion of Helicella (Xerothracia) pappi. Thanks
are also due to Dr. С. В. J. Dussart from
Christ Church College, Canterbury, U.K, for
his critical remarks, K. Asmi and Dr. A.

10 LAZARIDOU-DIMITRIADOU

Staikou for their technical help, and Dr. T.
Sofianidou for providing me with samples
from the area of Philippi before this study
was undertaken.

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Revised ms. accepted 28 November 1994

Be, >| i |
| o > = 2. 8
ME =
ny Shane ae ecu
414 EN ibe “abr x my pe
LUS

| de к” |

MALACOLOGIA, 1995, 37(1): 13-21

EGG-LAYING AND ASSOCIATED BEHAVIOURAL RESPONSES OF
LYMNAEA PEREGRA (MULLER) AND LYMNAEA STAGNALIS (L.)
TO CALCIUM IN THEIR ENVIRONMENT

Hilary Piggott & Georges Dussart

Canterbury Christ Church College, Canterbury, Kent CT1 1QU, United Kingdom

ABSTRACT

In laboratory trials, specimens of the freshwater pulmonate snail Lymnaea peregra from
Ullswater, a soft-water lake in the English Lake District (6.5 mg/l [Са?*], pH 7.1), showed a
significant preference for laying eggs on clean, dead conspecific snail shells (y? = 38.91, Р <
0.001 n = 20), thereby confirming anecdotal field observations of this behaviour.

A choice chamber was used to investigate to test the hypothesis that Iymnaeid snails might
be able to use calcium as a cue for orientation, for example as a stimulus to find target shells
for oviposition. Lymnaea stagnalis showed a more positive response to calcium than did L.
peregra (t = 4.2, Р < 0.05, п = 137). When snails from soft-water environments were reared in
soft water in the laboratory, specimens of L. peregra showed a strong preference for calcium
(x? = 19.6, Р < 0.001, п = 202), but specimens reared in hard water (84 mg/l [Ca?*], п = 44))
showed no such preference. The hypothesis that snails could use a calcium cue to select a shell
as an oviposition site was supported, and, in addition, the breadth of the chemical niche of L.

peregra appears to be wider than that of L. stagnalis.
Key words: egg-laying, calcium, preference, ecology, hardness, Lymnaea peregra, Lymnaea

stagnalis.

INTRODUCTION

The importance of calcium in the distribu-
tion of freshwater molluscs has been widely
reported (Boycott, 1936; Macan, 1950;
Okland, 1969; Williams, 1970; McKillop 8
Harrison, 1972). In his qualitative study of the
ecology of freshwater molluscs in Britain,
Boycott suggested that whereas some spe-
cies were restricted to waters with calcium
concentration exceeding 20mg/l, other spe-
cies were more tolerant and could occur both
in high calcium waters and elsewhere in low
calcium waters, L. peregra being a typical ex-
ample of such tolerance. Boycott (1936) also
highlighted the difficulty of separating water
chemistry from geographical distribution and
physical characters of the habitats. In quan-
titative studies of molluscan ecology in rela-
tion to water chemistry, Dussart (1976)
showed that general mollusc abundance was
greater in hard waters ([Ca*] > 40 mg/l), al-
though medium waters ([Са?*] 5-40 mg/l)
had greater species diversity. As well as af-
fecting distribution, calcium concentration
also affects such aspects of freshwater mol-
lusc biology as shell composition in planorbid
snails (Madsen, 1987), freshwater sphaeriid
clams (Burky et al., 1979), and the ampullariid

13

snail Marisa cornuarietis (L.) (Meier-Brook,
1978). Other evidence of a direct metabolic
response to environmental calcium was
demonstrated by Dussart & Kay (1980), who
showed that L. peregra reared in waters of
different hardness had different respiration
rates.

Dussart (1979) observed anecdotally that
in some soft-water habitats, L. peregra ap-
peared to show a preference for laying eggs
on the shells of other L. peregra from the
same generation, and it was proposed that
this behaviour could possibly *'... provide an
immediate source of nutrients to the off-
spring after the post-egg laying deaths of the
parent population.” These observations
therefore raised questions about whether
snails could behaviourally orientate to envi-
ronmental calcium. Thomas (1982) observed
that taxes of freshwater molluscs along inor-
ganic ion gradients had been little studied.
There is precedent for suggesting that fresh-
water molluscs exhibit chemoreception,
because, for example, the freshwater pulmo-
nate Planobarius corneus (L.) can discrimi-
nate between amino acids (Lombardo et al.,
1991), and Thomas et al. (1980) observed
species-specific responses to four amino
acids.

14 PIGGOTT & DUSSART

Lymnaea peregra 1$ probably the most
common freshwater snail in Europe (Fitter 8
Manuel, 1986); it occurs in all types of habi-
tats, including water with a calcium concen-
tration as low as 1 mg/l. By contrast, L. stag-
nalis is a calciphile species comprising larger
individuals, often sympatric with L. peregra in
harder waters. The objectives of the following
experiments were therefore, firstly to investi-
gate the hypothesis that soft-water snails
might lay eggs preferentially on the shells of
other individuals, secondly to investigate
whether L. peregra and L. stagnalis might ori-
entate to calcium in solution and thirdly, to
identify any species-specific differences in
such behaviour.

MATERIALS AND METHODS

Sources and Maintenance Conditions
of Snails

All samples were taken in June and July
1992, when L. peregra were obtained from
the River Stour in Kent (OS ref. TR1785990,
84 mg/l [Са?*], pH 7.4) and L. stagnalis were
collected from Monkton Nature Reserve (OS
ref. TR657295, 72mg/l [Са?*], pH 7.5). Spec-
imens of L. peregra were also collected from
Ullswater, a soft-water lake in the Lake Dis-
trict (OS ref. NY421205, 6.5 mg/l [Ca**], pH
7.1). In all the choice experiments, snails
were young-mature specimens, of about
8-11 mm maximum length for L. peregra and
25-35 mm maximum length for L. stagnalis;
because some of the snails came from the
field, age could only be estimated. Other-
wise, snails were four-six months old.

Snails were housed in the laboratory in
plastic tubs measuring 16 x 16 x 16 cm, each
tub containing one litre of water and ten
snails. Where snails from hard water were
being cultured in hard water, water from the
River Stour was used. The snails were main-
tained at room temperature, with continuous
aereation, in natural daylight and fed fresh
lettuce every three days; ten millilitres of fil-
tered pond water were added to each tub to
provide additional micro-nutrients. A small
quantity of washed, fine sand was provided
in each tub to aid digestion, and the water
was changed at three-day intervals. In the
experiment to investigate substratum choice,
two round, flint pebbles (10 mm diameter, the
approximate size of an L. peregra shell), and
two empty L. peregra shells were added to

each tub. Each tub thus comprised a micro-
cosm for which both the number of egg cap-
sules and the area of the surface types upon
which eggs were laid was recorded, includ-
ing the submerged walls of the tub.

Review of Choice-Chamber Designs for
Aquatic Snails

Various experimental designs have been
employed to examine chemoreception in
aquatic molluscs. For example, Uhazy et al.
(1978) investigated chemicals attractant to
Biomphalaria glabrata (Say) by using a grid of
10 x 5 units marked on a white enamel dish
with test material at one end of the dish and
control material at the other; Madsen (1992)
used a similar design in food location exper-
iments for Helisoma duryi (Wetherby) and Bi-
omphalaria camerunensis (Boettger). Lom-
bardo et al. (1990) used a ‘Y’ shaped maze to
investigate Planorbarius corneus, whereas
Thomas et al. (1980) used an olefactometer
to investigate the response of B. glabrata to
amino acids and related compounas.

There are however, many problems asso-
ciated with the design of such experiments;
firstly, diffusion causes dynamic change in
ionic concentrations, so that snails might be
responding to an ionic flow, as opposed to
absolute concentrations. Secondly, the ani-
mals might become satiated as they move up
the concentration gradient so that their be-
haviour changes accordingly. Thirdly, there
are problems of deciding when a choice has
been made; for example, Lombardo et al.
(1991) reported that a time limit was neces-
sary due to the slowness and sometimes in-
activity of the snails in their experiments.
Fourthly, the distinction between olfaction
and gustation is not clear in aquatic molluscs
(Kohn, 1961). Fifthly, when a liquid medium
rather than a solid object is the object of the
choice, there are problems of experimental
design because seiche phenomena can be
entrained at the start of each trial. Given the
nature of the animal, it is unlikely that any
aquatic choice-chamber design will be
wholly satisfactory for aquatic snails.

To circumvent these problems, Vareille-
Morel (1986) used a flowing water design in
studies of the response of Potamopyrgus
jenkinsi Smith to a nutrient source, and Dus-
sart (1973) used a static chamber with a
“starting box”; test specimens introduced
into the starting box had an instant choice of
waters when the starting gate was removed.

RESPONSE OF LYMNAEA TO ENVIRONMENTAL CALCIUM 15

Central baffle

Water flow

os

FIG. 1. Choice chamber used in the present study.

Design of the Choice-Chamber

Because L. stagnalis and L. peregra can be
amphibious, the choice-chamber shown in
Figure 1 was used, as it circumvents many of
the problems described above. Preliminary
dye experiments and water samples taken
during the trials showed that there was no
significant transfer of chemicals across the
central divider; the latter was made of wood
(ramin) because streaming was found to oc-
cur when plastic dividers were used.

Character of Water

For all choice experiments, solutes were
added to distilled de-ionised water to give
the following standard water (mg/l): magne-
sium, 15; sodium, 30; potassium, 7; ortho-
phosphate, 15; calcium, 0; pH 7.5; (pH ad-
justment with 2M НС). By contrast, the test
hard water included 50 mg/l calcium.

Methods of Testing Preferences

Testing one species at a time, up to ten
snails were placed on the divider, all facing
the same way. They could then move to the
left or right, or could remain on the divider. To
prevent extraneous bias, the location of
choice waters were systematically varied be-
tween runs. Because Green et al. (1992) had
shown significant anticlockwise-left move-
ment of L. peregra when out of water, runs
were repeated, with all the snails facing in the
opposite direction. There were thus four ori-
entations in each run and each run was re-
peated, so that approximately 80 snails were
involved in each trial. Similar numbers of
snails were used by Madsen (1992) in food
location experiments. After ten minutes, or
sooner if all the snails had moved, the snails
were removed from the choice chamber and
the dividers cleaned; there was no contact

between snails and no evidence of trail fol-
lowing.

y? tests with Yate's corrections were used
to see if there was a significant choice and
snails which stayed on the central divider
were not included in the analysis. For the final
investigation, L. peregra reared from eggs
produced by Ullswater snails were used. Two
batches of adult snails were separately main-
tained in soft water (5 mg/l [Са?*]), and two
batches were separately maintained in hard
water (84 mg/l [Са?*]), both being kept for
1-3 weeks before the trials.

RESULTS
Oviposition Choice of L. peregra

Besides the walls of the tub, the snails had
only two pebbles and two shells upon which
to lay, so that oviposition on these surfaces
needed a significant positive choice. After
correction of the data for available area, soft-
water snails imported from Ullswater showed
a significant bias towards laying eggs on
shells, and this bias was greater for snails
which, before the trials, had been kept in the
laboratory in soft water (y? = 38.91, P <
0.001) compared with those kept in hard wa-
ter (X? = 13.49, Р < 0.01). These results imply
that L. peregra from soft water showed a
preference for laying eggs on snail shells,
particularly having been maintained in soft
water. It is a statistical requirement of a chi-
square test that no observed values should
be less than five; because no eggs were laid
on the pebbles, this requirement was not
met, and the results should therefore be
treated with caution; it is conceivable that
eggs might never have been laid on a flint
pebble. It should be noted that in other batch
cultures however, eggs were intermittently
laid on such pebbles.

Choice Experiments

A number of preliminary tests were con-
ducted. Ten L. stagnalis were housed in soft
water and used in the following preliminary
experiments. When given a choice of sodium
and calcium cations but with the same con-
centrations of chloride anions, the null hy-
pothesis could be refuted at P < 0.05, imply-
ing that movement was significantly towards
the source of calcium (Table 1-Trial (1). To
determine whether this apparent bias was

16

PIGGOTT & DUSSART

TABLE 1. Preliminary investigation of choice behaviour. For the purposes of this report, a trial is defined
as an opportunity to make a choice by a single individual snail. Because some snails stayed on the
central baffle for the duration of an experiment, the number of movements 15 frequently less than the
number of trials. In experiments (i)-(iv), each of the ten snails was tested four times (i.e. four runs) in

the four orientations, giving 160 trials.

Preliminary trials—ten mature specimens of L. stagnalis previously housed in soft water for one week

Bias Towards.. vo Significance
(i) Comparison of cations
100 mg/l [CI ] cf 100 mg/l [CI ]
(as NaCl) (as Ca Cl,.2H,0)
Movements 81 56 [Cas] 4.2 Р < 0.05
(ii) Sodium against water
100 mg/l [Na*] ai HEC
(as NaCl)
Movements 73 50 [Na*] 3.9 Р < 0.05
(iii) Calcium against water
100 mg/l [Са?*] cf H,0*
(as Ca CI,.2H,0)
Movements 89 53 [Gaz] 8.8 Р < 0.01

Osmotic Potential Trial —40 mature specimens of L. stagnalis previously housed in soft water for one
week were tested in each of the four orientations, ¡.e. 160 trials.

(iv) [Ca?*] cf [Na*]
500 mg/l 370 mg/l
(аз Ca CI,.2H,0) (as NaCl)
Movements 92 45

prompted by an aversion for sodium, the
snails were given a choice between sodium
and de-ionised water in Trial (ii). Again, the
null hypothesis could be refuted at P < 0.05,
implying that the snails chose the sodium re-
gime and aversion had not been a factor in
Trial (i).

In Trial (iii), there was again an apparent
bias towards calcium, implying that the null
hypothesis could be refuted at P < 0.01; this
could represent an aversion to de-ionised
water, possibly due to the effects of osmotic
potential. Freshwater gastropods have body
fluids that are hyper-osmotic to the external
media and therefore have to cope with
the continual influx of water. To establish
whether the difference in osmotic pressure
between the test substances was affecting
the responses, in Trial (iv), snails were offered
a choice between calcium and sodium but at
concentrations that would each exert the
same osmotic potential. The result provides
evidence to refute the null hypothesis that
there is no significant difference in the re-
sponse of snails to different substances of
the same osmotic pressure at P < 0.01; the
snails apparently once again showed a sig-
nificant bias towards the water containing
calcium.

Bias Towards.. Ne Significance

[Ca?*] 15.4 Р < 0.001

To investigate whether there might be spe-
cies-specific differences in these move-
ments, 40 specimens each of L. stagnalis
from Monkton and L. peregra from the Stour
were maintained in hard and soft water for
periods of one to three weeks before inves-
tigation. The results of the main trials shown
in Table 2 allow comparison between their
behaviour. Trials (v) and (ix) were controls
showing that when exposed to identical test
substances, there was no significant bias in
choice for either species.

For Trials (vi) to (viii), L. stagnalis displayed
a highly significant positive response to cal-
cium, but in Trials (x) to (хи), L. peregra
showed a lower response (Table 2). A Stu-
dent's t-test of these data confirmed this dif-
ference; across the three trials, the mean
number of L. stagnalis choosing calcium was
42.17 with a comparable mean of 32.42 for L.
peregra, thereby indicating that L. stagnalis
were orientating more strongly than L. pere-
gra (t = 4.33). The responses for both species
towards calcium differed, depending on the
length of time they had been maintained in
hard or soft water before the experiment. For
example, the y? value for L. stagnalis that had
been kept in soft water for one week was
62.3, compared with 19.5 for snails that had

RESPONSE OF LYMNAEA TO ENVIRONMENTAL CALCIUM АГ

TABLE 2. Results of trials to investigate choices of forty individuals of each of L. stagnalis and L.
peregra. Snails were kept in either hard water for one week (H1), hard water for two weeks (H2), soft
water for one week (S1) or soft water for three weeks (S3). All calcium was presented as 100 mg/l

calcium as [Ca Cl,.2H,0]. Each snail was tested twice in each of the four orientations, i.e. 320 trials.

Trial Choice Available Result

L. stagnalis Bias Towards.. x? Significance
(v) H1 [Ca?*] cf [Са]

Movements 140 135 neither 0.08 N.S.
(vi) H2 [Са?*] ef H,O*

Movements 155 98 [Са || 12.4 Р < 0.001
(vii) $1 [Ga?*] cf 850%

Movements 186 61 [Cart] 62.3 P < 0.001
(viii) $3 [Ca?*] cf H,O*

Movements 165 93 [Ga?*] 19.5 Р < 0.001

L. регедга
(x) H1 [Gas] cf [Са?*]

Movements 135 139 neither 0.08 N.S.
(x) H2 [Gas] cf H,O*

Movements 124 128 [Ca**] 0 N.S.
(xi) $1 [Са?*] cf H,O*

Movements 132 95 [Са?*] 5.7 Р < 0.05
(xii) $3 [Ca] et 150:

Movements 133 69 Са | 19.6 Р < 0.001

DISCUSSION

been kept in soft water for three weeks; by
contrast, the equivalent y? values for L. per-
egra were 5.7 compared with 19.6.

It is possible that the snails were orientat-
ing to chloride rather than calcium. However,
Trial (x) for L. peregra showed no significant
preference between chloride and de-ionised
water; and, in fact 34 snails compared with
26 snails actually migrated into the de-
ionised water. There is always the possibility,
however, that this experiment shows a bal-
anced preference/aversion for both chloride
and de-ionised water. For example, aversive
behavioural and physiological responses to
salinity are well documented (Perkins, 1974).
The animals may have balanced their aver-
sion to salinity with an aversion to the os-
motic problems posed by deionised water,
and consequently made по significant
choice.

Those L. peregra that had been raised in
hard water showed no calcium preference
(Table 3). By contrast, L. peregra raised in
soft water showed a significant preference
(X? = 8.2, Р < 0.01) for calcium. For example,
over four runs, the mean number of snails
that had been raised in hard water and chose
calcium was 6.5, and the mean number of
snails that had been raised in soft water and
chose calcium was 8.75, the difference be-
tween the means being significant at (t =
2:67).

The molluscan shell is formed by the dep-
osition of calcium carbonate on a protein ma-
trix. For snails in a soft water environment, an
immediate source of calcium could be ben-
eficial to the developing juveniles and so
there is an a priori reason for expecting snails
to be able to detect and orientate towards
calcium. The possible attractant properties of
calcium were demonstrated here by the re-
sults of Trial (i) on [. stagnalis. lt appeared
that sodium chloride did not act as a repel-
lant, because movements away from this
compound were not significant in Trial (ii);
Madsen (1990) found that such snails as H.
duryi and Bulinus truncatus Audouin were not
adversely affected by low concentrations of
sodium chloride. However, there could have
been a significant aversion to the de-ionised
water in Trial (ii).

Young (1975) reported that in soft water, L.
peregra extracted 70% of the calcium re-
quirement from lettuce and in hard water, ex-
tracted only 46% from lettuce. This was
compared with L. stagnalis, which, although
it efficiently extracted 95% of the calcium
content of the lettuce, usually only took 20%
of the total requirement from this source, the
remaining 80% being derived directly from
the water. Thus, the relationships between
environmental Базе-юп concentration and

18 PIGGOTT & DUSSART

TABLE 3. Results for eight L. peregra reared in hard water and eight reared in soft water. The snails
were tested when they had reached an overall shell length of 6-8 mm. Calcium was presented as 100
mg/l [Ca Cl,.2H,0]. Each specimen was tested twice in each of the four orientations, i.e. 64 trials.

Choice Available

Snails reared in soft water

[Са?*] cf
Movements 35
Snails reared in hard water

[Ca?*] cf
Movements 22

snail biology appear to be complicated. For
example, ionic ratios might be involved; Har-
rison et al. (1966) invoked the ratio of cal-
cium/magnesium as a significant factor in
egg production. There are also contrary re-
sults; though Harrison et al. (1966) found a
curvilinear relationship between egg produc-
tion and calcium concentration for В+
omphalaria pfeifferi, Thomas et al. (1974)
found a positive linear relationship for Bi-
omphalaria glabrata. Nevertheless, it ap-
peared that in our experiments, the need to
respond to a source of environmental cal-
cium was less for L. peregra than for L. stag-
nalis, possibly because L. peregra obtains a
smaller proportion of its calcium require-
ments direct from the environment. This
proposition 1$ supported by the results
shown in Table 2: a gradually increasing re-
sponse to a source of calcium by L. peregra,
whereas the corresponding response of L.
stagnalis was immediately highly significant.
Even when kept, albeit temporarily, in hard
water (84 mg/l [Са?*]), it seems that the cal-
cium requirement of L. stagnalis was not be-
ing met in these relatively small containers.
Trials (v) to (xii) were designed to show
species-specific differences in behaviour and
were not particularly designed to distinguish
between the effects of calcium, chloride or
possible osmotic potential effects of de-ion-
ised water. However, the results of trial (x) for
L. peregra suggest that chloride and osmotic
potential were not playing respectively at-
tractive and aversive roles, because a major-
ity of snails chose de-ionised water. There is
also circumstantial evidence from Trials (i) to
(iv) and from the literature to suggest that cal-
cium 1$ a significant factor (e.g., Greenaway,
1971a, b). It is probable that ten L. stagnalis
housed in one litre of pond water would re-
duce the calcium concentration to such a
level that a significant response to a source
of calcium would be both essential for the

H,0*

H,0*

Result
Bias Towards.. x Significance
14 lea] 8.2 P < 0.01
22 neither 0 N.S

snail and observable by the experimenter.
For example, in our experiments, L. stagnalis
showed a massive response to the source of
calcium, having been in soft water for only
one week. Out of 247 observed movements,
186 were towards the source of calcium; if
osmotic potential were the only factor, it
might be expected that the response would
not have changed between trials. In addition,
at this stage, the shells of L. stagnalis were
becoming increasingly fragile. Nduku & Har-
rison (1976) suggested that snails cultured in
low calcium concentration are physiologi-
cally stressed and cannot carry out normal
metabolic processes.

Bielefeld et al. (1993) have implicated alka-
line phosphatase in the mantle epithelium as
a factor in shell mineralisation, and Green-
away (1971a, b) suggested that there was a
net movement of calcium from the environ-
ment into the blood, excess calcium being
deposited as carbonate in the shell. When
Greenaway cultivated snails in soft water,
there was a loss of calcium from the blood to
the external environment. To compensate, a
reverse flow of calcium occurred from the
shell to the blood. It is therefore likely that in
the later stages of our experiments, L. stag-
nalis were physiologically stressed.

Given that the test snails of both species
were of similar sizes, these results suggest
that either L. stagnalis has a greater calcium
demand than L. peregra, or that L. peregra
has a higher threshold for the inception of
calcium-mediated stress at lower concentra-
tions than L. stagnalis. The implication is
therefore that in terms of meeting calcium re-
quirements, the niche breadth of L. peregra is
wider than that of L. stagnalis. This hypothe-
sis 15 supported by Costil-Fleury (1991), who
used factor analysis to show that L. peregra
is less confined in terms of habit type than L.
stagnalis.

However, many other physico-chemical

RESPONSE OF LYMNAEA TO ENVIRONMENTAL CALCIUM 19

and biotic factors are as important as cal-
cium. Temperature, pH, macro-vegetation,
suspended solids and the nature of the
allochthonous input to the habitat have all
been suggested as crucial factors in mollus-
can distribution (e.g., Macan, 1974; Okland,
1983; Pip, 1986), and Dussart (1979) showed
that potassium, mud substratum-type and
rock substratum-type were major variables in
the distribution of Bithynia tentaculata (L.),
Gyraulus albus (Müller) and Planorbis planor-
bis (L.) respectively; magnesium was a major
water chemistry variable for L. peregra.

The results of the investigation using L.
peregra reared in different environmental
conditions tentatively support the suggestion
of Dussart (1979) that in soft waters, snails
might satisfy a metabolic need for calcium by
orientating towards, and laying eggs on
shells. There is anecdotal evidence that, es-
pecially in areas of base-ion deficiency,
aquatic snails will aggregate on shells, bones
and other calcium sources; it would be useful
to compare the calcium responses of snails
originating from soft-water environments
with snails of the same species from hard-
water environments. If raised in identical con-
ditions, any significant differences in re-
sponses persisting in the F1 and succeeding
generations would indicate the existence of
physiological races and plasticity. This in turn
might suggest that the process of speciation
for water type is under way.

The existence of a microhabitat at the sub-
stratum surface of, say, a shell or a pebble
needs to be recognised. Environmental cal-
cium concentration in the microhabitat could
be higher at interfaces due to mineralization
by bacteria, algae or fungi. Consequently, a
variety of factors could act as a cue for ovi-
position on shells, examples being biofilms
(auchwuss) on the shell surface, phero-
mones, amino acids from the shell protein, or
even physical contact with the shell as a sub-
stratum rather than the calcium itself. In the
experiments reported here, it is possible that
the differing physico-chemical nature of the
plastic, pebble and shell surface had facili-
tated the development of biofilms that might
have differentially encouraged egg deposi-
tion on these surfaces; this possibility needs
further investigation. Conversely, if an amino-
acid attractant was leaking from the shell sur-
face, it could be species-specific, because
the concentration of amino acids within
shells differs in different species of mollusc
and with waters of different hardness (Dus-

sart, 1973; 1983). Freshwater snails can
show chemoreception in relation to amino
acids exuded from a food source (Croll,
1983; Thomas et al., 1980), and the parent
snails could be following an amino acid gra-
dient to the target shell. In natural conditions,
the amino acids could act as a primary stim-
ulus and calcium could be secondary. The
behavioural response to the amino acids in
the shell of L. peregra could be examined by
other choice-chamber experiments. How-
ever, the possibility of synergism must be
recognised; for example, Uhazy et al. (1978)
noted a synergistic response of B. glabrata to
the amino acids proline and glutamine.

Despite the fact that Uhazy et al. (1978)
found that B. glabrata would orientate to-
wards magnesium but not calcium, our dem-
onstration of species-specific differences
has implications for control of snail vectors of
helminth disease; for example, species within
the Bulinus and Biomphalaria genera might
show similar variability. As suggested by
Thomas et al. (1980), Thomas (1982) and
Lombardo et al. (1991), baiting techniques
might be used to control helminth vectors,
and knowledge of the role of calcium could
be an important contributing factor.

In conclusion, it seems from this study that
there are differences in orientation behaviour
that might be symptomatic of the calciphile
distribution of L. stagnalis compared with the
eurycalcic distribution of L. peregra. In the
investigation of oviposition-site behaviour of
L. peregra, the bias towards laying eggs on
shells may have been due to such factors as
leakage of calcium or amino acids from the
shells, or mechanical quality of the shell sur-
face. Both species might be able to orientate
towards sources of calcium, though aversion
to a low osmotic potential and attraction to
chlorides are alternative, though less likely
hypotheses. Also, there appears to be a dif-
ference in response within a species when
the animals are raised under different envi-
ronmental conditions, suggesting that there
may be the capacity for physiological plas-
ticity within these metabolic requirements.

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RESPONSE OF [УММАЕА ТО ENVIRONMENTAL CALCIUM 21

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Revised Ms accepted 19 October 1994

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DR ir Ar

MALACOLOGIA, 1995, 37(1): 23-32

ANATOMICAL STUDY ON TONNA GALEA (ИММЕ, 1758) AND
TONNA MACULOSA (DILLWYN, 1817) (MESOGASTROPODA, TONNOIDEA,
TONNIDAE) FROM BRAZILIAN REGION.

Luiz Ricardo Lopes de Simone

Seçäo de Moluscos, Museu de Zoologia da Universidade de Sao Paulo,
Caixa Postal 7172, CEP 01064-970, Sáo Paulo, SP, Brazil

ABSTRACT

Tonna galea and Tonna maculosa, from Brazilian region, are described anatomically. Each
character is compared between the species and also with other known Tonnoidea. These
species differ anatomically in characters of the mantle color and collar, osphradium, hypobran-
chial gland, kidney, proboscis length, radular rachidian, lateral tooth, penis, vas deferens, and
female genital opening. Characters of the anterior region of the digestive system, heart, penis
and pallial oviduct are of particular interest in tonnoidean systematics.

INTRODUCTION

Tonna galea (Linné, 1758) is a very wide-
ranging species, occurring in the Pacific and
Atlantic oceans and in the Mediterranean
Sea. Tonna maculosa (Dillwyn, 1817), in con-
trast, only occurs in the tropical west Atlantic
(Rios, 1985: 70; Matthews et al., 1987: 33).
Tonna perdix (Linné, 1758), which is closely
related to Т. maculosa, occurs in the Indo-
Pacific region, and some authors consider
them to be synonyms (e.g., Morretes, 1949).
Two questions arise: (1) is “Топпа galea” а
single species, and (2) are Т. maculosa and Т.
perdix really separate species? These ques-
tions have been discussed (e.g., Turner,
1948; Matthews et al., 1987), with arguments
based on shell characters, but are still unre-
solved.

The objective of this paper is not to answer
these questions, but to be a step in this di-
rection, providing anatomical descriptions of
specimens identified as Tonna galea and Т.
maculosa from the Brazilian region. These
data could be used in comparisons with sup-
posed co-specific specimens from other ar-
eas to assist in clarifying the systematic
problems. Another objective of this paper is
_ to contribute to the understanding of system-
atic characters in the Tonoidea, identifying
some useful characters not previously uti-
lized in the systematics of this group.

Little has been published on the anatomy
of the Tonnoidea, and in particular the Ton-
nidae. What has been has mainly concerned
the alimentary canal and feeding habits. The

23

following information is available: Tonna
galea: Weber (1927) studied the digestive
system, and some of his data were repro-
duced by Hyman (1967); Turner (1948)
described the penis and figured the radula;
Matthews et al. (1987) figured the penis,
jaw and radular rachidian; and Bentivegna &
Toscano (1991: 37) figured an active speci-
men preying on Holothuria tubulosa and
Н. sanctori. Tonna maculosa: Turner (1948)
described the penis and figured the radula;
and Matthews et al. (1978) figured jaw
and radular rachidian. Other Tonnoidea de-
scribed anatomically and used here for com-
parision are: Reynell (1905) —description of a
male Cassidaria rugosa (Linné) (Ranellidae);
Day (1969) — digestive system of Argobuc-
cinum argus (Gmelin) (Ranellidae); Houbrick
8 Fretter (1969) — digestive system and
other organs of three species of Bursa (Bur-
sidae) and four of Cymatium (Ranellidae);
Lewis (1972) — anatomy of anterior region of
digestive system, head-foot complex and pe-
nis of Distorsio perdistorta Fulton (Ranell-
idae); and Hughes 8 Hughes (1981) — diges-
tive system of Cassis tuberosa (Linné),
(Cassidae).

MATERIAL AND METHODS

The specimens studied belong to malaco-
logical collection of the Museu de Zoologia,
Universidade de Sáo Paulo (MZUSP). They
are preserved in 70% ethanol.

All specimens were dissected using stan-
dard techniques. The buccal region, region of

24 SIMONE

female genital opening and pallial oviduct
were extracted, dehydrated in ethanol series,
stained by carmine, cleared, and fixed with
creosote. Radulae and jaws were examined
on slides with Hoyer. All drawings were made
using a camera lucida.

Anatomical terminology is based on Rey-
пе! (1905) and Hughes & Hughes (1981).
Conchological description and synonymy are
omitted, and can be found mainly in Mat-
thews et al. (1987) and Turner (1948).

Tonna galea (Linné, 1758)
(Figs. 1-19, 37)

Synonymy and types material: Turner
(1948: 173) and Matthews et al. (1987: 31)

Diagnosis

Shell of clear, homogeneous color; outline
globose; sculptured with strong, spiral,
somewhat isometrical ridges. Mantle border
thick; hypobranchial gland poorly developed.
Kidney large, with complex tissue arrange-
ment. Proboscis about half projecting from
rhynchodeum in fixed specimens. Central
cusp of radular rachidian smooth; main cusp
of lateral tooth smooth. Penis with a small
pointed papilla; anterior region of vas defer-
ens fused with seminal receptaculum. Fe-
male genital opening with larger inner divi-
sion for the bursa opening.

Description

Shell: Detailed descriptions of the shell given
by Turner (1948: 173 pl. 78) and Matthews et
al. (1987: 31, figs. 1, 2). Protoconch brown, of
almost four glassy, convex whorls.

Head-Foot Complex (Figs. 1, 2): Foot large,
solid, rounded posteriorly, notched anteri-
orly; propodium narrow, with anterior pedal
gland opening to ventral slit (Fig. 1, mp).
Operculum lacking in adult, present in young
(Rios, 1985). Tentacles long, fairly thick,
bluntly pointed. Black eyes on tubercles on
outer upper part of tentacle bases. Rhyncho-
deum with simple, rounded opening. Probos-
cis with about half of length projecting from
rhynchodeum in all specimens (Fig. 2). Head-
foot structures beige, with dark brown, irreg-
ular spots.

Pallial Complex (Fig. 12): Mantle edge entire,
simple, not reflected, thick, rounded, pale
cream in color. Pallial cavity occuping first

whorl. Siphon long, well developed, pale-
beige, with dark-brown, somewhat longitudi-
nal, irregular spots. Osphradium large, bipec-
tinate, on pallial roof at base of siphon;
osphradium leaflets lamellate, pigmented
brown. Ctenidium very large, monopectinate
(Fig. 12); leaflets very numerous, triangular,
low. Hypobranchial gland not well devel-
oped; some specimens with folds of this
gland running from the anterior and left re-
gion of rectum. Flaccid tissue covering rec-
tum (and pallial oviduct in females) on right
side of mantle cavity, allowing a well-devel-
oped ad-rectal sinus.

Excretory-Circulatory Systems: Kidney very-
large, on right side of pericardium, immedi-
ately behind pallial cavity, from which it is
separated by a thin, nearly transparent mem-
brane (Fig. 12, km); this membrane with slit-
like nephrostome, surrounded by muscular
fibres to form a sphincter (Fig. 12, ne). Kidney
traversed by intestine, which divides it into
two lobes, the largest (Fig. 13) anterior, the
smallest posterior. Internally, the kidney is
very complex, the outer divisions being
formed by green-brown lobes and tubes, its
anterior limit bulging the posterior region of
pallial oviduct glands of females. Nephridial
gland cream-colored, poorly developed, sit-
uated above nephrostome (Fig. 13, ng). Heart
(Fig. 13) with very thin, transparent, flaccid
auricle and a very thick, rounded ventricle
(Fig. 13). Ctenidial vein on right margin of gill,
entering auricle both anteriorly and posteri-
orly (Fig. 13). Central part of auricle inserting
directly in gill margin (Fig. 13, au).

Ad-rectal sinus very developed, apparently
continuous from chamber of kidney into an
aperture, with muscular walls, near the anus
(Fig. 37, at), similar to an ureter.

Digestive System: Similar to that described
by Weber (1927), typical of tonnoideans
(Hughes & Hughes, 1981). Some details of
insertion of proboscis gland duct and oe-
sophagial caecum duct in buccal complex
are shown in Figure 4; detail of buccal com-
plex (sectioned dorsally) shown in Figure 5;
transversal section of the mid region of ante-
rior oesophagus shown in Figure 6. Salivary
glands surrounding duct of proboscis gland
(Fig. 3). Duct of each proboscis gland looping
anteriorly to nerve ring in all specimens ex-
amined (Fig. 11). Radular rachidian tricuspate
(Fig. 7; Matthews et al., 1987); lateral teeth
with a flattened, irregular base and two
cusps, a long, large acuminate cusp and a

ANATOMICAL STUDY ОМ ТОММА GALEA 25

sheath exposed, scale = 10 mm; (3) anterior region of digestive system, dorsal view, scale = 10 mm; (4)
anterior extremity of opened proboscis, right-dorsal view, scale = 1 mm; (5) buccal complex opened
dorsally, inner view, scale = 2 mm; (6) transversal section in mid region of the anterior oesophagus, ventral
region down, scale = 1 тт; (7) rachidian tooth of the radula; (8) lateral tooth showing the accessory cusp
(ac); (9) inner marginal tooth; (10) outer marginal tooth, scale (Figs. 7-10) = 0.5 mm; (11) dorsal view of the
region of nerve ring, scale = 2 mm.

26 SIMONE

FIGS. 12-19: Tonna galea: (12) inner view of pallial cavity and viceral mass of a male, scale = 10 mm; (13)
detail of opened nephridial and pericardial chambers, scale = 2 mm; (14) viceral mass of a male, mantle
partially removed, scale = 2 mm; (15) ventral view of the penis, scale = 2 mm; (16) detail of insertion region
of the vas deferens in seminal groove, showing the receptaculum, scale = 2 тт; (17) pallial oviduct,
tegument removed, scale = 10 mm; (18) detail of the albumen gland showing the vesicles (ve), scale = 2
mm; (19) detail of female genital pore, tegument removed, scale = 2 mm.

ANATOMICAL STUDY ON ТОММА GALEA 27

minute cusp (Fig. 8, ac); base of inner mar-
ginal teeth (Fig. 9) a little longer than that of
outer marginal teeth (Fig. 10). Stomach
poorly developed, with two ducts to digestive
glands, without developed style sac, folds or
typhlosole.

Nervous System: Nerve ring (Fig. 11, nr) with
cerebral ganglia turned to left side in all spec-
imens; from these, three pairs of nerves run
anteriorly, fusing near the proboscis gland
duct loops to become only one pair, which lie
ventraly to the oesophagus (Fig. 11, np). In
mid oesophagial region, this pair of nerves
bifurcates, the median nerves (Fig. 6, in) in-
nervating radular bulb and the lateral nerves
(Fig. 6, on) innervating proboscis wall.

Genital System, Male: Testis (Fig. 14, tt)
branching into digestive gland, mainly on col-
umellar surface of viceral mass. Convoluted
seminal vesicle rather spheric (Fig 14, pt),
confined to anterior part of viceral mass just
anterior to testis. Vas deferens thin walled
(Fig. 14), its anterior region to right of the re-
ceptaculum, fused with its walls (Fig. 16). Re-
ceptaculum a modified, bulging region of the
spermatic groove posterior to vas deferens
insertion in spermatic groove (Fig. 16). In
floor of right margin of pallial cavity, sper-
matic groove thick walled near prostate
gland (Figs. 1, 14, 16). Penis large, somewhat
flattened, with open penial groove, which ter-
minates in a small, pointed papilla at central
region of penis tip (Fig. 15).

Female: Ovary branching into digestive
gland. Oviduct slender (Fig. 17), with a small
gonopericardial duct. Oviduct opening into a
short, thick-walled albumen gland. A series
of small, paired vesicles present in ventral
side of the albumen gland (Figs. 17, 18, ve).
Capsule gland long, curved, thick walled (Fig.
17). Bursa copulatrix long, claviform, sepa-
rate and to right of capsule gland. Posterior
limit of bursa sacciform, thin walled, the walls
gradually becoming thickly muscular anteri-
orly (Figs. 17, 19, bc). Genital pore small
(Figs. 17, 19, gp), to right, behind anus; pore
with two inner divisions: the largest and pos-

terior is the end of the bursa, the smallest and
_ anterior the end of pallial oviduct (Fig. 19).

Measurements
Length, width in mm and if mature (m) or

immature (i): MZUSP 27967: male, 78.5 by
66.5, m; female, 73.0 by 61.0, 1; male, 69.5

by 55.5, m; MZUSP 27984: male, 74.2 by
60.6, m; MZUSP 27968: female, 102.0 by
85.0, m; MZUSP 27986: male 118.5 by 104.0,
т: MZUSP 27969: female 135.0 by 115.0, т;
male 98.7 by 71.0, m.

Habitat

The specimens were obtained by diving,
burrowing in sandy sediment, near rocks or
reefs. Some specimens were also dredged
from about 150 m depth in muddy sediment.

Material Examined

BRAZIL. Espirito Santo: MZUSP 27970, 1
male and 1 female, Barra do Riacho (8/ix/72).
Sao Paulo: MZUSP 27967, 2 males and 1
female, Saco da Ribeira Beach, Ubatuba;
MZUSP 27983, 2 females, Enseada Beach,
Ubatuba (x/91); MZUSP 27984, 1 male, est.
42, otter traw (22/x/86); MZUSP 27985, 1
male, IOUSP-Veliger, ‘‘rede de pesca fixa 8”;
URUGUAY. off Maldonado: MZUSP 27968, 1
female, 35°18’S 52°32’W, “W. Besnard,”
station 1920, 150 m deep (30/x/72); MZUSP
27969, 2 males and 1 female, same data;
MZUSP 27986, 4 males and 1 female, ““W.
Besnard,’ station 1921 OT.9 (20/x/72).

Tonna maculosa (Dillwyn, 1817)
(Figs 20-36)

Synomymy and type material: Turner
(1948: 169) and Matthews et al (1987: 37)

Diagnosis

Shell dark-brown, spotted; outline fusi-
form; sculpture of low spiral ridges. Mantle
border thin. Hypobranchial gland developed.
Kidney with a smooth surface. Proboscis to-
tally within rhynchodeal cavity in fixed spec-
imens. Radular rachidian with crenulations
on base of central cusp; crenulation on main
cusp of lateral radular teeth. Penis without
papilla, with a flap of the tegument on tip.
Anterior region of vas deferens separated
from walls of receptaculum. Inner division in
female genital opening of capsule gland
larger than in 7. galea.

Description

Shell: Detailed decriptions of shell are given
by Turner (1948: 169-172, pl. 75, fig. 2, pl. 76,
figs. 1, 2) and Matthews et al. (1987: 37, fig.

28 SIMONE

6). Protoconch of almost four glassy, convex
whorls, brown in color (Figs. 20, 21).

Head-Foot Complex (Fig. 22): Foot solid,
large, rounded posteriorly, notched ащеп-
orly; propodium narrow, with anterior pedal
opening to ventral slit (Fig. 22, mp). Opercu-
lum lacking in adult. Tentacles long, fairly
thick, bluntly pointed (Fig. 22). Black eyes on
tubercles on outer upper part of tentacle
base. Rhynchodeum with simple, rounded
opening. Proboscis completely retracted
within proboscis sheath in all specimens.
Color of all head-foot structures beige, with
clear-brown irregular spots.

Pallial Complex (Fig. 31): Mantle edge entire,
simple, not reflected, thin, flattened, pale
cream in color. Pallial cavity occupying first
whorl. Siphon long, well developed, pale-
beige in color, with clear-brown irregular
spots. Osphradium large, bipectinate, pro-
portionaly larger than that of Т. galea, situ-
ated on pallial roof at base of siphon; osphra-
dium leaflets lamellate, pigmented brown.
Ctenidium very large, monopectinate, with
many low, triangular filaments. Hypobran-
chial gland developed, along left side of an-
terior region of rectum. Tissue covering rec-
tum and pallial oviduct of females less flaccid
than that of 7. galea, but allowing a well-de-
veloped ad-rectal sinus.

Excretory-Circulatory Systems: Kidney mod-
erately large (Fig. 30), forming a sac situated
like that of Т. galea. Well-developed, slit-like
nephrostome, surrounded by muscular fibres
to form a sphincter (Figs. 30, 31, ne). Internal
structures of kidney similar, but simpler than
in 7. galea, with a smooth surface and a
cream color; its anterior limit does not bulge
with albumen gland of females. Nephridial
gland somewhat inconspicuous, situated
dorsal to nephostome (Fig. 30, ne). Heart
(Figs. 30, 31) with a very thin, transparent,
flaccid walled auricle, and a very thick-walled,
rounded ventricle. Ctenidial vein and auricle
like those of T. galea.

Ad-rectal sinus well developed but less
than that of 7. galea; as in that species, sinus
apparently continuous from kidney chamber
into an aperture (Figs 35, 36, at), with mus-
cular walls, near anus, like an ureter.

Digestive System: Like that of Т. galea (Figs.
23-29). Structures within buccal bulb very
similar to those of 7. galea; a pair of ventral
jaws (Fig. 24, md), two dorso-lateral folds (If)
in buccal cavity, one on either side, with

opening of proboscis gland duct median to
fold and near its anterior end. Radula with
rachidian with a crenulation on base of its
central cusp; tip of this cusp slender and
smooth (Fig. 26). Lateral teeth with a series of
crenulations on main cusp (Fig. 25, cr); small
accessory cusp present (Figs. 25, 27, ac). In-
ner (Fig. 28) and outer (Fig. 29) marginal teeth
similar of those of Т. galea. Oesophagus and
its inner ventral folds and glands similar to 7.
galea, but much shorter (Figs. 23, 24). Oe-
sophageal caecum present, the folds and
glands of the oesophagus terminating in os-
tium of caecum (Fig. 24, eo). Posterior oe-
sophagus without distinct glands or crop.
Stomach poorly developed, with two ducts to
digestive glands, but without developed style
sac, folds, or typhlosole. Inner surface of
posterior oesophagus, stomach and intestine
with low longitudinal folds. Salivary glands,
proboscis glands and their ducts (Figs. 23,
24), similar of those of 7. galea (Weber, 1927).
All anterior structures of digestive system
maintained in position by a tridimensional net
of muscle fibres running to wall of vesopha-
gus, body wall and foot. Looping section of
ducts of proboscis gland lying anterior to
nerve ring, as in 7. galea.

Nervous System: Nerve ring (Fig. 23, nr) with
cerebral ganglia turned to left side in all spec-
imens examined. Ventral pair of nerves of
proboscis similar to those of 7. galea.

Genital System, Male: Testis branching into
digestive gland, concentrated mainly on col-
umellar surface of viceral mass. End of vas
deferens an enclosed, small, thin-walled
tube, lying to right of receptaculum, without
fusion except for insertion (Fig. 34). Recep-
taculum a modified, bulging region of sper-
matic groove, posterior of insertion of vas
deferens in floor of pallial cavity (Fig. 34).
Spermatic groove in right side of floor of pal-
lial cavity, thick walled, by the prostate gland
(Figs. 22, 32, 33, 34). Penis very large (Figs.
32, 33), curved backwards, somewhat flat-
tened, its free end curving downwards fol-
lowing curve of floor of pallial cavity, blunt at
the apex. At right side, near apex of penis
there is a flap of tissue (fig. 32: pf), under
which the penial duct opens; there is no pa-
pilla.

Female: Ovary branching into digestive
gland. Oviduct slender, opening into a short,
thick-walled albumen gland (Fig. 35). A series
of paired vesicles present in ventral side of

ANATOMICAL STUDY ON ТОММА GALEA 29

FIGS. 20-31; Tonna maculosa: (20) protoconch in profile; (21) protoconch, apical view, scale (Figs. 20, 21)
= 2 mm; (22) head-foot complex from male, scale = 5 mm; (23) dorsal view of anterior region of the digestive
system, proboscis opened, scale = 5 mm; (24) the same, oesophagus opened longitudinally, scale = 5 mm;
(25) lateral tooth of radula, showing the crenutation (cr) and the accessory cusp (ac), scale = 0.1 mm; (26)
rachidian tooth; (27) lateral tooth; (28) inner marginal tooth; (29) outer marginal tooth, scale (Figs. 26-29) =
0.2 mm; (30) detail of opened nephidial and pericardial champers, scale = 2 mm; (31) pallial cavity of a male,
inner view, scale = 5 mm.

30 SIMONE

FIGS. 32-36: Tonna maculosa: (32) penis and seminal groove, dorsal view; (33) the same, ventral view,
scale = 5 тт; (34) detail of the insertion of vas deferens in seminal groove, showing the receptaculum,
scale = 1 mm; (35) pallial oviduct ventral view, tegument partially removed, scale = 5 mm; (36) detail of
female genital pore, tegument removed, scale = 2 mm.

albumen gland, similar to those of 7. galea,
Capsule gland long, сигуеа, thick walled (Fig.
35, cg). Bursa copulatrix long, claviform,
slender, separate and to right of capsule
gland; posterior end of the bursa sacciform,
thin walled, the walls becoming thick and
muscular anteriorly (Fig. 35, bc). Small genital
opening at right and posterior to anus (Figs.
35, 36, gp). Genital opening with two inner
divisions, the smallest and posterior is end of
bursa (bc), and the larger and anterior is end
of capsule gland (cg) (fig. 36).

Measurements

MZUSP 27961, female = length 65.0 mm
by width 44.0 mm; male = 40.4 mm by 26.5
mm. MZUSP 27962, female = 45.3 mm by
31.0 mm.

Habitat
The collected specimens were found by

diving or at low tide, burrowing on sandy bot-
toms near reefs.

Material Examined

BRAZIL. Bahia: MZUSP 27961 (one male
and one female) Цариа Beach, Salvador (7/
vii/71); MZUSP 27962 (one female) Карча,
Salvador (29/ix/84).

DISCUSSION

Tonna galea differs anatomically from 7.
maculosa in having (1) a thick mantle border;
(2) darker spots on the epidermis; (3) a less-
developed hypobranchial gland; (4) a propor-
tionally smaller osphradium; (5) a more de-
veloped kidney, with more complex internal
structure; (6) proboscis extending 50% from
rhynchodeum (in 7. maculosa proboscis al-
ways completely retracted within proboscis
sheath) in fixed specimens; (7) oesophagus
and inner oesophagial structures much
longer; (8) central cusp of radular rachidian
teeth and lateral teeth without crenulations
(present in 7. maculosa); (9) penis with a pa-

ANATOMICAL STUDY ОМ TONNA GALEA 31

FIG. 37: detail of anal region of Tonna galea, scale
= 2 mm.

pilla (7. maculosa has a flap, without papilla);
(10) end region of vas deferens fused with the
receptaculum walls (in 7. maculosa this duct
is free); and (11) female genital pore with
larger inner opening to the bursa (7. таси-
losa has the larger opening leading to the
capsule glana).

The function of the aperture near the anus
(Figs 36, 37, at) in both species is unknown. It
probably controls the exit of the inner fluid of
the ad-rectal sinus, which 1$ apparently con-
tinuous to the kidney chamber. These struc-
tures resemble the ureter of the Viviparidae
(Hyman, 1967), for example, which have no
well-developed nephrostome as in Tonna
(Figs. 21, 31, ne). The fortuitous use of the
ad-rectal sinus as an ureter merits further
study.

Some differences between the literature
accounts (Turner, 1948; Mattews et al., 1987)
and the specimens studied here were: (1) the
protoconchs of both species are closely sim-
_ ilar, and of almost four whorls, in contrast

with the data of Matthews et al. (1987), in
which differences in number of whorls was
given; (2) the jaw lies ventral to the proboscis;
(3) the radular rachidian of 7. maculosa has a
crenulation only on the base of the central
cusp; the tip of this cusp is smooth and slen-
der (Fig. 26, cr); (4) the radular lateral tooth

has a small, but conspicuous accessory
basal cusp in all specimens examined of both
species (Figs 8, 25, 27, ac); (5) Turner (1948:
168) reported an extremely long and flagel-
late papilla in the penis of Т. maculosa, dif-
ferent from the penis described herein, in
which the papilla is lacking (Figs. 32, 33). Fur-
ther investigation is need to determine the
significance of these differences.

The proboscis of Tonna, as in all known
Tonnoidea, has a great development of the
buccal mass, this region taking most of the
proboscis length (Figs. 2, 23). The proposal
of the tonoidean proboscis as a distinct type
(Day, 1969) is perhaps not justified, but rather
it can be regarded as a specialized and mod-
ified pleurembolic type.

The auricle structure and pallial oviduct
may be considered as additional characters of
Tonnoidea, in addition to the anterior region of
the digestive system. This type of auricle 1$
found in Cassidaria rugosa (Ranellidae) (Rey-
nell, 1905). However, no reference to the pal-
lial oviduct has been found in the literature
except for Bursa cruentata (Houbrick & Fret-
ter, 1969: 417), but details are missing that
would allow a full comparison.

Besides radular aspects, other charac-
ters of the Tonna digestive system differing
from other Tonnoidea (Reynell, 1902; Day,
1969; Houbrick & Fretter, 1969; Lewis, 1972;
Hughes & Hughes, 1981) are (1) the presence
of a oesophagial caecum and (2) the absence
of a clear oesophagial gland (crop or bulb) in
the posterior oesophagus. These are per-
haps characters of Tonnidae.

LITERATURE CITED

BENTIVEGNA, F. 8 A. TOSCANO, 1991, Observa-
tion au laboratoire sur le comportement alimen-
taire de trois especes de la superfamille Ton-
noidea (Mollusca, Gastropoda). Revue Française
d’Aquariologie Herpetologie, 18: 33-38

DAY, J. A., 1969, Feeding of the cymatiid gastro-
pod, Argobuccinum argus, in relation to the
structure of the proboscis and secretions of the
proboscis gland. American Zoologist, 9: 909-
916

HOUBRICK, J. R. 8 V. FRETTER, 1969, Some as-
pects of the functional anatomy and biology of
Cymatium and Bursa. Proceedings of the Mala-
cological Society of London, 38: 415-429

HUGHES, В. М. & H. Р. 1. HUGHES, 1981, Mor-
phological and behavioural aspects of feeding in
the Cassidae (Tonnacea, Mesogastropoda). Ma-
lacologia, 20: 385-402

32 SIMONE

HYMAN, L. H., 1967, The invertebrates, Volume VI,
Mollusca |. McGraw-Hill Book Company. New
York, 792 pp.

LEWIS, H. 1972, Notes in the genus Distorsio (Cy-
matiidae) with descriptions of new species. Nau-
tilus, 86: 27-50

MATTHEWS, H. R.; J. Н. М. LEAL 8 A. С. S. CO-
ELHO, 1987, Superfamilia Tonnacea no Brasil.
Vli—Familia Tonnidae (Mollusca: Gastropoda).
Arquivos de Ciéncias do Mar, 26: 29-45

MORRETES, F. L., 1949, Ensaio de catálogo dos
moluscos do Brasil. Arquivos do Museu Para-
naense, 7: 2-216

REYNELL, A., 1905, Some account of the anatomy
of Cassidaria rugosa (Linn.). Proceedings of the
Malacological Society of London, 6: 292-299, pl.
6

RIOS, E. C., 1985, Sea shells of Brazil. Fundacáo
Universidade de Rio Grande, Fundacáo Cidade
de Rio Grande, Museu Oceanográfico. Rio
Grande 239 pp., 102 pls.

TURNER, R. D., 1948, The family Tonnidae in the
western Atlantic. Johnsonia, 2: 165-192

WEBER, H., 1927, Der Darm von Dolium galea L.
eine vergleichend anatomische Undersuchung
unter besonderer Berücksichtigung der Trito-
пит Arten. Zeitschrift für Morphologie und
Okologie der Tiere, 8: 663-804

Revised Ms. accepted 26 October 1994

ABBREVIATIONS
ac: accessory cusp of lateral tooth
ag: albumen gland
ao: anterior aorta
an: anus
at: aperture of ad-rectal sinus
au: auricle
Be: bursa copulatrix
bu: buccal complex
CE central fold of buccal complex
cm: columellar muscle
Cp: capsule gland
ch crenulated ridge
CV: ctenidial vein
dc: duct of oesophagial caecum
dg: digestive gland
dp: duct of proboscis gland
ес: oesophagial caecum

eo: ostium of oesophagial caecum

floor of pallial cavity
foot

gonopericardial duct
gill

female genital opening
hypobanchial gland
inner proboscis nerve
intestine

kidney

membrane between kidney and
pallial cavities

lateral fold of buccal complex
buccal lips

mantle border
mandibule (jaw)
muscle fibers
mid-ventral mucous gland of
oesophagus

mouth

anterior pedal gland
nephrostome
nephridial gland

nerve ring

anterior oesophagus
oesophagial folds
outer proboscis nerve
posterior oesophagus
osphradium

oviduct

posterior aorta
proboscis

pericardial chamber
penis

penian flap

penian seminal groove
proboscis nerve
proboscis gland
penian papilla

: convoluted seminal vesicle

radular complex
proboscis sheath
receptaculum seminalis
radular nucleus
rhynchodeum

rectum

salivary gland

seminal groove

siphon

cephalic tentacle

testis

vas deferens

vesicles of albumen gland
ventricle

MALACOLOGIA, 1995, 37(1): 33-40

A TAXONOMIC APPLICATION OF MULTIVARIATE MIXTURE
ANALYSIS IN PATELLIDAE

J. D. Acuña? & М. A. Muñoz*

ABSTRACT

Multivariate mixture analysis is a powerful tool that appears to be useful for dealing with
situations in which several traits with overlapping variation are used for species discrimination.
A multivariate mixture analysis technique is applied to discrimination of two sibling species of
the genus Patella (P. aspera Róding and P. caerulea Linnaeus). These species show a sub-
stantial overlap in the distribution of maximum shell width and shell height. In spite of this
overlap, the results of mixture analysis in a sample of 101 specimens, classified but treated as
unclassified for the purpose of the analysis, suggest the existence of two mixed distributions.
Moreover, examination of specimen classification derived from mixture analysis reveals that
these mixed distributions correspond to P. aspera and P. caerulea. The estimated values of
mixture parameters confirm a substantial overlap in the bivariate distribution of maximum shell
width and shell height of the two species. We hope that these results will contribute to making
multivariate mixture analysis more popular among taxonomists.

Key words: multivariate mixture analysis, taxonomy, species discrimination, Gastropoda,

Patellidae, Patella aspera, Patella caerulea.

INTRODUCTION

Individual variation in morphological traits
often provides valuable information for spe-
cies discrimination. However, the interpreta-
tion of variation is sometimes difficult due to
polymorphism, polytypy, and similarity be-
tween species.

A particularly difficult case arises when
species overlap in the distribution of contin-
uous morphologic traits used for their classi-
fication. Overlap in these traits can be mis-
takenly interpreted as evidence in favor of
interbreeding when, in fact, two or more non-
hybridizing species are involved.

The solution to this problem requires a de-
tailed analysis of trait variation which is fre-
quently made resorting to several statistical
methods. However, taxonomists have not
taken full advantage of recent advances in
statistics to deal with the problem of overlap-
ping variation. Mixture analysis techniques
offer a useful alternative to traditional meth-
ods of statistical analysis when overlapping
variation is a concern (Everitt 8 Hand, 1981;
Titterington et al., 1985; McLachlan 8 Bas-
ford, 1988).

Although originally developed to deal with
problems in the biological realm (Pearson,

1894), mixture analysis has received little at-
tention from biologists. This may be in part
due to difficulties in the computations re-
quired by the method. Recently, several au-
thors have confirmed the usefulness of mix-
ture analysis for species discrimination (Do &
McLachlan, 1984), the resolution of the age-
class structure of a population (Equihua,
1988), and the study of sexual dimorphism
(Flury et al., 1992).

In a previous герой (Muñoz & Acuña,
1994), we were able to discriminate between
two sibling species of the genus Patella (P.
aspera Róding and P. caerulea Linnaeus) by
means of a univariate mixture analysis tech-
nique. The specimens included in the sample
were actually classified, but for the purpose
of the analysis they were treated as unclas-
sified. The taxonomic trait that we selected
was shell height. Our aim was to show the
usefulness of mixture analysis as a tool for
species discrimination using a trait with over-
lapping variation. Here we attempt the dis-
crimination of these two species with the
same method but using two conchological
traits, which requires the use of multivariate
mixture analysis. Multivariate mixture analy-
sis provides powerful techniques for dealing
with situations in which several traits with

“Departamento de Biología Animal, Universidad de Valencia, Dr. Moliner, 50, 46100 Burjasot, Valencia, España.
“Departamento de Ciencias Morfolögicas |, Universidad Complutense de Madrid, Arcos de Jalón, s/n, 28037 Madrid,

España.

34 ACUÑA & MUÑOZ

overlapping variation are used for species
discrimination. We hope that our results will
contribute to making multivariate mixture
analysis more popular among taxonomists.

PATELLA ASPERA AND
PATELLA CAERULEA

Patella aspera Róding (= P. ulyssiponensis
Gmelin) and P. caerulea Linnaeus are two
very abundant European marine gastropods.
The first is found in the Mediterranean and is
also widely distributed along the Atlantic
coast, whereas the second 1$ restricted to the
Mediterranean. The two species live in very
similar habitats. Both are found on hard sub-
strata in the littoral zone and reach upper
subtidal levels, although P. aspera has a
lower distribution range in the littoral zone
and exhibits a preference for areas exposed
to wave action. From a taxonomic stand-
point, P. aspera and P. caerulea are consid-
ered two separate species, although their
specific status was a contentious issue for
some time.

Part of the protracted debate regarding the
taxonomic status of these two species came
about because there 1$ substantial overlap in
the distribution of traits (shell shape, orna-
mentation, and coloration) used for their clas-
sification. In the past, taxonomists interpreted
this overlap as arising from interbreeding and
accordingly rejected a specific distinction or
considered this to be an instance of incom-
plete speciation (Fischer-Piette, 1935, 1938;
Evans, 1953, 1958). Others, however, argued
that reproductive features (e.g., timing of the
breeding season) could be used as a basis for
discriminating between the two species (Fi-
scher-Piette, 1948). The controversy was fi-
nally settled when Fischer-Piette 8 Gaillard
(1959) reported clear-cut species differences
in the single cusp lateral teeth of the radula.
More recently, analyses of a variety of taxo-
nomic traits has confirmed the taxonomic va-
lidity of Patella aspera and P. caerulea. Such
is the case of studies that relied on caryotypic
(Cervella et al., 1988), electrophoretic (Sella et
al., 1989; Cretella et al., 1990) and soft-part
traits (Cretella et al., 1990).

A review of the literature reveals that con-
chological traits, when taken altogether, allow
for a separation of the two species. Indeed,
several authors have relied on conchological
traits to separate samples, the species mem-
bership of which was later confirmed by dif-

ferences in radular (Fischer-Piette & Gaillard,
1959), electrophoretic (Cretella et al., 1990), or
soft-part traits (Cretella et al., 1990). Never-
theless, taxonomists were reluctant to grant
these Patella their current specific status until
clear-cut differences were found in non-con-
chological traits. This may be partly due to the
difficulties involved in analyzing shell traits
with overlapping variation.

MATERIALS AND METHODS

Our study of shell height by means of
univariate mixture analysis (Muñoz & Acuña,
1994) used a very large sample. Part of this
material was judged suitable for the present
study.

The complete sample (detailed description
in Muñoz & Acuña, 1994) included over а
thousand specimens of Patella aspera and P.
caerulea obtained at Cabo Oropesa, Castel-
lón, España. Specimens of all available sizes
were collected randomly in a narrow band of
uniform characteristics located at the base of
the littoral zone. Sampling took place in May
1989. In the laboratory, shell length (distance
between anterior and posterior shell mar-
gins), maximum width (maximum distance
between lateral shell margins), and height
(distance between apex and line between an-
terior and posterior shell margins) were mea-
sured to the nearest 0.05 mm using calipers
on 1025 useful, whole shells. Specimens
were also assigned to either species using
non-conchological traits, mainly foot mor-
phology and color (Cretella et al., 1990). At
Cabo Oropesa, Patella aspera can be easily
recognized by its pyriform or oval foot, with
sole yellow or cream with no dark areas. Pa-
tella caerulea, on the other hand, has an oval
foot, with sole dark gray or bluish with edge
and center cream. Use of these diagnostic
characters resulted in 439 specimens being
classified as P. aspera and 581 as P. caer-
ulea. Five specimens could not be unambig-
uosly assigned to either species and were
classified as doubtful. Later, the range of
shell lengths in the total sample (5.30-35.10
mm) was divided into 31 intervals of 1 mm,
which were operationally considered as
growth stages. Specimens were, irrespective
of their specific identity, grouped into these
31 shell-length class intervals.

Because the performance of mixture anal-
ysis is greatly improved by a large sample

MULTIVARIATE MIXTURE ANALYSIS IN PATELLIDAE 35

size (e.g., Equihua, 1988), only the largest
subsample was considered for the analysis
performed in the present study. This sub-
sample included all specimens with shell
length between 14 and 15 mm (n = 103). Ac-
curate measurements of maximum shell
width and shell height could only be obtained
from 101 of the 103 specimens (37 belonging
to Patella aspera and 64 to P. caerulea) yield-
ing 101 bivariate data that were submitted to
the statistical analysis.

The subsample was considered a mixture
(superposition of density functions) of two
components with bivariate normal distribu-
tion as to maximum shell width and shell
height. Multinormality of the continuous phe-
notypic traits in a population is a frequent
theoretical assumption (Нат & Clark, 1989).
In a real mixture situation, testing for normal-
ity would be impossible because group
membership of the specimens 1$ not known.
In our sample of classified specimens, multi-
normality was tested for confirmation. It was
done for each species separately using (1)
Kolmogorov-Smirnov goodness-of-fit test
with Lilliefors correction (Sokal & Rohlf, 1981)
to test the univariate normality of the mar-
ginal distributions, and (2) Mardia’s test of
multinormality (Mardia, 1970; Appendix 1) to
evaluate joint normality. Although marginal
normality does not imply joint normality, the
converse is true. Hence, test (1) is useful for a
fast and easy detection of many types of
non-normality, whereas test (2) permits eval-
uation of multivariate normality when results
of the previous test are inconclusive (i.e., fails
to reject the hypothesis of normality).

In the Patella sample case, results of good-
ness of fit testing revealed no significant de-
partures from marginal normality and multi-
normality (P > 0.05) in the growth stage
(shell-length class interval) that is the focus of
this report, as well as in most other growth
stages of either species.

Statistical analysis of the mixture required
estimation of 11 parameters, including four
means, four variances, two covariances, and
one mixing parameter (proportion of either
component in the mixture). The estimation
was accomplished by means of a maximum-
likelihood approach. The EM algorithm
(Dempster et al., 1977) was used to compute
the maximum-likelihood estimates of the pa-
rameters using the equations studied by
Wolfe (1970). The procedure has been de-
scribed by Everitt & Hand (1981) for the gen-
eral case of a mixture of multivariate normal

distributions (an arbitrary number of mixed
distributions and variables, with all parame-
ters unknown) (Appendix 2).

This method, although slow, is easily pro-
grammable and very stable (i.e., shows little
dependency on the initial estimates of the
parameters). In the case studied here, includ-
ing two mixed distributions and two vari-
ables, the programming was particularly sim-
ple. Initial estimates of the parameters were
computed by the program directly from the
data. The variance/covariance matrices of
the components were assumed equal to the
variance/covariance matrix of the mixture,
and the means were calculated by imposing
a small deviation (10% of the range of the
variables) on both sides of the mixture
means. The means calculated in this way
were then grouped into vectors taking into
account the sign of the correlation of the data
in the mixture. Mixing proportions were set at
0.5. This approach has proved successful in
several runs with simulated data. In addition,
different initial estimates were used in order
to confirm that the results of the mixture anal-
ysis did not correspond to a local maximum
of likelihood (Appendix 2). The convergence
criterion of the iterative procedure was spec-
ified in terms of the Euclidean distance be-
tween successive estimates of the parameter
vector. Following Everitt (1984), the criterion
value was set at 0.0001.

A likelihood ratio test was used to evaluate
the goodness-of-fit of the mixture (Hassel-
blad, 1969; Everitt & Hand, 1981; Equihua,
1988). The statistic is given by

G=2 (Ly — №

where L, is the log-likelihood computed un-
der the null hypothesis (which assumes only
one distribution), and L, is the log-likelihood
under the alternative hypothesis (which as-
sumes the mixture of two distributions). Sta-
tistic G is asymptotically distributed as chi-
square with degrees of freedom equal to the
difference in the number of parameters be-
tween the two hypotheses (six in the present
case). This approach, however, is not devoid
of criticisms (Everitt & Hand, 1981; Tittering-
ton et al., 1985). Under the null hypothesis,
the mixing proportions fall in the boundary of
the parameter space, so that conditions for G
to be asymptotically distributed as chi-
square are not fulfilled. However, a satisfac-
tory performance of the test was found by
Hasselblad (1969) with mixtures of exponen-

36 ACUÑA & MUÑOZ

tial, Poisson, and binomial distributions.
Based on this evidence, Equihua (1988) has
argued that the likelihood ratio test can be of
assistance in assessing the number of com-
ponents in a mixture.

Once the analysis was completed, the
probabilities of membership (posterior prob-
abilities) of each datum (specimen) to each
component (species) in the bivariate mixture
were calculated by dividing the component
density function weighed by its proportion in
the mixture by the mixture density function.

The presence of two components in the
mixture could be taken as evidence of a spe-
cific discrimination. Alternative interpreta-
tions based on other phenomena that yield
mixed distributions in natural populations ap-
pear rather unlikely. For example, the species
are hermaphroditic (Bacci, 1947; Fretter &
Graham, 1976), thus sexual dimorphism is
not a likely explanation for the presence of
two components in the mixture. Mendelian
segregation in the shell dimensions of these
two species has never been reported, and it
is likely that, as is the case with other organ-
isms, traits related to body size are under
additive polygenic control resulting in a nor-
mal distribution of maximum shell width and
shell height (Falconer, 1989; Hartl 8 Clark,
1989). Also, because the samples were col-
lected in a uniform environment, a possible
effect of disruptive selection and/or differen-
tial reaction seems unlikely.

RESULTS

Figure 1 shows scatterplot of the two vari-
ables measured in the 14-15 mm shell-length
growth stage. Inspection does not permit
discrimination of two components. However,
the results of the mixture analysis (Table 1)
suggest the existence of two mixed multivari-
ate distributions. This can be inferred from (1)
the absence of an empty component in the
mixture; (2) the absence of mixing propor-
tions suggesting a component with a mar-
ginal representation as could arise from a
spurious frequency peak in the tails of the
distribution, and (3) the results of the likeli-
hood ratio test, which allow rejection of the
null hypothesis with P < 0.05 (G = 12.72; d.f.
= 6). Furthermore, the estimated values of the
parameters indicate a substantial overlap in
the bivariate distributions for the two compo-
nents.

The taxonomic interpretation of the results

SHELL HEIGHT (mm)

9 10 11 12 13
MAXIMUM SHELL WIDTH (тт)

FIG. 1. Scatterplot for the sample used in the
present study. Specimens belonging to Patella as-
pera are symbolized by closed dots, and those be-
longing to P. caerulea by open dots. Superim-
posed lines limit the regions with probabilities of
membership (posterior probabilities) > 0.95 in the
components detected through the use of mixture
analysis.

TABLE 1. Results of the mixture analysis.

Component 1 Component 2

(Patella aspera) (Patella caerulea)

Mixing

proportion 0.45 0.55
Maximum shell

width mean 10.71 11.62
Shell height

mean 4.76 3.88
Maximum shell

width variance 0.35 0.31
Shell height

variance 0.35 0.18
Maximum shell

width—Shell

height covariance 0.04 0.07

of the mixture analysis is straightforward if
one considers simultaneously the probability
that each specimens belongs to any one
component in the mixture and the results of
their specific diagnosis based on non-con-
chological traits (Table 2; Fig. 1). When strin-
gent probability levels are applied (e.g., 0.99
or 0.95), the classification derived from the
mixture analysis is congruent with that based
on non-conchological traits. With few excep-
tions, the component with the lowest maxi-
mum shell width mean and the highest shell
height mean (component 1) corresponds to
Patella aspera shells, whereas the compo-
nent with the highest maximum shell width

MULTIVARIATE MIXTURE ANALYSIS IN PATELLIDAE 37

TABLE 2. A comparison of the classifications arising
from mixture analysis and from specific diagnosis
based on non-conchological traits. PM: probability
of membership in a component; A: number of spec-
imens ascribed to either component; lA: number of
incorrect ascriptions; OIA: observed percentage of
incorrect ascriptions; ElA: expected percentage
of incorrect ascriptions.

PM A IA OIA(%) EIA(%)
0.99 31 0 0.00 1.00
0.95 59 5 8.47 5.00
0.90 70 5 7.14 10.00
0.75 89 8 8.99 25.00
0.50 101 11 10.89 50.00

mean and the lowest shell height mean (com-
ponent 2) corresponds to P. caerulea shells.
The number of incorrect ascriptions in-
creases at low probability levels (e.g., 0.75 or
0.50), but is always under reasonable values.
Therefore, it seems safe to conclude that the
two components detected through the use of
mixture analysis correspond to P. aspera and
P. caerulea.

Although the following discussion is mainly
based on the analysis of specimens between
14-15 mm in shell length, the same proce-
dure was also applied to other shell-length
class intervals in the sample. Only occasion-
ally did mixture analysis reveal two compo-
nents using data from a single interval [e.g.,
18-19 mm (n = 65)]. However, in some cases
discrimination was accomplished after pool-
ing data from two successive intervals [e.g.,
14-15 mm + 15-16 mm (n = 101 + 87), 17-18
mm + 18-19 mm (n = 68 + 65)].

DISCUSSION

The results of the mixture analysis pre-
sented here indicate that there are only slight
differences between the sibling species Pa-
tella aspera and P. caerulea as far as the bi-
variate distribution of maximum shell width
and shell height. The shell of P. aspera 1$
slightly narrower and taller than that of P.
caerulea. It is worth noting that a third vari-
able, namely shell length, was bearing on the
results of the bivariate analysis, because the
specimens were grouped into growth stages
according to shell length. Therefore, the spe-
cific differences revealed by the analysis con-
cern all three variables that determine the
general shape of the patelliform shells. The
shell of P. aspera is slightly more oval-conic

and elevated than that of P. caerulea. This
difference has been reported in type-materi-
als for a long time (Bucquoy et al., 1886;
Christiaens, 1973; Powell, 1973) and is po-
tentially interesting, because shell shape is a
taxonomic trait that was extensively investi-
gated in relation to adaptive value (Segal,
1956; Lowell, 1984).

The discrimination between Patella aspera
and P. caerulea extends the results of a pre-
vious report (Muñoz & Acuña, 1994) and il-
lustrates the use of multivariate mixture anal-
ysis to deal with situations in which several
traits with overlapping variation are used for
classification. Multivariate mixture analysis
proved capable of discriminating between
these two species in spite of a substantial
overlap in the mixed distributions and a mod-
erate sample size. We hope that these results
will encourage the use of multivariate mixture
analysis among taxonomists.

Moreover, mixture analysis seems prefera-
ble in many taxonomic applications to other
statistical procedures, such as discriminant
analysis and ordinary cluster analysis (Flury
et al., 1992). Mixture of distributions arise
when a population is subdivided into homo-
geneous components, but it is unknown from
which of the components any given observa-
tion originates. Mixture analysis models this
situation, attempting to estimate the statisti-
cal parameters of the components and their
proportions in the population by means of a
sample of unclassified observations. Later,
the observations can be classified using the
parameter estimates. In discriminant analy-
sis, the basic problem 1$ to assign a given
observation to one of two or more classes on
the basis of the value of this observation. The
procedure requires reference samples with
known group membership. Ordinary cluster
analysis attempts to partition the data into
homogeneous subgroups without consider-
ing a statistical model and assuming that the
subgroups are distinct and do not overlap.
Therefore, mixture analysis seems preferable
for dealing with taxonomic problems in which
(1) the statistical distribution of traits 1$ used
for discrimination, (2) overlapping variation is
present, and (3) reference samples are not
available. In these situations, mixture analy-
sis is a reasonable alternative provided that
the number of variables (and therefore pa-
rameters that need estimation) does not de-
mand an inordinate sample size. When the
number of variables is high, such methods as
principal component analysis can be used to

38 ACUÑA & MUÑOZ

reduce the dimensions of the variability to a
smaller number of meaningful and indepen-
dent variables.

However, a general discussion of the
methodological implications of mixture anal-
ysis is necessary. In particular, application of
mixture analysis to the problem of species
discrimination entails a risk of misinterpreta-
tion. The coexistence of populations leads,
indeed, to mixtures of distributions, but other
phenomena may be responsible for the pres-
ence of mixed distributions in populations.
Mendelian segregation, sexual dimorphism,
disruptive selection, and multiple environ-
mental reaction are examples of potential
sources of mixed distributions. Thus, mixture
analysis, like other more popular statistical
techniques, should be not be considered as
an alternative, but rather as an aid to tradi-
tional taxonomic methods.

ACKNOWLEDGMENTS

We are very grateful to J. D. Bermúdez (De-
partamento de Estadística e Investigación
Operativa, Universidad de Valencia), E. Font
(Departamento de Biología Animal, Univer-
sidad de Valencia), and M. Sendra (Departa-
mento de Estadística e Investigación Opera-
tiva, Universidad de Valencia) for providing
useful information and discussion of an ear-
lier draft of this manuscript. We also thank R.
T. Dillon and an anonymous referee for stim-
ulating discussion and constructive criticism.

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CERVELLA, P., L. RAMELLA, C. A. ROBOTTI 4 G.
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CHRISTIAENS, J., 1973, Révision du genre Patella
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CRETELLA, M., G. SCILLITANI, Е. ТОЗСАМО, P.
TURELLA & О. PICARIELLO, 1990, Comparative
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DEMPSTER, А. P., М. M. LAIRD & D. В. RUBIN,
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DO, K. & G. J. MCLACHLAN, 1984, Estimation of
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EVERITT, B. S. & D. J. HAND, 1981, Finite mixture
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géographie. Les patelles d'Europe et d’Afrique
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FISCHER-PIETTE, E. & J. М. GAILLARD, 1959, Les
patelles au long des côtes Atlantiques lbériques
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Gender identification of water pipits (Anthus
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ITA

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MULTIVARIATE MIXTURE ANALYSIS IN PATELLIDAE 39

MCLACHLAN, С. J. 8 К. E. BASFORD, 1988, Mix-
ture models. Marcel Dekker, New York. 253 pp.

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Revised Ms. accepted 17 January 1995

APPENDIX 1
Mardia’s Test of Multinormality

Mardia’s test is a simple but useful test of
multinormality (Mardia, 1970). If y,,..., y,
are a random sample of k-dimensional data
from a normal multivariate distribution of y,
the sample estimators of skewness and kur-
tosis, b, and b,, are

n

n
by = 1/12 Y Y In -w’S (y, - УВ
h=1 i=1

=)

62 = 1/n) [y -У)’5 * (y, -УР
i=1

where y is the sample estimate of the mean
vector and $ is the sample estimate of the
variance/covariance matrix. Then, for large
samples, we have, asymptotically,

A=1/6nb; -xf {= 1/6 k(k + 1)(k + 2)

Bee ran
[8k(k + 2) /n]

Although these distributions are only ap-
proximate for moderate sample sizes, they at
least indicate whether the data show any
marked departure from multivariate normal-

ity.

APPENDIX 2

Maximum Likelihood Estimation of the
Parameters of a Mixture of Multivariate
Normal Distributions by Means of the
EM Algorithm

ИС.,..., Cy are the g components
with multivariate normal distribution which
contribute to the mixture in proportions
ру, -.., Pg respectively, О < pj < 1

9
Ур =1
i=1

and f, (y) is the probability density function of

the distribution of y ~ М, (m, S) in С, (i=1,
., 9) given by
f(y) =1218,1 1

exp [-1/2(y — т)” s (y — m;)]

then, considering a random sample of n k-di-

mensional data y;,..., y, the density of y; is
given by

9
fly) = Ур) (j=1, ..., n)
i=1

and the maximum likelihood estimates of the
parameters of the mixture (Everitt & Hand,
1981) are given by

n
В, = У wi/n dei, 2.9) (1)
j=1
n
j=1
n
Si = > Wi (y; = mi) (y; = m;)’ /n Pi

iz

where

40 ACUÑA 8 MUÑOZ

Equations 1 through 3 are solved itera-
tively. Initial estimates of the parameters are
used in the right hand side of the equations to
obtain other estimates of the parameters op-
erating the data. The resulting estimates are
then used in the right hand side of the equa-
tions to yield new estimates, and so on. The
iterative procedure continues until two suc-
cessive estimates of the parameters differ
only by some arbitrarily small amount.

As is the case with other iterative pro-
cesses, with the EM algorithm there is no
guarantee that the global maximum of the
likelihood function will ever be found. It is
possible that, depending on the initial esti-

mates of the parameters, one or several local
maxima might result. In these instances, the
solution with the largest likelihood would be
accepted. However, EM algorithm shows,
compared to other algorithms, little depen-
dency on the initial estimates of the parame-
ters (Everitt, 1984).

Note: A copy of the program written to
compute the maximum likelihood estimates
of the parameters of a mixture of two bivari-
ate normal distributions is available from the
authors upon request. This program 1$ writ-
ten in Quick Basic and runs on IBM PC or
compatible computers.

MALACOLOGIA, 1995, 37(1): 41-52

THE LIFE CYCLE, POPULATION DYNAMICS, GROWTH AND SECONDARY
PRODUCTION OF THE SNAIL VIVIPARUS CONTECTUS (MILLET)
(GASTROPODA: PROSOBRANCHIA) IN THE MARSHES OF THE RIVER
STRYMONAS, SERRES, MACEDONIA, NORTHERN GREECE.

М. Eleutheriadis & М. Lazaridou-Dimitriadou

Section of Zoology, Department of Biology, University of Thessaloniki 54006,
Thessaloniki, Greece

ABSTRACT

The life cycle, population dynamics, growth and secondary production of the prosobranch
freshwater snail Viviparus contectus were studied in the marshes of Strymonas River upstream
of its entry into the artificial Lake Kerkini, Serres, Macedonia, northern Greece. Demographic
analysis of the population of V. contectus revealed that two or more generations existed in the
field throughout the year. The sex ratio was 1:1. Reproduction took place in the beginning of
spring or at the end of autumn, depending on the water level and water temperature in the study
area. Growth of newly born individuals mainly took place during spring and early summer. Von
Bertalanffy's method suggested that V. contectus may live up to five years to reach ¡ts maxi-
mum possible size (51 mm). Annual secondary production, calculated by Hynes' size frequency
method, revealed a mean annual density of three individuals/m?, a mean standing crop (В) of
4.19 g dry body weight/(m”.year), and an annual production (P) of 13.45 + 12.9 д dry body
weigh/(m?.year). Annual turnover ratio (Р/В) was 3.21 and turnover time was 113.7 days.

Key words: Viviparus contectus, Gastropoda, ecology, population dynamics, production.

INTRODUCTION

Viviparus contectus is a prosobranch snail
found in the marshes of Strymonas River
(old bed of Strymonas River) upstream of its
entry to Lake Kerkini, which lies 100 km from
Thessaloniki in Serres, Macedonia, northern
Greece. These marshes were formed in 1982
after the construction of a new bed of Stry-
monas River parallel to the old one. The lake
is an artificial water storage reservoir, con-
structed on the site of a natural marsh and
small natural lake. The area of the lake 1$ in-
cluded in the list of the internationally impor-
tant wetlands, edited by the United Nations
at Ramsar in 1971. It supports a numerically
rich, diverse wildlife (Pelecanus onocrotalus,
P. crispus, Hirudo medicinalis), including V.
contectus.

Studies on the biology and ecology of fam-
ily Viviparidae occurring in freshwater show
considerable differences between genus,
species and populations (Stanczykowska et
al., 1971; Samochwalenco 4 Stanczykowska,
1972; Young, 1975; Bernardi et al., 1976;
Browne, 1978; Vail, 1978; Jokinen et al., 1982;
ВЫ & Gebhardt, 1986). Studies of these dif-
ferences can provide worthwhile insights into
the strategies that the species follow (Young,
1975).

41

Although the prosobranch snail Viviparus
contectus (Millet) is widespread in Europe
(Zhadin, 1952), little is known about its life
history, apart from studies done by Samoch-
walenco & Stanczykowska (1972) in Poland.
К is one of the most important constituents in
the food web of lake fauna and also the main
intermediate host of trematode parasites.
This gastropod is relatively large and is nu-
merically and functionally a dominant mem-
ber of the second trophic level of the macro-
fauna of Lake Kerkini. Consequently, a study
on the life cycle, population dynamics,
growth, and secondary production of this
prosobranch snail was considered valuable.

MATERIALS AND METHODS

The abiotic characteristics of the study
area have been studied in detail, including
water level, precipitation, water temperature,
pH, dissolved oxygen, chloride, water hard-
ness, and PO,-P, in relation to the density
and growth of V. contectus (Eleutheriadis &
Lazaridou-Dimitriadou, in press.). Presented
here are the water level of Lake Kerkini at
weekly intervals during the study period (Fig.
1), which affects the marshes around the lake

42 ELEUTHERIADIS & LAZARIDOU-DIMITRIADOU

37

36

35

34

33

Water level

32

31

0
JFMAMJJASONDIJFM AM
1989

JJASONDJFMAMJJASOND
1990

1991
Months

FIG. 1. Water level at weekly intervals of artificial Lake Kerkini during the years 1989, 1990 & 1991.

where this snail lives, monthly water temper-
atures (Fig. 2) at the marshes, and monthly
changes of the concentration of dissolved
oxygen (Fig. 3), at the marshes, which seem
to affect the onset of hibernation and repro-
ductive period of this species.

The study started in October 1989 and
lasted for two years. Data from October 1989
to August 1991 were used for the demo-
graphic analysis of the populations of V. con-
tectus, which dominate the benthic, littoral
and sublittoral fauna. Samples were taken
using random coordinates (Lewis & Taylor,
1972) by use of a net “drag” based on the
Doulkeite sampler (illustrated in Lammote &
Bourlière, 1971) with a 1-m° surface, at ap-
proximately monthly intervals throughout the
year. Vegetation, debris and snails were hand
washed through 1-mm mesh sieves. Snails
were sexed (males have an enlarged right
tentacle used as a copulatory organ), mea-
sured and then returned as near as possible
to their initial places.

Measurements of the distance between
the apex of the shell and the farthest point on
the aperture (shell height) were made using
vernier calipers. In addition to the shell height
(H) of each specimen, the largest shell diam-
eter (D) and the perpendicular diameters of
the peristome (d & a) were also measured to
the nearest 0.1 mm.

The total number of samples were deter-

mined so that the percentage error (Table |)
was not more than 25% (Elliott, 1971).

Spatial distribution of the snails in the hab-
itat was examined by using Taylor's (1961)
power law. The parameter b from Taylor's
equation s* = ax” (where a = constant, $2 =
variance, х = mean number of snails found in
a sample unit, a and b = constants) was used
as an index of dispersion. Parameter b is
fairly constant and characterizes a species
(Southwood, 1966); it is independent of the
total number of samples and the total num-
ber of animals in the samples but 15 affected
by quadrat size (Elliott, 1971).

The class interval of the monthly size-fre-
quency histograms of shell height (H) was 3
mm, as determined by Goulden's method
(Cancela da Fonseca, 1965). Cohorts were
separated using probability paper (Harding,
1949). This method was valid because the
modes of the age classes were separated by
at least 2.5 standard deviations (Grant, 1989).
Although some age classes had less than 50
individuals, the modal values were consistent
from month to month, which confirms that the
modes were real and not the result of sam-
pling variation. This method has been used for
demographic analyses of the populations of
other molluscs (Hughes, 1970; Léveque,
1972; Daguzan, 1975; Lazaridou-Dimitriadou,
1981; Lazaridou-Dimitriadou 8 Kattoulas,
1985; Staikou et al., 1988, 1990, 1991).

LIFE CYCLE OF VIVIPARUS CONTECTUS 43

1990

FIG. 2. Monthly water temperatures at the marshes of Strymonas River from July 1989 to April 1992.

20
>
E
Е
>
> 10
O
8
>
o
2
TD
0 $ с
8 10
1989

FIG. 3. Monthly concentrations of dissolved охудеп (D.O.) at the marshes of Strymonas River from July

1989 to April 1992.

For the study of absolute growth, data
from the modal distribution of V. contectus
were used (e.g., the growth of one age class
was followed by the growth of the same age
class the following month taking into consid-
eration that time intervals always had to be
equal). For determination of the theoretical
growth curve, Bertalanffy's (1938) equation
was employed: H, = Ha, [1 e KE
where H, = the largest shell height at age t,
Hmax = the asymptotical maximum possible
largest shell height, К = growth rate coeffi-
cient, t = time in months, and t, = hypothet-
ical time when Н is equal to “zero.”

The coefficient К and H,„.. were deter-
mined according to Walford’s (1946) method.
Hmax is the intersection point of the growth
curve H,,, = f(H,) and the line drawn at 45°
through the zero point. The coefficient К 1$
equal to —loga* 2,30259 (where a = the slope
of Walford’s line). For the determination of
the date of birth of an age class on the time
axis, it is possible to use a secondary origin

(Г = 0) corresponding to the smallest snails
found in the biotope during the study period
of a species (in our case was H = 5.5 mm),
assuming that all the small snails of this spe-
cies have been captured with the same size
and that all age classes follow the same
growth laws. Consequently, it is possible to
draw the theoretical growth curve of shell
height in relation to time from the first capture
Hy = Hmax [1 — e 9], If the shell height
at the moment of birth is known from labora-
tory data (in our case was H = 3.5 mm), the
axes may be changed taking as origin birth
(zero point) (to = t’-t”). So Bertanlaffy’s
equation becomes H, = Hina, [1-е “* ‘J
and the life span of the studied species until
H max Can be estimated.

For the study of relative growth, the mor-
phometric criteria of shell diameter (D) in re-
lation to the perpendicular diameters of the
peristome (d & a) were used from all the an-
imals sampled during 1990 (Table |. Mayrat's
method (1965 a, b) was used to compare the

’

44 ELEUTHERIADIS & LAZARIDOU-DIMITRIADOU

growth of the shell diameter (D) in relation to
the peristome surface (P.) between immature
and mature male and female snails. Because
the peristome was almost an ellipse, its sur-
face area (P.) could be calculated by the for-
mula (Rey = 3.14: Ча

Annual production in 1990 was calculated
by the size-frequency method, because it
has an important advantage in that single co-
horts need not be identified to calculate pro-
duction (Krueger & Martin, 1980), although it
may produce an overestimate (Waters 4
Crawford, 1973). This method has been used
for the determination of gastropod produc-
tion, because it gives similar results to those
obtained by Russell-Hunter's (1970) method.
Both methods have been compared for B.
graeca (Eleutheriadis 8 Lazaridou-Dimitria-
dou, submitted). The formula as modified by
Benke (1979) and Krueger & Martin (1980)
can be written as:

Z (nn) (VW Win)"

P=a

+ ia.) - 365/CPI,

where Р = the mean annual production in mg,
a = number of size classes, п; = number of
snails at the size class | in number, Un; =
variance of ñ,, W, = mean individual dry body
weight+mean dry shell of organic material (in
mg), and CPI = cohort production interval in
days.

For the determination of the variance (U) of
P the following formula was used:

— 22] С? И.
U(P)=a*| GF U(n:)+ Z (G¡-G;¡ +) U(n;)
¡NA j

+ (W,—Ga-1)2U(na) |: (365/CPI)?,

where G = geometric mean of weight of pairs
of successive size classes. For the determi-
nation of n; and Un;, data from the population
dynamics of V. contectus were used and
snails collected during 1990 were grouped
into 15 size classes of 3-mm intervals.

To determine dry body weight and the dry
organic shell material, 60 snails representing
all size classes were brought to the labora-
tory in April 1990, where their shell height (H)
was measured in mm and they were dried in
vacuo at room temperature; dry body weight
(W) in mg was measured one week later. Dry
organic shell material was also estimated af-
ter successive treatments of immature and

mature shells (mature: H > 17 mm and imma-
ture: H < 17 mm, according to the study of
relative growth) with 5N HCl, as described in
Staikou et al (1988). Size classes >18 mm
contained two females and two males (ex-
cept for the last size class, which contained
only females), in the sex ratio 1:1 as in nature,
and in the size classes from 3 to 18 mm 20
snails were examined (in total 60 snails).

On March 1991, some females were re-
turned to the laboratory for observations on
birth. There the mean number of new-born
snails released per female was determined.
We also measured the mean shell height of
new-born snails. The differentiation of the
genitalia in relation to the age of the snails
was also studied under a stereoscope. The
maturation of sectioned female and male go-
nads in relation to the age of the snails were
studied under a light microscope.

RESULTS
Aspects of the Biology

The snails were sexually mature when the
largest shell height (H) exceeded 17 mm, as
verified by following gonad maturation. Ma-
turity was attained three months after birth if
the new-born appeared in early spring, but
eight months if they appeared in autumn. The
sex ratio in the three consecutive years of
study (1989, 1990, 1991) was equal to 1:1. In
1989, new-born snails appeared in October
when water level was high (Fig. 1) and the
water temperature was > 15°C (Fig. 2). In the
same season in October 1990, because wa-
ter level was low (Fig. 1) and water tempera-
ture < 15°C (Fig. 2), females carried fully
formed embryos through the winter and gave
birth during spring 1991. No females smaller
than 16 mm shell height contained develop-
ing embryos. The mean number of new-born
snails released per female was 16 + 8.8 (n =
8) and the new-born snails had a mean shell
height 5.15 + 0.58mm (n = 61). Fully devel-
oped young were found throughout the year
in the brood sacs of the adult females. Fe-
males attained larger sizes because of their
greater longevity. The snails in this popula-
tion were active during spring, summer and
until the end of autumn. At the end of autumn
and during winter, snails were buried in the
mud of deeper water (1-2 т). The population
began to move into shallower waters by early

НЕЕ CYCLE OF VIVIPARUS CONTECTUS

45

TABLE 1. Viviparus contectus population density in the marshes of Strymonas River from July 1989 to
August 1991 (п: number of samples; X: mean number of animals/m?; s: standard error).

1989
6/7 21/7 2/8 17/8 20/9 2/10 19/10
n 40 31 38 33 22 28 40
x 0:15 1.52 1.26 112 2.45 1253 0.77
$ 0.26 0.38 0.25 0.29 0.50 0.34 0.17
Percentage error D 34.6 25:2 20.0 26.3 20.3 22.1 22.4
1990
22/4 13/5 26/5 10/6 29/6 21/7 4/8 29/8 13/9
п 28 24 24 24 24 20 24 16 16
x 1.64 РУ 215 3.29 2.62 12 2.91 215 2.68
5 0.48 0.45 0.43 0.67 0.61 0.34 0.47 0.6 0.53
Percentage error D 29.7 16.7 Wes 20.1 23.4 28 16.5 21.6 19.7
1990 1991
24/9 22/10 7/11 24/3 21/4 16/5 3/6 26/6
п 16 22 24 14 8 16 15 21
x 2:5 1.59 1.41 4.78 8.75 3 2 2
Ss 0.74 0.48 0:37 1.02 2:23 0.6 0.47 0.32
Percentage error D 28.9 30.5 25.8 21.4 25:5 19.2 22.8 16.1
1991
12/7 1/8 24/8
n abe) 11 14
x 2.4 1.3 2.43
$ 0.52 0.56 0.83
Percentage error D 21.4 32 34

spring and occupied the highest 50 cm of
water. Migration to deeper waters occurred
again between October and November, so
that over-wintering of all members of the
population occurred in water of 1-2 m depth.

Population Density and Spatial Distribution

Population density fluctuated during the
study period, either from month to month or
from year to year (Table 1). Low values ap-
peared in July and in the end of autumn each
year, whereas high values were recorded in
March and April in 1991. The mean popula-
tion density of V. contectus during the study
period 1989-1991 was 2.84 + 2 (mean + stan-
dard deviation) snails m ?.

The spatial distribution of V. contectus was
found to be contagious, because parameter
b of Taylor’s power law was equal to 1.43 ($2
17189):

Demographic Analysis of the Population of
V. contectus

The analysis of size frequency histograms
(Fig. 4) from October 1989 to April 1991 with
probability paper showed the following (Fig.
5):

(a) Two cohorts were found in the habitat
throughout the year; a third was added after
the release of newly born individuals.

(b) In 1989, the newly born individuals ap-
peared in mid-autumn and in 1991 the new-
born appeared in the beginning of spring.

(c) Increased growth rate for newly born
young occurred during spring and in the be-
ginning of summer. Growth was continuous
until the end of autumn.

(d) One year after the new-born appeared,
the largest shell height was about 28 тт;
two years later, the snails were about 35 mm
(March 1991) and the third year about 37-40
mm (Gas).

46

ELEUTHERIADIS 8 LAZARIDOU-DIMITRIADOU

307 % 10/89
20
10
О 6 12 18 24 30 36 42
% 4/90
40
20
0
O 6 12 18 24 30 36 42
205 % 5/90
10
0
0 6 12 18 24 30 36 42
59] 2 6/90
20
10
0 6 12 18 24 30 36 42
30- + 7/90
20
10
0 6 12 18 24 30 36 42

304 + 8/90
20
10
0 6 12 18 24 30 36 42
307 0, 9/90
20
10
0 6 12 18 24 30 36 42
SU ca 11/90
20
10
O 6 12 18 24 30 36 42
407% 3/91
30
20
10
O 6 12 18 24 30 36 42
60 + % 4/91
40
20
O 6 12 18 24 30 36 42

401 % 5/91
20
O 6 12 18 24 30 36 42
SO Tg 6/91
20
10
0 6 12 18 24 30 36 42
407 + 8/91
30
20
10
O 6 12 18 24 30 36 42

D (mm)

FIG. 4. Size frequency histograms of Viviparus contectus in the marshes of Strymonas River from October
1989 to August 1991.

H(mm)

Hibernation

Hibernation

10
Months

11

Reproductive period

'

3/91 4 5 6 7

30%

55%

FIG. 5. Population analysis of the populations of Viviparus contectus at marshes of Strymonas River from
October 1989 to August 1991. Percentages denote the contribution of each cohort to the total population.
(G88 to G91 indicate when a generation started and when it ended). Dotted lines represents a decrease in
shell height (H) because of the death of the largest individuals.

LIFE CYCLE OF VIVIPARUS CONTECTUS 47

TABLE 2. Estimation of statistical parameters of the population of Viviparus contectus
[where a,b: constants, r: coefficient correlation, N: number of snails, D: the mean shell
diameter, Ps: 1/10 d - а, (а, а = the perpendicular diameters of the peristome, о: standard

deviation)] from Teissier's regressions.

Data Entire sample
a+o 0.511 =0.037
logb +0 0.657 + 0.047
г? 0.957

logD + o 1.304 + 0.137
logPs +o 1.266 + 0.268
М 819

Relative Growth

The study of the relative growth of D in
relation to Р. (for practical reasons we used P.
as 1/10 : d a) showed a positive correlation
between D and P, (r? = 0.957, п = 819) (Fig. 6).
Knowing that gonad differentiation was com-
plete (first appearance of spermatozoa and
mature oocytes) when the largest shell diam-
eter reached 13.5 mm (corresponding to 17
mm shell height), it was decided to examine
whether relative growth rate was the same in
the two size groups, that is those with D< 13.5
mm and those with D > 13.5 mm. A logarith-
mic transformation was applied to the data
because the coefficient of correlation was
higher than for raw data (0.957 and 0.950 re-
spectively). A statistical difference (P < 0.01)
was found between the slopes of the two re-
gression lines using Mayrat's method (1965а,
b) (Table 2). The intersection point of the two
regression lines, corresponded to D = 14.9
mm, near to the size that sexual maturity was
attained (Fig. 6). No statistical difference was
found between the regression lines of mature
female and male snails.

Absolute Growth

Knowing that after maturity growth rate was
the same between male and female V. con-
tectus, we decided to study absolute growth
in all snails. H,,,,., which represents the inter-
section point of Walford equation of a straight
line (H,,, = 0.941H, + 3.087) and the diagonal
H,=H,,,, was 52.3 тт. By using the slope of
the line (a = 0.941), which showed the growth
rate of the animals, the coefficient К was са|-
culated as 0.06. Knowing the minimum H of
the measured snails in the biotope during the
study period (H = 5.5 mm), the growth of the
snail shell height was calculated by Bertalan-
ffy's equation H, = 52.3 [1 — е %:08('+1.84)]
Because it was known from laboratory data

immature snails mature snails

0.421 + 0.012 0.556 + 0.0055
0.735 + 0.008 0.594 + 0.007
0.937 0.932
1.019 + 0.10 1.333 + 0.103
0.673 + 0.235 1.326 + 0.186
79 744

that the minimum H of new-born snails was
about 3.5 mm and their age t”, the age from
zero point t, = (Г — t”) was calculated. By
starting the curve at birth (Zero point), when
snails had their smallest H (equal to 3.5 mm),
the theoretical growth curve of H in rela-
tion to age was calculated: H, = 52.3
[1-е °`95+1-17)]. From this curve (Fig. 7), it
was found that V. contectus may live up to five
years before reaching its maximum size ac-
cording to Von Bertalanffy’s equation.

Secondary Production

The calculations of the size-frequency
method are listed in Table 3. The mean bio-
mass of each size class 15 expressed in dry
weight.

Applying Benke's correction, values of n
(mean annual density), B (mean annual crop)
and P (annual production) were calculated to
be 3 individuals/m?*, 4.19 g dry body weight/
п?.уеаг and 13.45 + 12.9 g dry body weight/
m*.year respectively. The annual turnover ra-
tio P/B was 3.21 and the turnover time was
113.7 days.

DISCUSSION

Populations of Viviparidae from various
habitats appear to differ in a number of life
history traits. Intraspecific and interspecific
differences exist in size of new-born snails,
number of broods/year, size of largest males
and females, time of birth, and life span of
females (Chaberlain, 1958; Samochwalenko
8 Stanczykowska, 1972; Young, 1975; Ber-
nardi et al., 1976; Browne, 1978; Vail, 1978;
Jokinen et al., 1982; Buckley, 1986; НЫ &
Gebhardt, 1986). Viviparus contectus is iter-
oparous and viviparous. Birth begins in mid
autumn or in the beginning of spring. Spring

48 ELEUTHERIADIS 8 LAZARIDOU-DIMITRIADOU

-.2 .2 .4 .6

.8 1 5 1.2 1.4 1.6 1.8
Ps=10.9 mm

FIG. 6. Relative growth of the shell diameter in relation to Ps of shell peristome in the whole population and
in mature and immature snails of Viviparus contectus (Teissier's regression lines).

55 4 mm

Shell height

Months

FIG. 7. Theoretical growth curve of Viviparus contectus.

reproduction 1$ also reported for other spe-
cies ofthe Viviparidae by Van Cleave & Cham-
bers (from Vail, 1978), Van Cleave & Altringe
(from Vail, 1978), Chaberlain (1958), Fretter &
Graham (1962), Young (1975), Bernardi et al.
(1976), Browne (1978), Vail (1978) and Jokinen
et al. (1982), possibly because in this season
the environmental conditions favour survivor-
ship and rapid growth of the new-born snails.
In contrast to most other molluscs, fecundity
of У. contectus is low, reflecting the conse-
quences of viviparity, as has also been re-
ported for V. georgianus by Browne (1978).

The number of new-born snails released per
female is variable. This number in the genus
Viviparus ranges from 2.5 to 90 snails. In
Poland, V. contectus released 4.5-10.2
new-born snails per female (Samochwalenco
8 Stanczykwoska, 1972), whereas in our
study, the number of newborn snails was 16
+ 8.8 (n = 8). Selection probably drives female
toward a larger size than males as a conse-
quence of the cost of viviparity. The repro-
ductive output of smaller females is not only
limited bioenergetically, but smaller size
places severe physical constraints on the

LIFE CYCLE OF VIVIPARUS CONTECTUS 49

TABLE 3. Calculation of production of Viviparus contectus by the size-frequency method. Annual
production based on sets of samples from April 1990 to April 1991 (where п; = number of snails at the
size class j in number; Un, = variance of п; W = mean individual dry body weight+mean dry shell of
organic matter (in mg); G, = geometric mean of weight of pairs of successive size classes; В = mean
standing crop or population biomass in mg; P = annual production in mg; P/B = annual turnover ratio; a
= number of size classes; CPI = cohort production interval.

Class —
range n,/m? Un; N, NA W, (mg)
3-6 0.42 0.0224 — 0.04 0.042
6-9 0.45 0.6475 0.22 0.100
9-12 0.23 3.7573 0.19 0.238
12-15 0.04 0.0157 0.01 0.406
15-18 0.06 0.0023 — 0.06 0.575
18-21 0.12 0.0196 —0.01 0.742
21-24 013 0.0191 — 0.01 0.910
24-27 0.14 0.0047 — 0.02 1.078
27-30 0.16 0.0469 — 0.03 1.376
30-33 0.19 0.0240 = 0.12 1.870
33-36 0:31 0.0975 —0.09 2.381
36-39 0.39 0.0554 0.14 2.780
39-42 0.26 0.0897 0.18 3.390
42-45 0.08 0.0185 0.05 3.970
45-48 0.02 0.0013 0.02 4.490
365 days 3

(B) Pp
__G [njWi] (N, ñ,,:)G)

(WW...) (mg m?) (mg m?)
64.8 ir — 2.5
154.3 45.4 34.7
310.9 54.6 58.1
483.2 17.3 —7.1
653.2 33.0 — 39.9
821.7 87.9 —12.0
990.4 12.1 Or
1217.9 153:0 — 18.8
1604.1 216.5 —4.9
2087.8 351.3 =250.3
2545.6 717.4 219.1
3069.9 1094.9 420.8
3668.6 870.4 659.1
4222.0 306.0 225.9
4490.0 105.9 105.9

4192.1mg/m? 897

P =а.Р”-365/СР! = 15:897-365/365 = 13455 mgm * or 13.455 gm ? in 365 days. U(P) = UN,(G;G,1)* - (865/CPI)? - a? =

41730400

Confidence limits of P = P + 2-[U(P)9*] = 13.455 + 12.92
P/B = 13.455/4.192 = 3.21

Turnover time = 113.7 days

number of embryos that can be maintained in
the uterus if discrete embryo size units are to
be maintained. By contrast, males following
the strategy of attempting to mate with as
many females as possible would have only
limited time for feeding, resulting in slow
growth and a much shorter life span than fe-
males. This has also been reported for V.
georgianus by Browne (1978).

Viviparus contectus in Lake Kerkini under-
take distinct seasonal migrations into deeper
water in November following the drop in
marsh temperatures, and they migrate back
into shallow water in early spring. Although
other species of Viviparidae appear to mi-
grate before a decrease in temperature
(Stanczykowska & Magnin, from Jokinen et
al., 1982), the decline in temperature appears
to trigger a migration of V. contectus into
deeper water in the marshes, bringing the
snails from summer feeding areas into the
hibernation area of 2 m depth. Snails migrate
to avoid areas of low temperatures (Skoog,
1971; Horst & Costa, 1975; Vincent et al.,
1981) and unfavourable ecological condi-
tions in the surface water (Coulet 8 Alfaro-

Tejera, 1985). Other factors that provoke mi-
gration might be seasonal habitat changes
(Lilly, 1953; Jokinen, 1985), changes in water
level (Skoog, 1971), food availability (Russell-
Hunter, 1953), or water currents that flow
faster as vegetation dies back (Lilly, 1953;
Fretter 4 Graham, 1962; Boss et al., 1984;
Lodge et al., 1987). Additionally, snails mi-
grate to areas of macrophytic vegetation as a
preferred habitat for the release of new-born
snails or egg deposition, as shown for Amni-
cola limosa by Horst & Costa (1975). The re-
cruitment of newly born snails to the popula-
tion during the breeding season in March and
April 1991 was the main reason for the rise in
population density in these months. By con-
trast, the low numbers found at the end of
autumn were probably due to migration to
the undersampled marsh bottom. The de-
crease after the breeding season was prob-
ably due to mortality of the new-born snails.
According to Taylor's law, the spatial distri-
bution of V. contectus was contagious, and
this behaviour, as Bovbjerg (1965), Duch
(1976) and Brown (1979) have demonstrated,
is characteristic of freshwater snails that tend

50 ELEUTHERIADIS & LAZARIDOU-DIMITRIADOU

to aggregate on filamentous algae and large
diatom clusters. Bithynia graeca in Lake Ker-
kini also shows a contagious distribution
(Eleutheriadis 8 Lazaridou-Dimitriadou, sub-
mitted). Growth of V. contectus stopped at
the end of autumn and in winter for about 4
months. Growth pauses have also been re-
ported for V. georgianus by Jokinen et al.
(1982) and Buckley (1986) in New York. In
contrast, Young (1975) for V. viviparus in En-
gland, Bernardi et al. (1976) for V. ater in Italy,
and Browne (1978) for V. georgianus in the
USA noted that growth was continuous
throughout the year, although growth rate
varied seasonally. Rapid growth during
spring may be due to the favourable water
temperatures (Fig. 2) and oxygen concentra-
tions (Fig. 3) that prevailed during this period
in the Strymonas marshes. Water level and
water temperature seem to influence the on-
set of birth. Fully developed young were
found throughout the year in the brood sacs
of the adult females, so it appears that adult
snails choose the most favourable environ-
mental conditions for the release of young
snails. The fact that both internal changes in
this species concerning the maturation of the
gonads correspond to external morphomet-
ric changes in the shell agrees with results
reported for other prosobranch species of
the family Littorinidae (Daguzan, 1975), for
Monodonta lineata (Daguzan, 1991), and for
B. graeca (Eleutheriadis 4 Lazaridou-Dimitri-
adou, submitted). Viviparus 1$ a genus show-
ing considerable variation in the duration of
life. Viviparus contectus has a multiyear life
cycle, and there are some differences in the
pattern of the cycle within the genus. Such
European investigators as Samochwalenco &
Stanczykwoska (1972) reported that V. con-
tectus and V. viviparus lives up to 4 years;
Young (1975) reported that V. viviparus lives
up to 2 years, and Ribi & Gebhardt (1986)
reported that V. ater lives 5 to 8 years. A Ca-
nadian population of V. malleatus was re-
ported to live for 5 years (Stanczykwoska et
al., 1971). In the USA, populations of V. geor-
gianus were reported to have a 2 to 3 year
life-span (Van Cleave & Lederer, from Vail,
1978; Browne, 1978; Jokinen et al., 1982)
and 4 years (Buckley, 1986). These studies
indicate that Viviparus life span may be a trait
determined by habitat factors. The compari-
sons in production among freshwater snails
were done by use the turnover times, be-
cause productivity rates are very sensitive to
difficulties of assessing environmental space

in calculating densities for biomass (Russell-
Hunter & Buckley, 1983). The turnover time
for V. contectus is short and reflects relatively
high levels of productivity. The value of 113.7
days is low compared with values from other
freshwater prosobranchs. For four popula-
tions of V. georgianus, the turnover time was
477, 510, 393 and 421 days (Browne, 1978),
for Bithynia tentaculata 337 and 314 days for
females and males respectively (Tashiro,
from Russell-Hunter & Buckley, 1983), and
for three population of Leptoxis carinata 372,
311 and 303 days (Aldridge, 1982). All these
populations are at least biennial. Low values
were reported by Lévéque (1973) in Lake
Chad for three annual prosobranchs snails
including 74.5-117 days for Melanoides tu-
berculata, 101-126 days for Cleopatra buli-
moides, 56-70.2 days for Bellamya unicolor,
and in Lake Kerkini 77.9 days for an annual
population of В. graeca (Eleutheriadis 4
Lazaridou-Dimitriadou, submitted). The dif-
ference in turnover time of V. contectus com-
pared to other biennial and triennial popula-
tions must be due to lack of competition with
other freshwater snails; in two years” study,
only small numbers of Valvata piscinalis and
Lymnaea stagnalis were recorded. There 1$
also little direct human intervention on these
marshes.

Viviparus contectus, being an A-strategist,
seems to be able to profit the favourable con-
ditions even when the climate shows a sud-
den difference and its parameters do not
follow the characteristic cycle of the Mediter-
ranean climate at the marshes of Strymonas
River.

ACKNOWLEDGEMENTS

We would like to thank Dr. G. Dussart from
Canterbury Christ Church College for his
comments on an earlier draft of this paper and
the two anonymous referees of this journal.

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Revised ms accepted 28 Nov 1994

MALACOLOGIA, 1995, 37(1): 53-68

COMPARATIVE LIFE CYCLE AND GROWTH OF TWO FRESHWATER
GASTROPOD SPECIES, PLANORBARIUS CORNEUS (L.)
AND PLANORBIS PLANORBIS (L.).

Katerine Costil & Jacques Daguzan

Laboratoire de Zoologie et Ecophysiologie (L. А. INRA & URA 1853), Université de Rennes |,

Campus de Beaulieu, Av. du General Leclerc, 35042 Rennes Cedex, France

ABSTRACT

The shell growth and life cycle of two natural populations of Planorbarius corneus and
Planorbis planorbis were studied for 16 and 17 months, respectively. These planorbid popu-
lations lived in ponds where they exibited great annual density variations, which could be
related to environmental conditions, snail behaviour and, especially, their life cycle. The pop-
ulation of Planorbis planorbis showed a life cycle with two breeding periods per year, and
consequently, two generations. Whereas both generations were semelparous and lived for 11
months, they grew with different seasonal patterns. But of the two species, Planorbarius cor-
neus growth seemed more influenced by the climate. For the population of this species, an
annual life cycle with two generations was observed in 1987, whereas only one spring gener-
ation seemed to be present in 1986 and 1988. In 1987, the vernal generation, showing an
estimated maximum life span of 18-20.5 months, was potentially iteroparous, and the estival-
autumnal generation living 15-21 months was basically semelparous. Life-cycle variations of

natural populations of freshwater snails are reviewed.
Key words: shell growth rate, life cycle, Planorbidae, population dynamics, reproduction.

INTRODUCTION

Life cycles of freshwater snails are of the
highest importance for the study of their bi-
ology because they can be viewed as a syn-
thesis of the main life-history traits of snail
populations. Such studies have been per-
formed for planorbid temperate species (Rus-
sell-Hunter, 1961; Boerger, 1975; Young,
1975; Eversole, 1978; Alfaro Tejera, 1982;
Lodge & Kelly, 1985; Byrne et al., 1989; Ca-
quet, 1993). Planorbis planorbis has been re-
ported to show an annual life cycle (De Coster
8 Persoone, 1970; Dussart, 1979), whereas
the life cycle of P. corneus is almost unknown.
In England, a population of P. corneus was
studied for only four months (Berrie, 1963).
Moreover, freshwater snails present great in-
traspecific variation in life-history patterns
(Calow, 1978; Brown, 1985), and this plastic-
ity has been considered to be of fundamental
selective value in their evolution. (Russell-
Hunter, 1961). An observed life history is the
result of both long-term evolutionary forces,
and the more immediate responses of an or-
ganism to the environment in which it is and
has been living (Begon et al., 1990).

The present data constitute part of a set of
results on the biology and the ecology of Ar-

53

morican freshwater gastropods (Costil, 1993,
1994; Costil 8 Daguzan, 1994). The aim of the
present study was to determine the life cy-
cles of the populations of P. corneus and P.
planorbis in order to compare them with life-
history strategies of other freshwater snail
populations. Special attention was given to
snail shell growth.

MATERIAL AND METHODS
Study Sites

The population of P. corneus was studied
for 17 months in pond Le Boulet, located 35
km north of Rennes, east Brittany (48°20’N,
1°38’W). The population of P. planorbis living
in pond La Musse, 35 km _ southeast of
Rennes (48°00’N, 1°58’W), was sampled for
16 months. These continuous studies were
preceded and followed by instantaneous ob-
servations made in 1986 and in 1989.

The sampling areas in the large ponds (Le
Boulet: 120 ha.; La Musse: 49 ha.) were the
shallowest (depth of 20-50 cm) and the most
eutrophic parts. Vegetation contained nu-
merous aquatic and amphibious macro-
phytes (Clement, 1986; Costil, 1993), and the
bottom was covered by mud. Table 1 sum-
marizes the results of water chemical analy-

54 COSTIL & DAGUZAN

TABLE 1. Water chemistry in the two study ponds; the unit is mg/l except for conductivity (uS/cm) and
pH (pH unit); S.M. = matter in suspension; the analyses were performed in May 1988.

Total
S.M. CON СЕР” Mg?* pH Total iron mangenese
Le Boulet 80 160 13:8 6.0 ES SU 1.4
La Musse 150 160 12.6 4.3 8.7 MOS ES
№№. N-NO, N-NH,* Total P-PO,? Total
Or 10m nitrogen (035 phosphorus
Le Boulet 1253 1.4 9.0 21 2.1 5.6 107
La Musse 0.39 1.3 8.0 1.86 0.9 2:4 11078

ses. Because of human interferences, the
water level was subject to great variations (up
to + 180 cm at the study site in La Musse).
Planorbarius corneus lived with 10 coexisting
gastropod species (4 planorbids, 3 lym-
naeids, 1 physid and 2 prosobranchs),
whereas P. planorbis shared its habitat with 7
species (3 planorbids, 3 lymnaeids and 1
physid). In both ponds, birds and fishes that
include molluscs in their diets were present,
e.g. Anas platyrhynchos L. and Aythya
fuligula L., Rutilus rutilus L. and Abramis
brama L.

Sampling

Samples were taken fortnightly from May
5, 1987, to October 18, 1988, for P. corneus,
and from April 14, 1987, to August 16, 1988,
for P. planorbis. Sampling was sometimes
impossible because of ice cover or flooding.
Random sampling took place in a limited
zone of 50 т? surface. In mid September
1987 in pond La Musse, the deterioration of
the environment leading to the disappear-
ance of the studied population obliged us to
transfer the study site from the south to the
north bank, a few meters away. The snails,
however, from both banks belonged to the
same population.

The snails were collected using a pond net
(1 mm mesh) and a wood frame delimiting a
sampling surface of one square meter. This
sampling method was chosen especially be-
cause it respects the environment and allows
long-term study. On every sampling date, we
tried to collect at least 100 individuals. The
number of sampled т? depended on the
snail density and environmental conditions,
and fluctuated from 3 to 10 for P. planorbis,
and from 3 to 15 for P. corneus. The number
of collected snails sometimes reached 380
(P. planorbis) and 389 (P. corneus), but sam-

pling difficulties (particularly ice cover or
flooding) did not always allow us to collect
100 snails. In the field, living snails and empty
shells were counted (рег m*), and measured
(maximum shell diameter) using a vernier cal-
liper (to the nearest of 0.05 mm). The snails
were then returned to the study site, while the
empty shells were removed. Egg capsules
were sampled to determine periods of repro-
duction.

Sample Treatment

The results were recored in 1-mm size
classes. The number of individuals in each
size class was expressed as a percentage of
total number, establishing size-frequency
histograms. The cohorts, represented by
gaussian structures, were discriminated us-
ing Bhattacharya's (1967) method. On every
sampling date, a mean shell diameter (+ stan-
dard error) was calculated for each cohort.
Each cohort was assumed to be representa-
tive of one generation resulting from a spe-
cific reproductive period. The empty shells
were attributed to a particular cohort if pos-
sible.

The observed life span of a cohort corre-
sponded to the time from recruitment until
death. For the “real life span,” we had to add
the time necessary for the snails to reach the
recruitment size. This time was known if the
water temperature in the field and the growth
results of the laboratory experiments were
taken into consideration (Costil, 1994).

Von Bertalanffy’s model was used to de-
scribe the growth of the vernal and estival or
autumnal cohorts (Von Bertalanffy, 1938). It is
given by the equation:

D, = De [t= ee

where D, is the diameter at time t; О„.„„, the
asymptotic maximum diameter; K, a growth

max

PLANORBID LIFE CYCLES 55

rate coefficient; t,, the hypothetical time
when О is equal to “zero” (minus the duration
of the egg stage). К and L,,,,, are determined
according to Ford-Walford's method by the
slope and intercept on the x-axis of the re-
gression line of growth rate on size, respec-
tively.

RESULTS
Population Density

The mean densities of P. corneus and P.
planorbis were respectively 20.0 (S.E. = 3.5)
and 37.8 (S.E. = 5.9) individuals/m”, but great
annual variations were noted for both species
(Fig. 1). The maximum density was the same
(127 individuals/m?), but it was observed at
different dates (P. planorbis: 07-07-87; P.
corneus: 20-10-87).

Life Cycle

Two reproduction periods were found, a
vernal for both species and an autumnal or
estival-autumnal (16 weeks) for P. planorbis
and P. corneus, respectively (Table 2). For P.
planorbis, autumnal reproduction was only
observed on two sampling dates. On Octo-
ber 13, 1987, the site was inaccessible and
perhaps egg capsules were present (repro-
duction period for a minimum of two, possi-
bly up to four weeks).

The percentages of empty shells in relation
to sampled live snails varied from 0.4 to
26.8% for P. corneus, and from 0.0 to 32.4%
for P. planorbis (Fig. 2). They were especially
high in July 1987 for the first species, and in
August 1987 and in April 1988 for the latter.

For two consecutive years, the maximum
shell diameter of the population of P. corneus
was observed in May (20.9 mm on 19-05-87
and 18.9 mm on 31-05-88), whereas the min-
‘imum (6.9 mm) was noted in March 1988. For
P. planorbis, the minimum and maximum
shell diameters were 4.2 mm (23-05-87) and
11.8 mm (26-05-87), respectively. These
changes in mean diameter of the population
reflected the succession of cohorts that ap-
peared, grew and died. Each cohort repre-
sented a generation stant after a particular
reproductive period and descended from the
same parent generation.

At the beginning of the study, the popula-
tion of P. corneus corresponded to cohort
G1, the mean diameter of which was 18.9

mm (Fig. 3). One month later, the diameter
reached 23.7 mm and the snails reproduced.
Some individuals of the cohort G1 lived until
September 22, and some until mid Novem-
ber. The very large shells (25-28 mm) were
thicker and more eroded than the large shells
(22-23 тт). Cohort G2 appeared on June 2,
1987, and the largest snails laid egg capsules
in summer 1987. The individuals of G2 over-
wintered and, after May, except for observa-
tions made on September 20, 1988 (24 indi-
viduals in a restricted area), the number of
survivors was low. Cohort G3 appeared on
August 25, 1987, continued to be alive until
the end of the study. Individuals of G3, but
also survivors of G2, which had laid egg cap-
sules in previous summer, were the parents
of cohort G4. This cohort, recruited on May
31, seemed not to reproduce, although three
small snails were collected on October 4,
probably belonging to cohort G5.

In the case of P. planorbis, the population
showing a mean diameter of 8.6 mm on April
14, 1987, was mainly made up of one cohort
(G1) (Fig. 3). This cohort coexisted with a
more aged cohort (GO; mean size of 15.7
mm), which was no longer found in the mid-
dle of June. G1 retained a relatively large
number of snails until June 9, when the first
newly hatched snails of cohort G2 appeared.
After ovipositing in autumn (mean diameter of
8.2 mm), the individuals of G2 did not disap-
pear, and a lot of them were collected on
March 15. Snails of G3 were seen for the first
time on October 27, 1987. However, consid-
ering their relative large size and the period
when egg capsules were observed, it
seemed that recruitment had already oc-
curred on October 13. (On this date, we
could not sample because of flooding.) In
spring 1988, individuals of G3 gave the co-
hort G4. From July 5, 1988, the snails of co-
hort G3 contributed less than 10% to the to-
tal population.

For P. corneus, the observed life span of
cohort G2 was 70 weeks (about 16 months).
The number of these snails was sometimes
very low or null on certain sampling dates.
Moreover, the individuals of cohorts G1 and
G2 were very similar in size at the beginning
of May for two consecutive years. So, G2 ap-
peared to be homologous to G1, and its
members could survive six weeks after Oc-
tober 4, as was the case of G1 in 1987. The
observed life span of G2 was estimated to be
from 70 to 76 weeks. If the correction factor
and the size of G2 on the recruitment date

56 COSTIL 8 DAGUZAN
180 A

120

Population density (snails/m2)
00
©

8 as = a 8 as as A =F == AS AAs E am as Aa
5-6. 7_8 9 101112 1234 5 6 7 в
Study dates (days & months)

180

160

120

Population density (snails/m2)
5

Study dates (days € months)

FIG. 1. Changes in population density with time: Mean + S.E.. A: Planorbarius corneus (from April 1987 to
October 1988); B: Planorbis planorbis (from May 1987 to August 1988).

were taken into consideration, the “real life snails belonging to cohort G3 measured 24
span” was then from 79 to 85 weeks (18 to mm, that is, 2 mm lower than the maximum
20.5 months). At the end of the study, the size noticed for G2. Their growth curve did

PLANORBID LIFE CYCLES 57

30 А
25
e
a 20
©
<=
on
> 15
dd
о.
Е
4
æ 10
о
o
г
5
0
8 as = an an as as as =8 == A Aas = m= ae Ss as =
5. “6 2711869, 10: 11121234 5 6 7 8 9 10
Study dates (days & months)
= B

No. of empty shells (%)
S

м Ро.
Study dates (days & months)

FIG. 2. Variation of the number of collected empty shells in relation to the number of live snails collected.
A: Planorbarius corneus; B: Planorbis planorbis.

58 COSTIL & DAGUZAN

TABLE 2. Features of reproduction period (В. P.) in Planorbarius corneus and Planorbis planorbis; the
water temperatures, recorded on every study date at midday, correspond to minimum and maximum

values during the reproduction periods.

Water
Start of the End of the Duration temperature
Rip: R.P. (weeks) (°C)
Planorbarius 19-05-87 16-06-87 4 16-20
corneus 28-07-87 17-11-87 16 9.5-28
17-05-88 28-06-88 6 15-22
Planorbis 28-04-87 23-06-87 8 12-22
planorbis 15-09-87 29-09-87 2 14.5-18.5
(13-10-87?) (4?)
11-05-88 19-07-88 10 15-29

not show a plateau, typical of an asymptotic
growth. Moreover, the individuals of G3 were
not found in samples collected during spring
1989. The observed life span was estimated
to be from 60 to 80 weeks. The “real life
span” varied then from 67 to 87 weeks (15 to
21 months).

For P. planorbis, the cohort G2 showed an
observed life span of 44 weeks and a “real
life span” of 48 weeks. The population study
was stopped in mid-August 1988, when five
individuals of G3 were collected. If we con-
sider G3 as the homologue of G1, we could
say that G3 lived until the beginning of Sep-
tember 1988. So, the cohorts G2 and G3 had
the same observed (44 weeks) and “real life
span” (48 weeks or 11 months).

Growth of the Vernal and
Estival-Autumnal Cohorts

In P. corneus, strongly different growth
patterns were noted, as evidenced by the co-
horts (Fig. 4). In the first sample, the mean
diameter of individuals of cohort G2 was 6.4
mm. After 6 weeks, it reached 53% of the
maximum size, that 1$, 13.8 mm, and the
maximum growth rate attained 2.1 mm/week
(Fig. 5). A second peak (1.5 mm/week) was
observed for planorbids one year old. On the
other hand, from December 15 to April 19
(age: 29-45 weeks), shell growth gain did not
exceed 2.3% (rates below 0.035 mm/week).
For the growth of G3 cohort in the field, two
periods appeared to be particularly favour-
able: at the time of cohort recruitment (rates
higher than 1.05 mm/week), and from April
19 to mid-June (age: 29-45 weeks, mean
rate of 1.2 mm/week, and so the size
changed from 13.6 to 20.1 mm). During the
winter and in July and August, shell growth
was slow or null. The mean growth rate was

slightly higher for G3 than for G2 (0.35; S.E. =
0.09 and 0.29; S.E. = 0.08). Nevertheless, in-
dividuals of both cohorts reached a size of
about 25 mm at the end of their life.

The growth rate coefficient of Von Berta-
lanffy's model (К) was equal to 0.046. For the
entire population of P. corneus, the shell
growth in terms of individual age was given
by the relation:

О; = 37.31 1 = et SIE
time unit of 14 days.

The theoretical diameter for maximum age
estimated in the field (32.4 for snails 20
months old) was higher than maximum size
observed in the field (28 mm) (Table 3).

In the case of P. planorbis, the snails of
cohort G2 reached 65% of their maximum
size eight weeks after recruitment; maximum
rates were of 0.85 and 0.80 mm/week (Figs.
4, 6). Thirty-two weeks were necessary to at-
tain 89% of their size before growth started
again and the death of the cohort. For the
cohort G3, a long recruitment period influ-
encing the growth rates was observed. The
snails reached the two-thirds of the maxi-
mum size after 26 weeks, and two peaks
(more than 0.7 mm/week) were noted in
spring and in summer 1988. The mean
growth rates of the cohorts G2 and G3 were
0.21 (S.E. = 0.05) and 0.19 (S:E- = 0:05), mn
week, respectively.

The theoretical shell growth of the popula-
tion in relation to snail age was given by the
following relation:

D, = 13.4 1 = & о:
time unit of 14 days.

According to Von Bertalanffy's model, the
maximum mean diameters of G2 (11.3 mm)
and G3 (11.6 mm) conferred on these cohorts
a life span of 11-12 months (Table 3).

PLANORBID LIFE CYCLES 59

Egg laying
NS

Diameter (mm) A

30
25
20
15
10

1987 Study dates (days & months)

Egg laying
ER
Diameter (mm) B

15

10

1987 1988
Study dates (days & months)

FIG. 3. Changes in mean shell diameter of Planorbarius corneus (A) and Planorbis planorbis (B) (+ S.E.) with
time (from May and April 1987 to October and August 1988, respectively), at the Le Boulet and La Musse
ponds respectively. The numerals correspond to percentages denoting the contribution of each cohort to
the total population. For P. corneus, G1, G2 and G4 represent the spring generations, and G3 the estival-
autumnal one. For P. planorbis, the spring generations are GO, G2 and G4, whereas the autumnal gener-
ations are G1 and G3.

COSTIL 8 DAGUZAN

©
o

—#— Cohort G2
=--0-- Cohort G3 A

A
/
Pooog “voce ps
$

Diameter/maximum diameter (%)
in
©

0 5 10 15 20 25 30 35 40 45 50 55 6 6.
Observed age (weeks)

Diameter/maximum diameter (%)

0 5 10 15 20 25 30 33 40 45
Observed age (weeks)
FIG. 4. Relation between the snail age and the percentage of maximum diameter attained by the individuals

belonging to the vernal (C2) and the estival or automnal (C3) cohorts. A: Planorbarius corneus; B: Planorbis
planorbis.

PLANORBID LIFE CYCLES 61

Cohort G2

Water temperature Growth rates
(°С) (mm/week)
26 Sp Su f Ww Sp Su 22
a PAT а [20

| 1,8
1,6
1,4
152
1,0

0.5 10 15 20 25 30 35 40 45.50 55: 60 65 70
Observed age (weeks)

Cohort G3

Water temperature Growth rates
(°C) (mm/week)
26 Su f Ww Sp Su f 722

247 % : 2,0
22 a+ gt 1,8

h , у 1,6

1,4

1,2
1,0
RN
Осьеэх 19:15 20725.30 355.04 60
Observed age (weeks)

0,8
0,6
0,4
02
0,0

FIG. 5. Variation of the water temperature at Le Boulet pond, and changes in growth rates for the vernal (C2)
and the automnal (C3) cohorts of Planorbarius corneus. Sp = spring; Su = summer; F = fall; W = winter. The
value corresponding to C3 snails at the age of 13 weeks is put in brackets because it is an abnormal value
due to sampling difficulties.

62 COSTIL 8 DAGUZAN

TABLE 3. Theoretical relation between the age and the shell diameter (D) of Planorbarius corneus and
Planorbis planorbis according to Von Bertalanffy's model, H = hatching.

Р. Age (months) H 1 3 6 12 18 20

corneus D (mm) 172 4.0 10.3 11742 26.3 31.2 32.4

22 Age (months) H 1 3 6 10 12

planorbis D (mm) 0.8 РЕЙ 5:5 8.4 10.6 11.4
DISCUSSION nately declines as the water level increases.

Density Variations

Variation in population structure appears to
be a major factor explaining variation in snail
density. In our study, the recruitment of newly
hatched snails had a strong effect on the den-
sity, whereas the cohort disappearance af-
fected the density more or less progressively.
We do not know exactly how long the resi-
dence time of the empty shells in the studied
ponds is and when the corresponding snails
were dead. Such a topic has not been exam-
ined much. Hunter (1990) has reported that
adverse water chemistry (low pH and/or low
calcium concentration) has a much greater
effect on shell dissolution than does presence
orabsence of periostracum. Moreover, empty
shells do not have an absolute value, because
they could accumulate in certain areas. Nev-
ertheless, the percentages of empty shells
and above all their variation helped us to de-
termine the life cycle of both populations, and
to explain the variations in their density. For P.
planorbis, great numbers of empty shells and
low densities could be attributed to death of
cohorts. In summer, the disappearance of G1
and G3 took place, whereas in spring the dis-
appearance of G2 occurred. On July 14 and
28, 1987, the density of P. corneus was low
and the percentage of empty shells high (20-
27%). The reproductive effort of the largest
individuals seemed to be responsible for their
death, whereas the high mortality of small in-
dividuals might be dueto predation. The pred-
ators seem to be small invertebrates unable to
eat whole shells, such as insects or leeches,
because a lot of empty shells were pierced.
Nevertheless, the major predation on fresh-
water snails 15 exerted by vertebrates (Lodge
et al., 1987). According to Eisenberg (1966),
the whole predation could lead to the death of
93% to 98% of the young Lymnaea elodes
(Say).

The snails were sampled using a pond net
which respects the environment, but the ef-
ficiency of this sampling method unfortu-

This is especially illustrated in winter for P.
corneus, and on June 7, 1988, when the den-
sity of P. planorbis population was less than
6 snails/m” at the flooded study site. On the
other hand, during dry periods, snails could
be seen on the mud and easily collected. Such
was the case in March 1988 for P. planorbis
and in September 1988 for P. corneus. Snail
behaviour also influenced their density. When
the environmental conditions became unfa-
vourable in winter, we observed some indi-
viduals of P. corneus sinking into substratum
crevices or even into mud. It was then very
difficult to collect them. Such behaviour has
been reported for Lymnaea catascopium (Say)
(Pinel Alloul, 1978). At the same time, some
individuals of P. corneus were seen moving on
the ice cover on December 1 and 15, 1987.
Nevertheless, the extreme climatic conditions
(cold and hot) appeared particularly unfavour-
able for the population density of P. corneus.
In summer, lack of dissolved oxygen into wa-
ter was a problem, as was the case in late July
1987, when chlorophytes pullulated at the
study site and no individual of P. planorbis
was sampled. Moreover, Eisenberg (1966)
emphasized the importance of food for the
regulation of density in a natural population of
L. elodes. The death and then the regrowth of
submerged macrophytes led to changes in
the densities of those snails inhabiting them
(Lodge et al., 1987). Lymnaea peregra and
Valvata piscinalis (Müller) exhibited low resis-
tance to habitat disturbance (decline of sub-
mersed macrophytes), but high adjustment
following the disturbance. The maximum den-
sity for P. corneus (127 snails/m*) was noted
three days after a storm occurred in Brittany
with winds of 140 km/h. The accumulation of
individuals of Anisus rotundatus (Роге) in lim-
ited areas attributed to wind was also re-
ported by Marazanof (1970).

Life Cycle

Different reproductive periods were deter-
mined according to the cohort to which the

PLANORBID LIFE CYCLES 63

Cohort G2
Water temperature Growth rates
(°С) (тт/меек)
28 Su f Ww Sp 1,0

0 3 10 15 20: “25 30 35 40 45
Observed age (weeks)

Cohort G3
Water temperature Growth rates
(°С) (mm/week)
287 № Ww Sp : su 1,0
26 N:
24 A 6
> Pa 0,8
20 |
18
16 | oP
14
12
10 0,4
8
6 0,2
4
/\
0 0,0

0 5 10 15 20:7 25 307 35 40 45
Observed age (weeks)

FIG. 6. Variation of the water temperature at La Musse pond, and changes in growth rates for the vernal (C2)
and the automnal (C3) cohorts of Planorbis planorbis. Sp = spring; Su = summer; F = fall; W = winter.

64 COSTIL & DAGUZAN

majority of the parents belonged. In the stud-
ied population of P. corneus, the egg-laying
period began in May and ended in Novem-
ber, but we accepted two periods. The esti-
val-autumnal reproduction period lasted for
16 weeks (the parents belonged to the cohort
G2). However, at the end of this period, we
could not exclude an egg-laying by the larg-
est individuals of G3 estival cohort (13 mm on
November 17, 1987). Moreover, no egg cap-
sules were found in August or in October
1987. Perhaps some were in the field, but it
was very difficult to see them among the
dense vegetation. The length of the egg-lay-
ing period was due to the inter-individual
growth variation. In L. catascopium, all the
snails belonging to spring generation were
not mature at the same time, and they con-
tinued to reproduce as long as the water tem-
perature allowed it (Pinel Alloul 4 Magnin,
1979). According to Berrie (1963), the repro-
duction period of P. corneus was short. On
the other hand, De Coster & Persoone (1970)
sampled very small P. corneus (0-2 mm) from
April to September. Concerning P. planorbis,
the spring reproduction periods were longer
than the autumnal period. In northwest En-
gland, P. planorbis also showed two egg-lay-
ing periods, one in February and the other in
July (Dussart, 1979).

According to Precht (1936), P. corneus did
not reproduce in autumn or in winter, and an
endogenous rhythm had to be suggested, be-
cause the snails brought in the laboratory at
23°C did not lay eggs either. This author
added that P. corneus began to reproduce
when the temperature reached 12°C. The
minimum threshold temperature for the re-
production of freshwater snails appears to be
between 7°C and 12°C (Boerger, 1975; Dun-
can, 1975; Eversole, 1978). Three egg cap-
sules were laid by P. corneus reared at 8°C
(De Wit, 1955). In our study, the reproduction
period of both species began when temper-
ature reached 15-16°C, which was above the
minimum threshold. In spite of suitable tem-
peratures, the genital organs could be imma-
ture. Such a result was observed by Russell-
Hunter (1961) in Gyraulus albus (Müller).
Moreover, the freshwater snails could be
more sensitive to temperature variations than
to absolute temperatures.

The main life-cycle patterns encountered in
natural populations of freshwater snails have
been described by Russell-Hunter (1961,
1978) and reviewed by Calow (1978). During
our study, P. planorbis showed a type C as
defined by Calow (1978), that is, an annual life

cycle with reproduction periods occurring in
spring and in autumn with parent generation
replacement. The snails of the vernal cohort
G2 did not survive until the next reproductive
period. However, these individuals coexisted
with the snails of the autumnal cohort for 28
months. Both cohorts corresponding to two
generations were semelparous, and their life
span was estimated at 11 months. A life span
of 12-13 months and an annual life cycle have
been reported respectively by De Coster &
Persoone (1970) and Dussart (1979). How-
ever, the second author described a life cycle
with egg-laying periods completely brought
forward in comparison with our results (re-
cruitment of newly hatched snails in February
and July instead of early May and September).
Such an annual life cycle with two generations
per year occurred in the following species:
Lymnaea peregra (Müller) (Russell-Hunter,
1961; Lambert, 1990), Lymnaea palustris
(Müller) (Hunter, 1975), Physa fontinalis (L.)
(De Wit, 1955; Russell-Hunter, 1961), Heli-
soma trivolvis (Say) (Eversole, 1978); Bathy-
omphalus contortus (L.) (De Coster & Per-
soone, 1970), Armiger crista (L.) (Alfaro Tejera,
1982), and A. rotundatus (Marazanof, 1970).
For the latter planorbid species, Caquet
(1993) observed a semelparous life cycle with
a maximum life span apparently reaching
17-18 months.

From a four-month study, an annual life cy-
cle was attributed to P. corneus (Berrie, 1963).
According to different authors, the life span of
this species in the field was 13 months
(Hilbert, 1911), 2 or 3 years (Boycott, 1936;
Boettger, 1944), and 4 years (Frómming,
1956). Russell-Hunter (1978) has reported
that biennal life-cycle patterns involve such
larger pulmonates as P. corneus and Lym-
naea stagnalis (L.) in higher latitudes or more
oligotrophic waters (1.е., poorer temperature
or trophic conditions). Like for P. planorbis,
the life cycle of P. corneus in 1987 was annual
showing two generations. The estival-autum-
nal generation was not replaced by the new
generation (type B). So, other differences
were also observed between the two species:
longer life span of both generations and an
annual life cycle close to a biennial (18-20.5
months for G2; 15-21 months for G3, these
life spans could be very similar); a longer sec-
ond reproductive period; a spring cohort
probably iteroparous to a certain extent and
an estival-autumnal cohort probably semel-
parous (the second egg laying period proba-
bly missing in 1986 and 1988). At the begin-
ning of May 1987, only one cohort was

PLANORBID LIFE CYCLES 65

observed (G1). Its mean diameter was the
homologous of the spring cohort G2 diame-
ter at the same date in 1988. In autumn 1988,
no egg capsule was present at the sampling
site, although three small snails were col-
lected on October 4 (mean diameter of 6.1
mm). The second reproduction period might
occur every other year or might be far less
predictable, depending on annual climatic
variations. In the pond Le Boulet in 1988, the
water temperature was apparently not as
high as in 1987, and shell growth was slower
than in 1987. The climatic hypothesis was put
forward by Vincent & Harvey (1985) to explain
both types of life cycle (short and long) en-
countered in a Canadian population of Bithyn-
ia tentaculata (L.). The number of genera-
tions per year can also be related to the
length of dry periods (Duncan, 1959; Ma-
razanof, 1970). Lymnaea catascopium exhib-
ited an annual life cycle with one generation
in hard or medium waters, and two genera-
tions in soft waters (Pinel Alloul, 1978). In L.
peregra, snails from the exposed habitats
matured earlier and put more effort into it
than snails from the sheltered habitats; these
differences in growth and reproduction could
be explained in terms of differences in selec-
tion pressure between habitats of varying
exposure (Calow, 1981). Individuals of H.
trivolvis from the eutrophic sites grew faster
and exhibited an annual life cycle, whereas
those from the mesotrophic site grew slowly
and lived about two years, breeding in both
summers (Eversole, 1978). From reciprocal
transfer experiments, Brown et al. (1985)
concluded: “genetic divergence among pop-
ulations of L. elodes explained a compara-
tively small proportion of the variation in life
histories in comparison with proximal factors
like habitat productivity. Nevertheless, snails
from the vernal pond always grew more
slowly, matured at a smaller shell length, and
had higher fecundities than other popula-
tions. These smaller differences may still be
important over evolutionary time scales.”
These examples show that life-cycle varia-
tions can be adaptative. The adaptative plas-
ticity allows the populations to have maxi-
mum productivity in given conditions and
then to compensate for bad years (Huben-
dick, 1958; Russell-Hunter, 1961).

Snail Growth

The shell growth of freshwater pulmonates
is continuous until death (indeterminate

growth). For both studied species, the growth
differed according to time when a cohort had
been recruited. The shell growth did not only
depend on age of the animals, but the ob-
served differences could be due to various
physiological potentialities that remain to be
studied. For B. tentaculata, growth rates fluc-
tuated from 0.17 mm/week to 0.80 mm/week
with the seasonal trophic conditions (Pinel
Alloul 4 Magnin, 1971). In L. peregra and L.
palustris, the vast majority of the interpopu-
lation variation of shell growth rate appeared
to result from non-genetic ecophenotypic en-
vironmental influences and in particular from
habitat productivity (Byrne et al., 1989). Cli-
mate seemed very important for growth, es-
pecially in the case of P. corneus.

Spring is very favourable to the shell
growth of P. corneus and other species (Ma-
razanof, 1970; Pinel Alloul, 1978). In June
1987, a marking experiment (with painting)
allowed us to observe that some shells had
grown from 0.75 of whorl to 1.75 whorls in
two weeks (diameters attaining respectively
10.1 and 18.1 mm). In spring, the shell growth
of young Lymnaea humilis (Say) was very fast
reaching 7% per day (MacCraw, 1961). Dur-
ing autumn, the studied snails grew with in-
termediate rates. In summer (July-August)
and in winter (December to March), a lower
rate of shell growth was noticed for L. pere-
gra (Lambert, 1990) and P. corneus inhabit-
ing Le Boulet pond. In summer, the high tem-
peratures (up to 26°С) monitored probably
exceeded the optimum, which was close to
20°С for the growth of the two studied spe-
cies reared in the laboratory (Costil, 1994).
For a population of P. corneus in England,
Berrie (1963) recorded a mean size increase
of 4% between July and August, and in-
creases of 70% and 33% respectively be-
tween June-July and August-September. In
our study, the growth of individuals belong-
ing to cohort G2 slowed down at the same
period that the snails reproduced. Concern-
ing the homologous cohort G4, no summer
reproduction period was observed in 1988,
but growth did not slow down. These facts
about the cohorts G2 and G4 suggested the
energy allocation theory which, on the other
hand, was not confirmed by the cohort G3
(growth slowed down after reproduction pe-
riod). A slow or null shell growth during winter
has been reported for many freshwater snails
(Russell-Hunter, 1961; Calow, 1973; Vincent
et al., 1981). Individuals of В. contortus sam-
pled in winter and then brought to the labo-
ratory grew fast again (Calow, 1973). Natural

66 COSTIL 8 DAGUZAN

population of aquatic molluscs can exhibit
degrowth (decrease in unit mass of structural
proteins through time) (Russell-Hunter,
1985); according to this author, degrowth
can be associated with reproductive or sea-
sonal stress, an example being the overwin-
ter starved individuals of Р. corneus, which
lost 44% of their tissue biomass over 126
days (with 33% mortality). For the same spe-
cies, Emerson (1967) has reported similar
losses: 62.3% with 56% mortality after 58
days of starvation; at this time, 95% of the
original polysaccharides, 49% of the proteins
and 22% of the lipids were metabolized. A
very omnivorous diet could be responsible
for the winter shell growth observed in P. plan-
orbis and also L. peregra (Russell-Hunter,
1961), Physa integra and P. gyrina (Clampitt,
1970). In the diets of both P. corneus and
Planorbis carinatus (Müller) (close to P. plan-
orbis), detritus formed over 85% of the food
ingested and sand grains about 10% (Reav-
ell, 1980). In comparison with Р. corneus, Р.
planorbis seemed far less affected by the cli-
matic variations. The spring cohorts (G2 and
С4) grew the fastest in July and August. Their
renewal of growth at the end of life could be
due to the late death of the largest individu-
als.

According to Von Bertalanffy's model, the
largest individuals of P. corneus in the field
were 14 months old. However, this model
does not take into account the environmental
conditions and their variations. lt appears
more suitable for P. planorbis than for P. cor-
neus. The first species showed higher growth
constant (k = 0.069) than the second (k =
0.046), which illustrates a faster develop-
ment. For comparison with other freshwater
snails, some values of k are given (same time
unit: 14 days): 0.024 for A. crista (Alfaro Te-
jera, 1982), 0.057 to 0.485 for snails from
Lake Tchad (Lévêque, 1971), 0.095 for L. per-
egra (Lambert, 1990).

The maximum size reached by animals in-
habiting in a given site depends on many fac-
tors, such as environmental conditions, but
also genetic makeup and parasitic infection.
Before death, the cohorts G1 and G2 of P.
corneus respectively measured 27.4 or 25.9
mm, and the size of the largest individuals
reached 28 mm. These sizes were lower than
the value that is usually given for this species
(35 mm). According to the cohorts of P. plan-
orbis, the mean diameter attained before
disappearance fluctuated from 11.3 to 14.3
mm. The maximum diameter recorded was

16.1 mm, whereas the maximum size re-
ported for this species is 20 mm. It is difficult
to know to what extent the environment con-
ditions influence the maximum size of the
snails of a given species. In both ponds, the
water mineralization was rather low, and
flooding and drying occurred quite often,
making the environment unstable. According
to Byrne et al. (1989), the extended adult sur-
vival and oviposition was a life-history trait
that allows L. palustris to survive in marginal,
unstable habitats. Such an observation could
be made for P. corneus showing a potentially
itereparous cohort and a very long annual life
cycle, but not for P. planorbis. Nevertheless,
for both species the spring cohort growth
was rapid enough to allow a second repro-
duction period per year (every year in the P.
planorbis population and in 1987 for P. cor-
neus). Moreover, the trophic conditions in
both studied ponds appeared favourable for
the coexistence of a great number of gastro-
pod species. The environmental factors are
of the highest importance in the life-history
strategies of the freshwater snails. The great
range of strategies used by these snails т-
habiting various types of environment make
them particularly useful for studies about ev-
olutionary biology.

ACKNOWLEDGMENTS

We are grateful to Maria Lazaridou-Dimitri-
adou for her comments on early draft of this
manuscript. We also thank Stacy Payne for
checking the english and Jean Luc Foulon for
technical help.

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chiology, 30: 253-271.

RUSSELL-HUNTER, W., 1961, Life-cycle of four
freshwater snails in limited populations of Loch
Lomond, with a discussion of intraspecific vari-

ation. Proceedings of the Zoological Society of
London, 137: 135-171.

RUSSELL-HUNTER, W., 1978, Ecology of fresh-
water pulmonates. Pp. 355-383, in: V. FRETTER &
J. PEAKE, eds., Pulmonates, Vol. 2, Academic
Press, London.

RUSSELL-HUNTER, W., 1985, Physiological, eco-
logical and evolutionary aspects of molluscan
tissue degrowth. American Malacological Bulle-
tin, 3: 213-221.

VINCENT, В & М. HARVEY, 1985, Dynamique de
deux populations du gasteropode Bithynia ten-
taculata. Verhandlungen Internationale Vereini-
gung für Theoritische und Angewandte Limnol-
ogie, 22: 3288-3291.

VINCENT, B., G. VAILLANCOURT & М. HARVEY,
1981, Cycle de développement, croissance, ef-
fectifs, biomasse et production de Bithynia ten-
taculata L. (Gastropoda : Prosobranchia) dans le
Saint-Laurent. Canadian Journal of Zoology, 59:
1237-1250.

VON BERTALANFFY, L., 1938, A quantitative the-
ory of organic growth. Human Biology, 10: 181-
213:

YOUNG, М. R., 1975, The life-cycles of six species
of freshwater Molluscs in the Worcester Birming-
ham Canal. Proceedings of the Malacological
Society of London, 41: 533-548.

Revised Ms. accepted 15 December 1994

MALACOLOGIA, 1995, 37(1): 69-110

WHEN SHELLS DO NOT TELL: 145 MILLION YEARS OF EVOLUTION IN
NORTH AMERICA'S POLYGYRID LAND SNAILS, WITH A REVISION
AND CONSERVATION PRIORITIES

Kenneth C. Emberton

Department of Malacology, Academy of Natural Sciences of Philadelphia, 1900 Benjamin
Franklin Parkway, Philadelphia, Pennsylvania 19103-1195 U.S.A.

ABSTRACT

A phylogenetic hypothesis for the 274 known polygyrid species is presented, based on 115
anatomical, behavioral, and shell characters (39 of which are newly discovered or newly as-
sessed), as corroborated in part by published allozyme data. The hypothesis differs little from
that of Emberton (1994a), except for greatly increased resolution in the tribes Stenotremini and
Polygyrini. A corresponding revision is presented that assigns species to 50 subgenera, 24
genera, seven tribes, three infrafamilies, and two subfamilies. All taxa in the revision, as well as
unnamed clades, are defined phylogenetically by shared derived characters. Nine new subge-
neric names are introduced: Neohelix (Asamiorbis), N. (Solemorbis), Triodopsis (Brooksorbis),
T. (Pilsbryorbis), T. (Macmillanorbis), T. (Vagvolgyorbis), Stenotrema (Archerelix), S. (Pils-
brelix), and Millerelix (Prattelix).

A separate phylogenetic analysis of polygyrid subgenera was conducted based on shell
morphology alone, using 71 character states in 14 characters, all new or newly assessed,
including ontogenetic characters scored from shell x-rays. The resulting shell-based hypothesis
had only one-fourth the phylogenetic resolution of, and showed major topological discrepan-
cies from, the anatomy-behavior-shell hypothesis. Thus, at present level of knowledge, poly-
gyrid shells are inadequate for reconstructing phylogenetic history, and identifications of pre-
Miocene fossils should be considered dubious at best. Remaining to be evaluated, however,
are shell-surface microsculptures and ultrastructural layers.

Museum and field surveys discovered the closest known polygyrid convergences in sympa-
try on flat, umbilicate, and tridentate shell forms: Patera laevior and Xolotrema fosteri at Hawes-
ville, Harlan County, Kentucky; Appalachina sayana and Allogona profunda on Pine Moun-
tain, Hancock County, Kentucky; and Inflectarius inflectus and Triodopsis fallax in Vinton
County, Ohio. These provide starting points for analyses of naturally replicated experiments
in evolutionary morphology, such as those already conducted on the globose shell form
(Mesodon normalis and Neohelix major at numerous sites in the Southern Appalachian Moun-
tains).

Adult Triodopsis tridentata from which apertural barriers had been removed lost water
27% faster than controls when retracted, and 9% faster when extended. Controls were 83%
more successful than barrierless snails in forming epiphragms. Epiphragms reduced the rate
of water loss by 3% in controls and by 38% in barrierless snails. Ramsay's (1935) hypothesis
is extended to suggest that barriers and epiphragms slow evaporation not directly, but indi-
rectly by interrupting natural convection currents between the aperture and the retracted snail's
body. Because 7. tridentata and other triodopsins have larger barriers in more humid climates,
barriers should also serve at least one other function, such as impeding invertebrate predators.
A new hypothesis is proposed for the function of extremely obstructed shell apertures: to
exclude water, allowing the snails to float to safety during floods. Polygyrids have many con-
vergences on apertural obstruction by barriers, some extreme examples of which are illustrated
together.

Based on the phylogenetic hypothesis/revision, the most urgent remaining targets for
polygyrid conservation are (1) Colombia's Isla de Providencia, where deforestation threatens
the ancient, relic, uniquely live-bearing, endemic Giffordius; and (2) the northern coves of Pine
Mountain, Harlan County, Kentucky, which harbor North America's most diverse land-snail
communities, including four simultaneous cases of polygyrid convergence in sympatry.

Key words: cladistics, phylogenetic taxonomy, shell ontogeny, paleontology, fossils, con-
vergence in sympatry, functional morphology, Gastropoda, Pulmonata, Stylommatophora.

69

70 ЕМВЕАТОМ

INTRODUCTION

The Polygyridae are an endemic North
American family of land snails that are remark-
able for their shell convergences in sympatry
(due to iterative, punctuated evolution cou-
pled with phenotypic and genetic, adaptive
parallel environmental responses); their 145-
million-year biogeographic history (paralleling
that of, for example, plethodontid salaman-
ders); and their evolution of hermaphroditic,
external sperm exchange via several behav-
iorally bizarre intermediates that remain ex-
tant (Pilsbry, 1940; Webb, 1974; Solem, 1976;
Emberton, 1981, 1988a, 1991a,b 1994a,b,
1995a,b; Asami, 1988, 1993). Polygyrids cur-
rently comprise 271 nominal and three unde-
scribed species (see below), with a maximum
sympatric diversity of ten species reported
from Pine Mountain, Harlan County, Ken-
tucky, U.S.A. (Hubricht, unpublished; Ember-
ton, 19950), with many more species yet to be
described and discovered in Mexico and Cen-
tral America (F. Thompson, pers. commun.),
and probably with some undiscovered spe-
cies remaining in the Pacific Northwest (T.
Frest, pers. commun.). Phylogenetic hypoth-
eses for the family as a whole (Webb, 1974;
Pratt, 1979; Emberton, 1994a) have been to
genus only, and the most recent hypothesis
omitted shell characters completely and had
major unresolved polytomies.

Polygyrid fossil shells of the Pleistocene
and Pliocene are nearly all identical with
Recent species (Hubricht, 1985, and pers.
commun.; Emberton & Bogan, unpublished).
Miocene and later fossils are generally as-
signable to genus (Pilsbry, 1940; Auffenberg
8 Portell, 1992; Roth 4 Emberton, 1994). Ear-
lier polygyrid fossils, however, which are rare
but presumably valuable in reconstructing
the first 125 million years of polygyrid history,
could be difficult to identify because of multi-
convergent, punctuated-equilibrium evolu-
tion (Emberton, 1994a). One way of testing
the value of early fossils is to make an hon-
est, concerted attempt to reconstruct poly-
gyrid phylogeny based on Recent shells
alone. This has not previously been at-
tempted. One possible source of new and
useful characters for such an analysis—and
for fossil identification—is shell ontogeny as
viewed in x-rays (Ramirez, 1993). For exam-
ple, a recent study discovered, using x-rays,
a significant difference in whorl-expansion
rate between the polygyrid tribes Triodopsini
and Mesodontini (Emberton, 1994a). Thus a

more general x-ray survey of polygyrid shell
ontogeny could be productive.

Non-mimetic sympatric convergences are
important as naturally replicated experiments
in evolutionary morphology (Emberton,
1995a). Polygyrid non-mimetic convergences
in sympatry fall into four distinct shell forms
(Pilsbry, 1940) that have been called globose,
flat, tridentate, and umbilicate shell-static
clades (Fig. 1; Emberton, 1991b, 1994a). The
globose clades show the closest conver-
gences in sympatry (Solem, 1976; Emberton,
1981; Asami, 1988, 1993), of which the most
extreme 1$ between Neohelix major and Ме-
sodon normalis. Recent analyses ofthese two
species have yielded important new insights
into the ecological, genetic, and natural-
selective influences on shell morphology
(Emberton, 1994b, 1995a). The most extreme
cases of sympatric convergence on flat, tri-
dentate, and umbilicate shell forms have not
been analyzed or even reported, however.

Polygyrid shell evolution is further note-
worthy for its convergences in extreme aper-
tural obstruction by denticles and other shell
structures (Fig. 2; Pilsbry, 1940; Zilch, 1959-
60). Such apertural barriers appear in many
forms among numerous land-snail clades
(Zilch, 1959-60). Four hypotheses have been
proposed concerning the function of these
barriers: (1) to deter attacks by predatory in-
sects (Cook, 1895; Boettger, 1921, 1935;
Solem, 1972, 1974; Falkner, 1984); (2) to re-
tard evaporative water loss (Boettger, as
cited by Goodfriend, 1986; Rees, 1964; Paul,
1974; Christelow, 1992); (3) to strengthen the
aperture against accidental breakage (Paul,
1974); and (4) to orient the shell during crawl-
ing (Paul, 1974). A fifth hypothesis, proposed
here for the first time, is suggested by some
polygyrid cases of extreme apertural ob-
struction existing in flood-prone environ-
ments (Hubricht, 1985; Emberton, 1986): that
some barriers function (5) to trap air within
the submerged shell, preventing drowning
and enhancing gene flow by allowing the
snail to float downstream. None of these hy-
potheses has been adequately tested using
living snails (Goodfriend, 1986; pers. ob-
serv.).

Also untested is the likely hypothesis that
some barriers perform two or more functions
simultaneously. In the polygyrid tribe Triod-
opsini, there is a correlation in several lin-
eages— including that of the common spe-
cies Triodopsis tridentata—between greater
apertural obstruction and increased environ-

71

POLYGYRID SHELL EVOLUTION

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72 ЕМВЕАТОМ

FIG. 2. Convergent shell-apertural obstruction among various polygyrid subgenera. Species are (left to
right, top row): Triodopsis (Haroldorbis) henriettae (ANSP 109734), Triiobopsis гореп (150631), Ashmunella
mudgii (319743), Stenotrema (Stenotrema) unciferum (171138), Stenotrema (Stenotrema) maxillifer
(170141); (bottom row): Millerelix (Millerelix) tamaulipasensis (317942), Millerelix (Prattelix) plicata (143448),
Daedalochila (Upsilodon) hippocrepis (84629), Daedalochila (Daedalochila) uvulifera (10990), Inflectarius

(Inflectarius) rugeli (116893).

mental moisture (Vagvolgyi, 1966; Emberton,
1988a). This implies that if apertural barriers
retard evaporative water loss in T. tridentata,
then they should also serve at least one ad-
ditional function, such as deterring predators
or trapping air.

Polygyrid conservation is relatively ad-
vanced, with at least five rare taxa currently
listed as endangered or threatened: Patera
clarki nantahalae, Fumonelix archeri, F. or-
estes, F. jonesianus, and Triodopsis playsay-
oides. Thanks to this recognition, these taxa
are receiving the protection they need to sur-
vive. Phylogenetic analysis, however, can
provide insights into conservation priorities
beyond the rarity of individual taxa. Polygyrid
conservation has never been considered in a
phylogenetic context.

The purposes of this paper are (1) to con-
duct a phylogenetic analysis of the Polygy-
ridae to the species-group/subgeneric level;
to use the resulting phylogenetic hypothesis
(2) to taxonomically revise the family; (3) to
conduct an independent phylogenetic analy-
sis based on shells alone, incorporating
x-rays; to use the resulting cladogram (4) to
assess the reliability of fossils for recon-

structing polygyrid phylogeny; (5) to find and
report the closest polygyrid convergences in
sympatry on the flat, tridentate, and umbili-
cate shell forms; (6) to test for retardation of
evaporative water loss by shell apertural bar-
riers in living Triodopsis tridentata; and (7) to
assess remaining conservation priorities for
the Polygyridae from a phylogenetic per-
spective.

MATERIALS AND METHODS
General Phylogenetic Analysis

Standard phylogenetic methods of charac-
ter analysis were used (Hennig, 1966; Wiley,
1981; Wiley et al., 1991; Brooks & McLennan,
1991). The outgroups used were the type
genera of the bradybaenids, helminthoglyp-
tids, thysanophorids, camaenids, and sag-
dids, plus Cepolis to represent the xanthony-
cids (Emberton, 1991c). Data sources were
from anatomical and behavioral characters
compiled by Emberton (1994a: appendix A),
with added anatomical character analyses
using illustrations of Pilsbry (1940), Archer

POLYGYRID SHELL EVOLUTION 73

(1948), Metcalf & Riskind (1979), Pratt
(1981a), and Emberton (1988a, 1991a), and
shell character analyses using the collection
of the Department of Malacology, Academy
of Natural Sciences of Philadelphia (ANSP).
Shell characters were limited as much possi-
ble to those involving complex apertural
barriers. No anatomy of Pilsbry’s (1940) “Poly-
gyra plicata group” had ever been ade-
quately published (W. G. Binney's [1878:
plate 15, fig. I] sketch of the genitalia of “Poly-
gyra troostiana Lea” is uninformative and un-
trustworthy, appearing as it does beside a
totally inaccurate depiction of the genitalia of
“P. tridentata [Say]’’). Therefore, all the (lim-
ited) ANSP alcohol material of “Р. plicata
Say” was examined, and one adult was cho-
sen for dissection and illustration of the shell
and the reproductive system, which were in-
cluded in the general character analysis.

Based on experience with anatomical-shell
comparisons in the context of an anatomical-
allozymic phylogenetic hypothesis among all
species of both the Triodopsini and the Me-
sodontini (Emberton, 1988a, 1991a), anatom-
ically unknown polygyrid species were tenta-
tively assigned to subgenus Бу shell
characters alone. Thus, subgenus was used
as the operational taxonomic unit for this
phylogenetic analysis, even though any given
subgenus may have been represented by up
to eleven species that were each scored for
each character. For this reason, all '“autapo-
morphies” defining subgenera were retained
in the data matrix.

Polygyrid allozyme data (Emberton, 1988a,
1991a, 1994a) were not used in this analysis
because data were lacking for many subgen-
era and because existing data were from two
separate analyses involving only partially
overlapping sets of loci. Instead, relevant
portions of this phylogentic hypothesis were
visually compared for topological congru-
ency with previously published, allozyme-
based hypotheses (Emberton, 1988a, 1991a,
1994a).

Phylogenetic analysis was conducted by
hand and was interactive with construction of
the data matrix. The analysis progressed
along successively more restricted ingroups,
each of which was compared with its closest
possible outgroup. Thus, the resulting cla-
dogram and its coordinate subgenus-by-
character-state matrix were superficially free
of certain homoplasies that would have re-
sulted from a non-hierarchical, single-out-
group analysis.

Taxonomic Revision

The revision was based on the general
phylogenetic hypothesis and followed basic
principles of phylogenetic taxonomy (Queiroz
& Gauthier, 1990), which are becoming well
established among vertebrate systematists
and have recently been introduced to mala-
cology by Roth (1995). The method, as em-
ployed here, designates as taxa clades that
are defined by shared derived characters (sy-
napomorphies), regardless of subsequent
evolutionary modifications of those charac-
ters. For a discussion of the naturalness of
this method and its great advantages over
traditional taxonomy, see Queiroz & Gauthier
(1990).

Shell-Based Phylogenetic Analysis

Previous studies had shown the extents
and limits of intraspecific variation in polygy-
па shell morphology (Emberton, 1988a,
1988b, 1994a, 1994b), which were consid-
ered during character analysis. In general,
one intact adult shell of the type species was
chosen to represent each subgenus, but for
subgenera with extreme shell variation, two
or more representative species were se-
lected. Outgroups were represented by one
shell each of the type species of the type
genus of each of the six closest polygyrid
outgroup families (Emberton, 1991c), except
when the type genus happened to have a
highly derived shell morphology.

Shells were mounted in the planes simul-
taneously of both the rotational axis and the
aperture (Emberton, 1988a: fig. 29b, d) on
clear acetate sheets using thick rubber ce-
ment, then x-rayed over single-coat SR-5 In-
dustrex R film. Contact prints were made
from the x-ray negatives. Shell “apertures” at
half-whorl intervals were drawn, with accom-
panying 3-mm scale lines, from the contact
prints using a camera lucida mounted on a
Wild M-5 dissecting microscope. The draw-
ings were inked, reduce-xeroxed until all of
about the same size, and mounted in regular
array, arranged by a former classification
(Webb, 1974; Richardson, 1986) that has
since been revised (Emberton, 1994a, this
paper).

The demounted shells themselves, as well
as the x-ray drawings of their ontogenies,
were compared in a standard phylogenetic
character analysis (Wiley, 1981; Wiley et al.,
1991; Brooks & McLennan, 1991: chapter 2).

74 ЕМВЕАТОМ

Conchological differences known to occur
among closely related species within sub-
genera of the Triodopsini and the Mesodon-
tini (Emberton, 1988a, 1991a) were dis-
counted as characters.

The resulting shell-character by subgenus
matrix was analyzed phylogenetically using
Hennig86 (Farris, 1988), assigning equal
weights to all characters. A Nelson semi-
strict consensus tree (Farris, 1988) was com-
puted from the set of resulting, equally and
maximally parsimonious cladograms.

Reliability of Fossils

To qualitatively assess the reliability of fos-
sils for reconstructing polygrid phylogeny,
the shell-based and anatomy-behavior-shell-
based phylogenetic hypotheses were com-
pared for their degrees of phylogenetic res-
olution among subgenera. Resolution in both
cases was quantified as the number of nodes
relative to the maximum number of possible
nodes for the given number of taxa.

Closest Convergences in Sympatry

Closest potential polygyrid shell conver-
gences in sympatry on the flat, tridentate, and
umbilicate shell forms were searched for us-
ing Pilsbry (1940), Hubricht (1985), and the
collections of the Field Museum of Natural
History, Chicago. Actual sympatry was tested
by field work conducted during the spring
months of 1979, 1981, 1982, and 1983.

Shell Barriers and Water Loss

Twenty-five adult Triodopsis tridentata
were collected during five field trips in the
spring and fall of 1970. Twenty-three were
from Athens County, Ohio, and two from
Monongahela County, West Virginia. Initial
weights ranged from 0.39 to 1.06 gm. From
ten of these snails, all three apertural denti-
cles were completely removed using a table-
mounted dentist's drill. These barrier-less
nails were given 20 hours to recuperate from
the operation, and seven appeared at that
time to be uninjured and normally active.

All snails were weighed and placed into in-
dividual rubber-stoppered Erlenmeyer flasks
in which were suspended 10.0 gm о the des-
iccant calcium sulfate in a cheesecloth bag.
Temperature was maintained at 14-15°C in a
walk-in refrigerator. Each snail was weighed
on a tortion balance every hour for 18 hours.

Weighing took about two minutes per snail.
At each weighing, the snail's activity state
was recorded as (a) fully extended, (b) par-
tially extended, (c) retracted without epi-
phragm, (d) retracted with partial epiphragm,
or (e) retracted with complete epiphragm. Pe-
riods of individual fully extended activity were
also recorded when they occurred between
weighings.

Each recorded weight was expressed as a
percentage of the snail's initial weight, then
the percent decrease from the previous
hour's weight was calculated for each “snail-
hour” that proved usable. Mean percent de-
creases were calculated for each of eight ap-
erture/activity categories. Differences among
categories were expressed as percent
changes in rates of evaporative water loss.
The study was conducted some 25 years
ago, and since that time the raw data were
lost, so no statistical analysis could be per-
formed.

Remaining Conservation Priorities

Phylogeny-based conservation priorities
were assessed for the categories of (1) radi-
ating, endemic clades; (2) extremely autapo-
morphic, endemic taxa; (3) relic sister groups
to major clades; and (4) sites rich in conver-
gences in sympatry (Emberton, 1992). Prior-
ities in these categories were judged from the
general polygyrid phylogenetic hypothesis/
revision (this paper; Emberton, 1988a, 1991a)
and from known species distributions (Hu-
bricht, 1985) and sites of known high diver-
sity (Solem, 1976; Emberton, 1995c, unpub-
lished; Hubricht, unpublished). The current
conservation status of each high-priority
taxon or site was evaluated based on the
U.S. Federal Endangered Species List and
the locations of national and regional parks
and forest preserves.

RESULTS
General Phylogenetic Analysis

Table 1 defines the 115 characters used for
phylogenetic analysis, with references to il-
lustrations of the characters. | had not previ-
ously used or detected some of these char-
acters (39, or 34%) (Emberton, 1988a, 1991a,
1994a), primarily those dealing with (a) penial
morphology within the tribe Stenotremini
(Fig. 3), (b) reproductive-system morphology

POLYGYRID SHELL EVOLUTION 15

TABLE 1. Anatomical, behavioral, and shell characters used for a phylogenetic hypothesis (Fig. 8) and
a revision (text) of the Polygyridae. Citations refer to papers of Emberton, unless otherwise indicated.

OO —J O O1 R © ND —

. Nodulose fertilization pouch-seminal receptacle complex (1994a: fig. 2: character-state 2.2).

. Anteriorly united retractor muscles (1994a: fig. 2: character-state 3.2).

. Deeply excavated ureteric interramus (1994a: fig. 2: character-state 4.2).

. Single, dorsal, penial pilaster (1994a: fig. 2: character-state 5.2).

. Proximally swollen, internally lamellate vas deferens (1994a: fig. 2: character-state 8.2).

. Terminally papillate penial verge (1988a: figs. 2f, 3b, 4b, 5е, 7b, 7d, 8c).

. Two, flat terminal papillae on a terminal penial verge (1988a: fig. ба).

. Distinct but pustulate lappets (= transversely, partially fused pustules) on the penial pilaster

(1988a: figs. 2b, 5с, 58.

. Doubled density of pustulate lappets on the penial pilaster (1988a: figs. 5a, 50).
. Smooth lappets (= transversely, completely fused pustules) on the penial pilaster (1988a: fig.

2е).

. Halved density of smooth lappets оп the penial pilaster (1988а: figs. 2d, 4a).
. Smooth penial-pilastral lappets, plus in adulthood a short vas deferens (only about twice as

long as the penis) and a penial-retractor-muscle attachment near the penis (on the vas deferens
within one-third penis-length of the apex of the penis) (1988a: table 4).

. Greatly enlarged pustules on the penial pilaster (1988a: fig. 7).
. Penial-wall columns that merge mid-ventrally into 6-10 U-shapes that are tapered and slightly

separated and that bear unequally sized pustules (1988a: fig. 8).

. Penial-pilastral pustules forming a single column of abutting cubes (1988a: fig. 8a).
. A ventrally subterminal penial verge (1988a: fig. 7).
. Penial-pilastral pustules that are knob-like, unfused, and abruptly larger than the penial-wall

pustules (1988a: figs. 9a, 9c), or derivatives thereof.

. 15-20 penial-wall columns unmerging and radiating directly from the ejaculatory роге (1988a:

figs. 9a, 9c), or derivatives thereof.

. Club-shaped penis with a ventrally subterminal ejaculatory pore about 1/5-way from the apex

and indented into the penial wall (Webb, 1959: figs. 22, 27, 34, 38; 1988a: fig. 9), or derivatives
thereof (characters 28-30).

. Penial-pilastral pustules fused into two interdigitating columns of rectangular boxes (1988a: fig.

12).

. Penial-pilastral polygons 4-10 times the size of penial-wall pustules and armed with pustule-

sized knobs, or derivatives thereof (characters 22-24).

. Penial-pilastral polygons fused into a single mass or into large, irregular masses (1988a: fig. 11).
. Ventral penial-wall columns with pustules indistinct (1988a: fig. 11).
. Penial pilaster 3/4 the length of the penis and bearing polygons armed with blunt spurs (1988а:

figs. 13a, 14a, 14b, 16a).

. Indistinct pustules on the ventral-most radiating penial-wall columns (1988a: fig. 18b).
. Penial-wall columns merging mid-ventrally into 5-7 acute, equilateral, widely separated

V-shapes bearing equally sized pustules (1988a: figs. 14a, 15b, 16a).

. Extremely long, narrow penis (at least 25 times as long as wide) (1988a: fig. 13).
. Mace-shaped penis with a ventrally subterminal ejaculatory pore and with a sub-pore region

erectile as a fleshy peduncle, and derivations thereof (characters 29, 30).

. Ejaculatory-pore position approximately 1/4-way from the penial apex, peduncle small (Webb,

1948: fig. 4; Webb, 1959: figs. 14, 25а, 40, 43; 1988a: figs. 14a-b, 15-17).

. Ejaculatory-pore position approximately 2/5-way from the penial apex, peduncle large (Webb,

1959: figs. 12, 13, 15, 41; 1988a: figs. 14c-d, 18а, 180).

. A discernible clasping disc during mating (1994a: fig. 2: character-state 1.4), or derivatives

thereof.

. An unfanned origin of the penial retentor muscle (1994a: fig. 2: character-state 7.3).
. An epiphallus and flagellum (1994a: fig. 2: character-state 9.2), or derived loss thereof.
. А constriction in the epiphallus from the penial apex part-way toward the flagellum (1994a: fig.

2: character-state 9.3).

. А Незпу protuberance near the apex of the penis (Roth 8 Miller, 1992: fig. 6).

(continued)

76

EMBERTON

TABLE 1. Anatomical, behavioral, and shell characters used for a phylogenetic hypothesis (Fig. 8) and
a revision (text) of the Polygyridae. Citations refer to papers of Emberton, unless otherwise indicated.

(Continued)

36. Conical, non-papillate verge at the apex of the penis (Pilsbry, 1940: fig. 512Bb; Webb, 1970:
plate 35, fig. 5; Roth & Miller, 1993: figs. 19, 24).

37. Clasping disc during mating that is as voluminous as the inserted portion of the penis (1994a:
fig. 2: character-state 1.5), or derivatives thereof.

38. Minutely pustulate sculpture on the dorsal penial wall (1994a: fig. 2: character-state 5.4).

39. Paired dorsal penial pilasters (Pilsbry, 1940: fig. 496Aa, Bb’, D), or derivative thereof (character
42).

40. Rugose clasping disk (Webb, 1990: plate 6, figs. 1—4).

41. Nearly effaced paired dorsal pilasters (1994a: appendix B, character state 5.4).

42. Single dorsal pilaster, plus a clasping disc that is twice as voluminous as the inserted portion
of the penis (1994a: fig. 2: character-state 1.5).

43. One or more small, pointed, fleshy processes on the clasping disc, and derivatives thereof
(characters 44, 45).

44. Asingle large, pointed, fleshy process on the clasping disc (Pilsbry, 1940: fig. 506, 1d-e; Webb,
1948: figs. 3, 3a).

45. Two large, pointed, fleshy processes on the clasping disc (Pilsbry, 1940: fig. 510: 2a, 5a).

46. Clasping disc divided peripherally into two or three broad, unpointed lobes (Webb, 1965: plate
27, figs. 1-4).

47. Epiphallus longer than the prostate-uterus (Pilsbry, 1940: figs. 524, 525).

48. Chitinous, ornate spermatophore (Webb, 1954).

49. No penial insertion during mating (1994a: fig. 2: character-state 1.8).

50. Lateral pilaster(s) on the clasping disc (= basal penis) (1994a: fig. 2: character-state 5.5).

51. A (secondarily) slender spermathecal duct (1994a: fig. 2: character-state 10.4).

52. (Basal) penial lateral pilasters apically modified into two fleshy-walled cups half as long as the
penis (Fig. 3E).

53. Shell aperture with a complete basal lamella having a central trough half or more as broad as
the lamella, and derivatives thereof.

54. Shell bearing a straight, even-height parietal apertural denticle isolated from both the umbilicus
and the aperture (Fig. 6: character-state 5a), and derivatives thereof.

55. During mating, bearing an everted female organ that receives ejaculate from a pocket at tip of
the penis (1994a: fig. 2: character-state 1.8).

56. Apertural-basal-lamellar central trough about one-third or less as broad as the lamella (Fig. 6:
character-state 4b), and derivatives thereof.

57. Parietal apertural denticle extending from the umbilicus into the aperture (Fig. 6: character-state
5b), and derivatives thereof.

58. A single penial lateral pilaster apically modified into a symmetrical, fleshy-walled cup one-third
as long as the penis, with a medial branch leading to a medial fleshy protuberance (Fig. 3C).

59. Lower apertural shell lip joined to the basal body whorl as a thin callus (Fig. 5, character-state
3b), and derivatives thereof.

60. Two large, fleshy penial lateral pilasters, both bearing apical V- or U-shaped structures, one up
and one down (Fig. 3A).

61. Two large, fleshy penial lateral pilasters, neither bearing apical V- or U-shaped structures, and
one or none bearing an apical, cup-like depression (Fig. 3T, S, P), and derivatives thereof.

62. One of the two large, fleshy penial lateral pilasters bearing an apical, cup-like depression about
one-tenth as long as the penis (Fig. 31).

63. The two large, fleshy penial lateral pilasters free of apical, cup-like structures (Fig. 3S).

64. One of the two large, fleshy penial lateral pilasters bearing an apical, cup-like depression
one-fifth to one-half as long as the penis (Fig. 3P).

65. Penial-retractor-muscle insertion on or near penial apex (1994a: fig. 2, character-state 7.5-7.8).

66. Penial sheath (secondarily) completely absent (1994a: fig. 2, character-state 7.8).

67. Only a vestigial flagellum near the penial apex (1994a: fig. 2, character-state 9.5), and deriva-
tives thereof.

68. Shell bearing a triangular parietal denticle (Fig. 6: character-state 6a), and derivatives thereof.

69. Ovoviviparity (Pilsbry, 1930a).

(continued)

POLYGYRID SHELL EVOLUTION 1%

TABLE 1. Anatomical, behavioral, and shell characters used for a phylogenetic hypothesis (Fig. 8) and
a revision (text) of the Polygyridae. Citations refer to papers of Emberton, unless otherwise indicated.
(Continued)

70.

fale

Shell apertural expansion rate abruptly increasing then decreasing such that successive whorls
are nearly equal in volume (Fig. 6: character-state 2b), and derivatives thereof.

At least one full whorl of growth beyond the expansion-rate increase (Fig. 6: character-state
2c-right).

. Shell bearing a V- to U-shaped parietal denticle (Fig. 6: character-state 6b).

. Shell with a narrow baso-palatal interdenticular notch (Pratt, 1981a: fig. 3i).

. Complete loss of pustulation in the penial apex (1994a: fig. 2, character-state 5.6).

. Asmall, sac-like, glandular diverticulum of the lower penis (Fig. 5Ln), and derivatives thereof.
. Patches of glandular cells on the penial wall above the diverticulum (Fig. 5Ln).

. Penial diverticulum large, one-third to equal the volume of the penis (Fig. 5Lb).

. Shell apertural expansion rate (Secondarily) regular throughout ontogeny.

. A secondary lobe on the penial diverticulum (Fig. 5Lb).

. A (secondarily) globose shell with the aperture entirely free of denticles (Pilsbry, 1940: figs.

425-429).

. Adnate penial diverticulum (Fig. 5PrF).

. Penial diverticulum long and at least twice the volume of the penis (Fig. 5PrP, PrX).

. Shell depressed, broadly umbilicate (Zilch, 1959-1960: fig. 2036).

. Spiral, threadlike sculpture on the embryonic shell (Pilsbry, 1940: 689).

. A bifurcate or trifurcate penial retractor muscle (Fig. 5PrP).

. A vestigial epiphallus without a flagellum (Fig. 5MiM, МР).

. А slender penis (width < 0.12 length) with an apical, pendant, conical projection (Fig. 5MiM,

MiP), and derivatives thereof.

. An extremely long and slender penis (width < 0.06 length) (Fig. SMiM).

. A greatly enlarged, muscular, proximal vas deferens (Figs. 7МГР, 13).

. Even-diameter vas deferens with no trace of epiphallus (Fig. 500, DU).

. A stout penis (length/diameter <3.5) with a straight apex (Fig. 5DU).

. Amoderately long penis (4 < length/diameter < 10) with a bent or convoluted apex (Fig. 5DD).
. A downward curve on the lower limb of the parietal apertural denticle (Pilsbry, 1940: figs.

384-387).

. A raised parietal callus (Pilsbry, 1940: figs. 384-387).
. А penial apical chalice formed by the junction of lateral pilasters (1991a: fig. 27).
. An even-diameter distal vas deferens with no trace of flagellum or epiphallus (1994a: fig. 2:

character-state 9.6).

. An arched parietal apertural denticle (Fig. 13: character-state 9b), and derivatives thereof.
. Adepressed, hairless shell (height/diameter 0.4-0.6) (1991a: figs. 49, 50), and derivatives thereof.
. A regularly oval-shaped aperture with the reflected lip uniform in width throughout its palatal

and basal regions, with no basal dentition (1991a: fig. 48a).

. A basal apertural lamella (1991a: figs. 46, 47).

. A pronounced, blade-like parietal apertural denticle (1991a: figs. 46, 47).

. A straight basal region of the aperture with only a vestigial lamella (1991a: fig. 10b).

. A barrel-shaped, solid pedestal underlying the penial chalice (1991a: fig. 45c-d).

. Asmall, globose or subglobose shell (diameter 8-15 mm, height/diameter 0.6-0.7) (1991a: figs.

35a,b, 40a,b), and derivatives thereof.

. A globose, hairless shell (1991a: figs. 35a,b).
. A subglobose shell (height/diameter 0.5-0.6) bearing periostracal scales, with the umbilicus

broadly covered by an extension of the basal apertural lip, and with palatal and basal apertural
denticles (1991a: figs. 40a,b), and derivatives thereof.

. A thick-walled, hooded, cup-shaped penial chalice (1991a: figs. 7, 8, 9b).
. Penial chalice with a higher left than right wall (1991a: fig. 27, transformation 21).
. Shell very broadly umbilicate (1991a: figs. 39a,c).

Dorsal penial sculpture (1991a: fig. 28, transformations 31-33).
Dorsal penial sculpture consisting of 4-10 cord-like, subparallel, anastomosing ridges, running
longitudinally to 30-degrees obliquely (1991a: figs. 4, 6, 16a).

(continued)

78 EMBERTON

TABLE 1. Anatomical, behavioral, and shell characters used for a phylogenetic hypothesis (Fig. 8) and
a revision (text) of the Polygyridae. Citations refer to papers of Emberton, unless otherwise indicated.

(Continued)

112. Dorsal penial sculpture consisting of 8-12 cord-like, subparallel, anastomosing ridges, running
longitudinally to 30-degrees obliquely, many of which are contiguous with one or both lateral
pilasters, and many of which enlarge basally to form a network of large basal bulges (1991a:

figs. 2b, 11b,c, 156).

113. Penial chalice a deep, thin-walled scoop, with the left wall much higher than the right (1991a:

figs: 2b, bie, 156).

114. Dorsal penial sculpture consisting of about 8-12 thin parallel ridges, equal in diameter, which
is constant or gradually increases basally (1991a: figs. 15a,b, 16b,c).
115. Penial chalice a thick-walled, rounded or pointed ear-like flap, flared to the left (1991a: figs.

15a,b, 16b,c).

within the tribe Polygyrini (Figs. 4, 5), and (c)
shell ontogenetic and apertural morphology
within—primarily—the tribes Stenotremini
and Polygyrini (Fig. 6).

Figure 7 shows the distributions of all 115
characters among polygyrid subgenera (as
revised below).

Figure 8 gives the maximum-parsimony
cladogram of polygyrid subgenera (as ге-
vised below), based on the data presented in
Figure 7. Each node in the cladogram is sup-
ported by one to three characters, numbered
as in Table 1.

Taxonomic Revision

The following revision exactly follows the
phylogenetic hypothesis of Figure 8. For
brevity's sake, definitions employ character
numbers as defined in Table 1. Species with
only tentative assignment to subgenus are
preceded by a question mark.

Family POLYGYRIDAE Pilsbry, 1894
Definition: The first stylommatophoran pul-

monate gastropod to possess characters #1,
2, and 3, and all of its descendants.

Subfamily TRIODOPSINAE Pilsbry, 1940
Definition: The first Polygyridae to possess

characters #4 and 5, and all of its descen-
dants.

Tribe TRIODOPSINI

Definition: as for the subfamily.

Unnamed Clade Comprising Webbhelix,
Neohelix, and Xolotrema

Synonym: “Xolotrema Rafinesque” (Webb,
1952)

Definition: The first Triodopsinae to pos-
sess character #6, and all of its descendants.

Genus Webbhelix Emberton, 1988

Type species: Helix multilineata Say, 1821,
by original designation.

Definition: The first Triodopsinae to pos-
sess character #7, and all of its descendents.

Species: W. multilineata (Say, 1821).

Genus Neohelix Ihering, 1892

Type species: Helix albolabris Say, “1816”
1817, by subsequent designation (Pilsbry,
1930a).

Definition: The first Triodopsinae to pos-
sess character #8, and all of its descendents.

Subgenus Neohelix (Asamiorbis) subgen. n.

Type species: Helix dentifera Binney, 1837.

Definition: The first Neohelix to possess
character #9, and all of its descendents.

Etymology: Dr. Takahiro Asami, land-snail
ecologist and geneticist, who has worked
extensively with N. (A.) dentifera and other
polygyrids in Virginia (Asami, 1988, 1993); or-
bis (Latin) “disc” or “coil.”

Species: N. (A.) dentifera (Binney, 1837); N.
(A.) divesta (Gould, 1851); N. (A.) lioderma
(Pilsbry, 1902).

Unnamed clade comprising subgenera
N. (Neohelix) s.s. and
N. (Solemorbis) subgen. n.

Definition: The first Neohelix to possess
character #10, and all of its descendents.

POLYGYRID SHELL EVOLUTION 79

FIG. 3. Character-state analysis of penial-functional-surface anatomy of the Stenotremini; see Table 1,
characters #52, 58, 60-64. Characters are delineated by solid lines; arrows are hypothesized transforma-
tions among characters. Anatomical figures (at different size scales) are from Archer (1948). A, Stenotrema
(Archerelix) subgen. n.; С Stenotrema (Cohutta); E, Euchemotrema; P, Stenotrema (Pilsbrelix) subg. n.; $,
Stenotrema (Stenotrema); T, Stenotrema (Toxotrema); aa, $. altispira altispira; ad, $. altispira depilatum; bb,
$. barbatum; bd, $. blandianum; bg, $. barbigerum; bv, $. brevipila; cd, $. caddoense; ch, $. cohuttense;
dc, S. deceptum; eg, $. edgarianum; ev, $. edvardsi; ex, $. exodon; ext, $. exodon turbinella (= $. turbinella);
fl, 5. florida; ft, E. fraternum; h $. hirsutum; |, $. labrosum, ma, E. monodon aliciae; mg, $. magnifumosum;
mn, E. monodon, mx, $. maxillatum; pb, $. pilsbryi; pl, $. pilula; sp, $. spinosum; st, $. stenotrema; и, $.
unciferum.

80 ЕМВЕАТОМ

FIG. 4. Reproductive system (minus ovotestis) and shell (umbilical view) of Millerelix (Prattelix) plicata (ANSP
A2423-A) from Knox County, Tennessee. a = atrium, ag = albumen gland, ap = apical, pendant, conical
projection of the penis, fpsc = fertilization pouch-seminal receptacle complex (= talon = carrefour), gp =
gonopore, hd = hermaphroditic duct, mf = mantle-cavity floor, о = oviduvt, р = penis, pr = penial retractor
muscle, pt = prostate, sd = spermathecal duct (= bursa copulatrix duct = gametolytic duct), u = uterus, v

= vagina, vd = vas deferens.

Subgenus Neohelix (Neohelix) s.s.

Synonym: Neohelix albolabris group (Em-
berton, 1988).

Definition: The first Neohelix to possess
character #11, and all of its descendents.

Species: N. (N.) albolabris (Say, 1817); N.
(N.) major (Binney, 1837).

Subgenus Neohelix (Solemorbis) subgen. n.

Type species: Neohelix solemi Emberton,
1988.

Synonym: Neohelix alleni group (Ember-
ton, 1988).

Definition: The first Neohelix to possess
character #12, and all of its descendents.

Etymology: The late Dr. Alan Solem, long

POLYGYRID SHELL EVOLUTION 81

МР

FIG. 5. Character-state analysis of the lower-reproductive-tract anatomy of the Polygyrini; see Table 1,
characters #75-92. Characters are delineated by solid lines; arrows are hypothesized transformations
among characters. Anatomical figures (at different size scales) are from Pilsbry (1940), Metcalf 8 Riskind
(1979), Pratt (1981a), and this paper (Fig. 4). DD, Daedalochila s.s.; DU, Daedalochila (Upsilodon); G,
Giffordius; Lb, Lobosculum; Ln, Linisa; ММ, Millerelix s.s.; МР, Millerelix (Prattelix); Po, Polygyra; PrF,
Praticolella (Farragutia); PrP, Practicolella (Praticolella); PrX, Praticolella (Filapex); be, Pr. berlandieriana; bu,
D. burlesoni; ch, D. chisosensis; cr, Po. cereolus; df, Mi. doerfeuilliana; hp, D. hippocrepis; Ip, D. leporina;
Iw, Pr. lawae; mb, Pr. mobiliana; mp, D. multiplicata; mr, Mi. тоогеапа; pi, G. pinchoti; pl, Mi. plicata; pu,
Lb. pustula; sp, Po. septemvolva volvoxis; tm, Ln. tamaulipasensis; tx, Ln. texasiana; uv, D. uvulifera.

82 ЕМВЕАТОМ

FIG. 6. Character analysis of shell morphologies of the Stenotremini and the Polygyrini; Table 1, characters
#53, 54, 56, 57, 59, 70-72.

83

POLYGYRID SHELL EVOLUTION

TTOOO
OOTTO
0000T
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
STI
=ETT

TOTOO
TOTOO
TOTOO
OTTOO
000TO
0000T
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
OTT
-90T

00000
00000
00000
00000
00000
0т000
TTOOO
00TTO
0000T
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
SOT
-TOT

OOOTT
OOOTT
OOOTT
OOOTT
OOOTT
OOOTT
OOOTT
TOTTT
TOTIT
OTTIT
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00-96

T0000
T0000
T0000
70000
T0000
70000
T0000
T0000
T0000
T0000
OTTTO
0000T
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
S6-T6

T0000
T0000
70000
10000
10000
70000
T0000
T0000
10000
70000
TOOOT
TOOOT
OTOTT
OOTTT
00000
00000
00000
00000
00000
00000
00000
т0000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
06-98

00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
TOOTO
OTOTO
OOTTO
0000T
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
S8-T8

00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
TOTTO
TOTTO
TOTTO
TOTTO
OTTTO
0000T
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
08-94

00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
OTTTO
OTTTO
OTTTO
OTTTO
TITIO
TITTO
TITTO
TITTO
TITTO
TTTTO
0000T
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
SL-TL

OOOTT
OOOTT
OOOTT
OOOTT
OOOTT
OOOTT
OOOTT
OOOTT
O00TT
OOOTT
TOTTT
TOTTT
TOTTT
TOTTT
TOTTT
TOTTT
TOTTT
TOTTT
TOTTT
TOTTT
TOTTT
OTTTT
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
02-99

70000
70000
70000
70000
T0000
T0000
70000
T0000
70000
T0000
10000
T0000
T0000
T0000
70000
70000
T0000
70000
70000
70000
70000
70000
OTOOT
OOTOT
OOOTT
00000
00000
00000

‘00000

00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
S9-T9

‘sdno1Byno pue елэцэбап$ puABAjod Buowe (| э|ае1) заэзоелецо вр | JO SUONNQUISIC ‘7 "Ol

00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
OTOTT
OTOTT
OTOTT
TTOTT
OOTTT
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
09-95

0000T
0000T
0000T
0000T
0000T
0000T
0000T
0000T
0000T
0000T
0000T
0000T
0000T
0000T
0000T
0000T
0000T
0000T
0000T
0000T
0000T
0000T
OTTTT
OTTTT
OTTTT
OTTTT
OTTTT
ETT ET
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
SS-TS

TTOOO
TTOOO
TTOOO
TTOOO
TTOOO
TTOOO
11000
TTOOO
TTOOO
TTO00
TTO00
TTOOO
TTOOO
TTOOO
TTOOO
TTOOO
TTOOO
TTOOO
TTOOO
TTOOO
TTOOO
TTO00
TTOOO
TTOOO
TTOOO
TTOOO
TTOOO
TTOOO
OOTTO
00000
00000
00000
0000T
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
05-97

00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00007
00000
00000
000T0
TOTTO
OTTTO
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
ЗУ-ТУ

000T0
000T0
000T0
000T0
000T0
000T0
000TO
000T0
000T0
000T0
000T0
000T0
000T0
000T0
000T0
000T0
000T0
000T0
000T0
000T0
000T0
000T0
000T0
000T0
000T0
000T0
000T0
000T0
000T0
OTTTO
OTTTO
TTTTO
OOTTO
OOTTO
OOTTO
0000T
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
07-95

OOTTT
OOTTT
OOTTT
OOTTT
OOTTT
OOTTT
OOTTT
OOTTT
OOTTT
OOTTT
OOTTT
OOTTT
OOTTT
OOTTT
OOTTT
OOTTT
OOTTT
OOTTT
OOTTT
OOTTT
OOTTT
OOTTT
OOTTT
OOTTT
OOTTT
OOTTT
OOTTT
OOTTT
OOTTT
OOTTT
OOTTT
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ТЕТЕ
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181919151
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(uoposayw) ‘en
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xTTauoung
(snr18309T3UI) ‘I
(sntqyotaqny) ‘т
(эталоэтерзБеч) ‘ed
(елэзеа) ‘ed
(eza3ediadsaA) ‘ва
(eTreyooTepseg) ‘а
(иороттзап) ‘а
(хттеззела) ‘ти
(хттедеэттти) ‘ти
(еттетортзела) ‘ла
(хэ4еття) ‘ла
(эзпрлепря) "Id
(етэпбеллея) ‘ла
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едАБАТОа
5птрлоуутэ
(xTTS1qsTId) *5
(ешэ130и935) *5
(ews130x0L) *s
(хттелэечолу) *5
(23314092) "Ss
PUSIJOWSUINT
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(зталортоден) “PL
(зталоиетттшоеи) “PL
(sTGIOA1ASTIA) “PL
(sTqitosyoo1g) “PL
(stq2opXoJTaus) “PL
(ешэдзотох) *х
(зталохоэттм) *х
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84 ЕМВЕАТОМ

В Bradybaena
X Cepolis
H Helminthoglypta outgroups
Т Thysanophora
с Pleurodonte
Ss Sagda
—_—_ + Webbhelix
| 10-11—NN Neohelix s.s.

6 8—— 12—NS Neohelix (Solemorbis)
| 9—_NA Neohelix (Asamiorbis)
4,5 13,14——15—XW Xolotrema (Wilcoxorbis) Trio-
16—XX Xolotrema s.s. dop-
19——_———————_TdS Triodopsis (Shelfordorbis) sini
17,18420—————————TGB Triodopsis (Brooksorbis)
21-22,23————TdP Triodopsis (Pilsbryorbis)
24-25——————TÓM Triodopsis (Macmillanorbis)
1-3 26-27 TdH Triodopsis (Haroldorbis)
28-29 —TdT Triodopsis s.s.
30—TdV Triodopsis (Vagvolgyorbis)
34————35—H Hochbergellus Vesperi-
36—V___Vespericola colini
40—CB Cryptomastix (Bupiogona)
3 8 ———— 39 —— HCC Cryptomastix s.s. Allo-
31-33 41—CM Cryptomastix (Micranepsia) go-
r46 Tb Trilobopsis nini
42—43—44—А1А Allogona s.s.
45—A1D Allogona (Dysmedoma)
47 48—— АЗ Ashmunella Ashmunellini
55——E Euchemotrema
52-544 58————5С Stenotrema (Cohutta) Steno-
37 56/97 60 SA Stenotrema (Archerelix) tremini
59] 64—SP Stenotrema (Pilsbrelix)
61—463-SS Stenotrema s.s.
62—ST Stenotrema (Toxotrema)
49-51 69 ——-—_ TE Giffordius
68 VPO Polygyra
76—————Ln Linisa
70 75— 79———Lb Lobosculum
77,78 81 PrF Praticolella (Farragutia)
80—| 83-PrE Praticolella (Eduardus) Poly-
72-74 82-84-PrX Praticolella (Filapex) gyrini
85-PrP Praticolella s.s.
65-67 87-88-MiM Millerelix s.s.
85 ————] 89-MiP Millerelix (Prattelix)
90-91-DU Daedalochila (Upsilodon)
92-9400 Daedalochila s.s.
98—799——PaV Patera (Vesperpatera)
100-101PaP Раёега s.s.
102,103PaR Patera (Ragsdaleorbis)
95-97———————-1047105—IH Inflectarius (Hubrichtius)
| 106— ТТ Inflectarius s.s. Meso-
107————_F Fumonelix dontini

109——App Appalachina
1081 111MeA Mesodon (Aphalogona)
110+23MeK Mesodon (Akromesodon)

1145MeM Mesodon s.s.

FIG. 8. Phylogenetic hypothesis for polygyrid subgenera based on the data in Fig. 7. Synapomorphies
supporting each node and defining each subgenus are numbered as in Table 1. “23MeK” = 112, 113 MeK.
“1145MeM” = 114,115 MeM.

POLYGYRID SHELL EVOLUTION 85

one of the world's leading and most prolific
specialists on land snails, who did some im-
portant work on polygyrids (Solem, 1976); or-
bis (Latin) “disc” or “coil.”

Species: N. (S.) alleni (Sampson, 1883); N.
(S.) solemi Emberton, 1988.

Genus Xolotrema Rafinesque, 1819

Type species: Helix denotata Ferussac,
1821 (= Helix notata Deshayes, 1830) by sub-
sequent designation (Pilsbry, 1940).

Definition: The first Triodopsini to possess
characters #13 and 14, and all of its descen-
dents.

Subgenus Xolotrema (Wilcoxorbis)
Webb, 1952

Type species: Polygyra appressa fosteri F.
C. Baker, 1932, by original designation.

Definition: The first Xolotrema to possess
character #15, and all of its descendents.

Species: X. (W.) fosteri (F. C. Baker, 1932);
X. (W.) occidentalis (Pilsbry & Ferriss, 1907).

Subgenus Xolotrema (Xolotrema) s.s.

Definition: The first Xolotrema to possess
character #16, and all of its descendents.

Species: X. (X.) caroliniensis (Lea, 1834); X.
(X.) denotata (Ferussac, 1821); X. (X.) ob-
stricta (Say, 1821).

Genus Triodopsis Rafinesque, 1819

Type species: Helix tridentata Say, “1816”
1817, by original designation.

Definition: The first Triodopsinae to pos-
sess characters #17 and 18, and all of its
descendants.

Subgenus Triodopsis (Shelfordorbis)
Webb, 1959

Type species: Triodopsis fraudulenta vul-
gata Pilsbry, 1940, by original designation.

Synonym: species group Triodopsis vul-
gata (Emberton, 1988).

Definition: The first Triodopsis to possess
character #19, and all of its descendents.

Species: Т. (S.) claibornensis Lutz, 1950; Т.
(S.) fraudulenta (Pilsbry, 1894); Т. (S.) picea
Hubricht, 1958; 7. (S.) vulgata Pilsbry, 1940.

Subgenus Triodopsis
(Brooksorbis) subgen. n.

Type species: Polygyra platysayoides
Brooks, 1933.
Synonym: species group Triodopsis

platysayoides (Emberton, 1988).

Definition: The first Triodopsis to possess
character #20, and all of its descendents.

Etymology: The late Dr. Stanley T. Brooks,
who described the type species; orbis (Latin)
“ISO OF GO

Species: Т. (B.) platysayoides (Brooks,
1933).

Unnamed Clade Comprising Triodopsis
Subgenera 7. (Pilsbryorbis) subgen. n., Т.
(Macmillanorbis) subgen. n., Т.
(Haroldorbis), T. (Triodopsis), and Т.
(Vagvolgyorbis) subgen. п.

Definition: The first Triodopsis to possess
character 421, and all of its descendents.

Subgenus Triodopsis (Pilsbryorbis)
subgen. n.

Type species: Polygyra tridentata tennes-
seensis Walker 8 Pilsbry, 1902.

Synonym: species groups Triodopsis ten-
nesseensis and Т. burchi (Emberton, 1988a).

Definition: The first Triodopsis to possess
characters #22 and 23, and all of its descen-
dents.

Etymology: The late Dr. Henry A. Pilsbry,
for some 70 years the world's best known
and most productive land-snail specialist,
who wrote the definitive monograph on poly-
gyrids (Pilsbry, 1940); orbis (Latin) “disc” or
“coll.”

Species: Т. (P.) burchi Hubricht, 1950; 7.
(P.) complanata (Pilsbry, 1898); Т. (P.) tennes-
seensis (Walker & Pilsbry, 1902).

Unnamed Clade Comprising Triodopsis
Subgenera T. (Macmillanorbis) subgen. n.,
T. (Haroldorbis), T. (Triodopsis), and T.
(Vagvolgyorbis) subgen. n.

Definition: The first Triodopsis to possess
character #24, and all of its descendents.

Subgenus Triodopsis (Macmillanorbis)
subgen. n.

Type species: Triodopsis tridentata rugosa
Brooks & MacMillan, 1940.

Synonym: species group Triodopsis rug-
osa (Emberton, 1988).

86 ЕМВЕАТОМ

Definition: The first Triodopsis already hav-
ing character #24, to possess character #25,
and all of its descendents.

Etymology: The late Gordan K. MacMillan
of the Carnegie Museum, Pittsburg, who col-
lected and coauthored the type species; or-
bis (Latin) “disc” or “coil.”

Species: Т. (M.) fulciden Hubricht, 1952; Т.
(M.) rugosa Brooks & MacMillan, 1940.

Unnamed Clade Comprising Triodopsis
Subgenera 7. (Haroldorbis), T. (Triodopsis),
and 7. (Vagvolgyorbis) subgen. n.

Definition: The first Triodopsis already hav-
ing Character #24, to possess character #26,
and all of its descendents.

Subgenus Triodopsis (Haroldorbis)
Webb, 1959

Type species: Triodopsis cragini Call,
1886, by original designation.

Synonyms: “Triodopsis соре! (Wetherby)”
(Vagvolgyi, 1968; species group Triodopsis
cragini Call (Emberton, 1988a).

Definition: The first Triodopsis already hav-
ing characters #24 and 26, to possess char-
acter #27, and all of its descendents.

Species: Т. (H.) cragini Call, 1886; Т. (H.)
henriettae (Mazyck, 1877); T. (H.) vultuosa
(Gould, 1848).

Unnamed Clade Comprising Triodopsis
Subgenera 7. (Triodopsis) and Т.
(Vagvolgyorbis) subgen. n.

Definition: The first Triodopsis already hav-
ing Characters #24 and 26, to possess char-
acter #28, and all of its descendents.

Subgenus Triodopsis (Triodopsis) s.s.

Synonym: species groups Triodopsis tri-
dentata (Say) and Т. fallax (Say) (Emberton,
1988a).

Definition: The first Triodopsis already hav-
ing characters #24, 26, and 28, to possess
character #29, and all of its descendents.

Species group 7. (T.) tridentata s.s.: Т. (T.)
anteridon (Pilsbry, 1940); Т. (T.) tridentata
(Say, 1817).

Species group Т. (Т.) fallax: Т. (T.) alabam-
ensis (Pilsbry, 1902); Т. (T.) fallax (Say, 1825);
Т. (T.) hopetonensis (Shuttleworth, 1852); Т.
(T.) obsoleta (Pilsbry, 1894); T. (T.) palustris

Hubricht, 1958; Т. (T.) soelneri (Henderson,
1907); Т. (T.) vannostrandi (Bland, 1875).

Subgenus Triodopsis (Vagvolgyorbis)
subgen. n.

Type species: Polygyra tridentata juxtidens
Pilsbry, 1894b.

Synonym: species group Triodopsis juxti-
dens (Pilsbry) (Emberton, 1988).

Definition: The first Triodopsis already hav-
ing Characters #24, 26, and 28, to possess
character #30, and all of its descendents.

Etymology: Dr. Joseph Vagvolgyi, author
of a conchological monograph on the Triod-
opsinae and Cryptomastix (Vagvolgyi, 1968);
orbis (Latin) “disc” or “coil.”

Species: Т. (V.) discoidea (Pilsbry, 1904); Т.
(V.) juxtidens (Pilsbry, 1894b); Т. (V.) neglecta
(Pilsbry, 1899); T. (V.) pendula Hubricht,
1952.

Subfamily POLYGYRINAE s.s.

Definition: The first Polygyridae to possess
characters 431, 32, and 33, and all of its de-
scendents.

Tribe VESPERICOLINI Emberton, 1994

Type genus: Vespericola Pilsbry, 1939, by
original designation.

Definition: The first Polygyrinae to possess
character #34, and all of its descendents.

Genus Hochbergellus Roth & Miller, 1992

Type species: Hochbergellus hirsutus Roth
8 Miller, 1992, by original designation.

Definition: The first Vespericolini to pos-
sess character #35, and all of its descen-
dents.

Species: H. hirsutus Roth & Miller, 1992.

Genus Vespericola Pilsbry, 1939

Type species: Polygyra columbiana pilosa
Henderson, 1928, by original designation.

Definition: The first Vespericolini to pos-
sess character #36, and all of its descen-
dents.

Species: V. armigera (Binney, 1885); V. co-
lumbianus (Lea, 1838); У. euthales (Berry,
1939); V. hapla (Berry, 1933); V. karokorum
Talmadge, 1962; У. marinensis Roth & Miller,
1993; У. megasoma (Dall, 1905); “М. sp. п. 1”
(Roth 8 Miller, 1993); “М. sp. п. 2” (Roth 4
Miller, 1993); V. orius (Berry, 1933); V. pilosus

POLYGYRID SHELL EVOLUTION 87

(Henderson, 1928); V. pinicola (Berry, 1916);
V. pressleyi Roth, 1985; V. shasta (Berry,
1921); V. sierrana (Berry, 1921).

Unnamed Clade Comprising Tribes
Allogonini and Ashmunellini and
Infrafamily Polygyrinai

Definition: The first Polygyrinae to possess
character 437, and all of its descendents.

Tribe ALLOGONINI Emberton, 1994

Type genus: Allogona Pilsbry, 1939, by
original designation.

Definition: The first Polygyrinae already
having character #37, to possess character
#38, and all of its descendents.

Genus Cryptomastix Pilsbry, 1939

Type species: Polygyra mullani olneyae
Pilsbry, 1928, by original designation.

Synonym: Triodopsis Rafinesque (in part)
(Pilsbry, 1940; Vagvolgyi, 1968).

Definition: The first Allogonini to possess
character #39, and all of its descendents.

Subgenus C. (Bupiogona) Webb, 1970

Type species: Polygyra mullani hendersoni
Pilsbry, 1928, by original designation.

Definition: The first Cryptomastix to pos-
sess character #40, and all of its descen-
dents.

Species: С. (В.) hendersoni (Pilsbry, 1928).

Subgenus С. (Cryptomastix) s.s.

Definition: (as for the genus).

Species: C. (C.) devia (Gould, 1846); C. (C.)
mullani (Напа & Cooper, 1862); С. (C.) san-
burni (Binney, 1886).

Subgenus C. (Micranepsia) Pilsbry, 1940

Type species: Helix germana Gould, 1851,
by original designation.

Definition: The first Cryptomastix to pos-
sess character #41, and all of its descen-
dents.

Species: C. (M.) germana (Gould, 1851).

Unnamed Clade Comprising Allogona
and Trilobopsis

Definition: The first Allogonini to possess
character #42, and all of its descendents.

Genus Allogona Pilsbry, 1939

Type species: Helix profunda Say, 1821, by
original designation.

Definition: The first Allogonini already hav-
ing Character #42, to possess character #43,
and all of its descendents.

Subgenus A. (Allogona) s.s.

Definition: The first Allogonini to possess
character #44, and all of its descendents.
Species: A. (A.) profunda (Say, 1821)

Subgenus A. (Dysmedoma) Pilsbry, 1939

Type species: Helix townsendiana Lea,
1838, by original designation.

Definition: The first Allogonini to possess
character #45, and all of its descendents.

Species: А. (D.) lombardii Smith, 1943; A.
(D.) ptychophora (Brown, 1870); A. (D)
townsendiana (Lea, 1838).

Genus Trilobopsis Pilsbry, 1939

Type species: Helix loricata Gould, 1846,
by original designation.

Definition: The first Allogonini already hav-
ing Character #42, to possess character #46,
and all of its descendents.

Species: Т. loricata (Gould, 1846); Т. peni-
tens (Hanna 4 Rixford, 1923); T. roperi (Pils-
bry, 1889); Т. tehamana (Pilsbry, 1928); Т. tra-
chypepla (Berry, 1933).

Tribe ASHMUNELLINI Webb, 1954

Type genus: Ashmunella Pilsbry 8 Cocker-
ell, 1899, by original designation.

Definition: The first Polygyrinae already
having character 437, to possess characters
#47 and 48, and all of its descendents.

Genus Ashmunella Pilsbry & Cockerell,
1899

Type species: Polygyra miorhyssa Dall,
1898 [= Ashmunella rhyssa miorhyssa (Dall,
1898)], by subsequent designation (Pilsbry,
1905).

Definition: (as for the tribe).

Species (n = 49): A. altissima (Cockerell,
1898); A. angulata Pilsbry, 1905; A. ani-
masensis Vagvolgyi, 1974; A. ashmuni (Бай,
1896); A. auriculata (Say, 1818); A. bequaerti
Clench & Miller, 1966; A. binneyi Pilsbry &
Ferriss, 1917; A. carlbadensis Pilsbry, 1932;

88 ЕМВЕАТОМ

А. chiricahuana (Dall, 1896); А. cockerelli Pils-
bry 4 Ferriss, 1917; A. danielsi Pilsbry 8 Fer-
riss, 1915; A. edithae Pilsbry 8 Cheatum,
1951; A. esuritor Pilsbry, 1905; A. ferrissi Pils-
bry, 1905; A. harrisi Metcalf 8 Smartt, 1977;
А. hawleyi Metcalf, 1973; A. hebardi Pilsbry &
Vanatta, 1923; A. intricata Pilsbry, 1948; A.
jamesensis Metcalf, 1973 (fossil); A. juarezen-
sis Pilsbry, 1948; A. kochi Clapp, 1908; A.
lenticula Gregg, 1953; A. lepidoderma Pilsbry
8 Ferriss, 1910; A. /evettei (Bland, 1881); A.
macromphala Vagvolgyi, 1974; A. mearnsi
(Dall, 1896); A. mendax Pilsbry 8 Ferriss,
1917; A. meridionalis Pilsbry, 1948; A. mog-
ollonensis Pilsbry, 1900; A. montivaga Pils-
bry, 1948; A. mudgei Cheatum, 1971; A. or-
ganensis Pilsbry, 1936; A. pasonis (Drake,
1951); A. pilsbryana Ferriss, 1914; A. proxima
Pilsbry, 1905; A. pseudodonta (Dall, 1897); A.
rhyssa (Dall, 1897); A. rileyensis Metcalf &
Hurley, 1971; A. ruidosana Metcalf, 1973
(fossil); A. salinasensis Vagvolgyi, 1974; A.
sprouli Fullington 4 Fullington, 1978; A. tegil-
lum Metcalf, 1973; A. tetrodon Pilsbry & Fer-
riss, 1915; A. thomsoniana (Ancey, 1887); A.
todseni Metcalf 8 Smartt, 1977; A. tularosana
Metcalf, 1973; A. varicifera Ancey, 1901; A.
walkeri Ferriss, 1904; A. watleyi Metcalf €
Fullington, 1978.

Infrafamily POLYGYRINAI s.s.

Definition: The first Polygyrinae already
having character #37, to possess characters
#49, 50, and 51, and all of its descendents.

Tribe STENOTREMINI Emberton, 1994

Type genus:
1819.

Definition: The first Polygyrinai to possess
characters #52, 53, and 54, and all of its de-
scendents.

Stenotrema Rafinesque,

Genus Euchemotrema Archer, 1939

Type species: Helix monodon Rackett,
1821, by subsequent designation (Pilsbry,
1940).

Definition: The first Stenotremini to pos-
sess character #55, and all of its descen-
dents.

Species: E. fasciatum (Pilsbry, 1940); E.
fraternum (Say, 1824); E. hubrichti (Baker,
1937); E. leai (Binney, 1840); E. monodon
(Rackett, 1821); E. occidaneum Roth 8 Em-

berton, 1994 (early Miocene fossil); E. wichi-
torum Branson, 1972.

Genus Stenotrema Rafinesque, 1819

Type species: Stenotrema convexa
Rafinesque, 1819 [nomen nudum = Helix
stenotrema Pfeiffer, 1842].

Definition: The first Stenotremini to pos-
sess characters #56 and 57, and all of its
descendents.

Subgenus Stenotrema (Cohutta)
Archer, 1948

Type species: Polygyra cohuttensis Clapp,
1914, by original designation.

Definition: The first Stenotrema to possess
character #58, and all of its descendents.

Species: S. (Cohutta) cohuttensis Archer,
1948

Unnamed Clade Comprising Stenotrema
(Archerelix), S. (Pilsbrelix), S. (Stenotrema)
s.s., and S. (Toxotrema)

Definition: The first Stenotrema to possess
character #59, and all of its descendents.

Subgenus Stenotrema (Archerelix)
subgen. n.

Type species: Helix barbigera Redfield,
1856.

Description: The first Stenotrema to pos-
sess character #60, and all of its descen-
dents.

Etymology: The late Dr. Allan F. Archer,
who contributed “information, notes, and
manuscript”? (Archer, 1948: 8) to Pilsbry’s
(1940) monograph on Stenotrema and wrote
a revision and ecological manual on the ge-
nus (Archer, 1948); helix (Latin) “coil” or
“snail.”

Species: $. (A) barbigerum (Redfield,
1856); S. (A.) edgarianum (Lea, 1841); S. (A.)
edvardsi (Bland, 1858); S. (A.) pilsbryi (Fer-
riss, 1900).

Unnamed Clade Comprising S. (Pilsbrelix),
S. (Stenotrema) s.s., and S. (Toxotrema)

Definition: The first Stenotrema to possess
character #61, and all of its descendents.

POLYGYRID SHELL EVOLUTION 89

Subgenus $. (Toxotrema) Rafinesque, 1819

Type species: Helix hirsuta Say, 1817, by
subsequent designation (Pilsbry, 1930).

Definition: The first Stenotrema to possess
character #62, and all of its descendents.

Species: 5. (T.) barbatum (Clapp, 1904); $.
(T.) hirsutum (Say, 1817); S. (T.) labrosum
(Bland, 1862); ?S. (T.) simile Grimm, 1971.

Subgenus Stenotrema (Stenotrema) s.s.

Synonyms: Stenotrema (Stenostoma)
Rafinesque, 1831 (Archer, 1948); Stenotrema
(Maxilliter) Pilsbry, 1940 (Archer, 1948);
Stenotrema (Coracollatus) Archer, 1948.

Definition: The first Stenotrema to possess
character #63, and all of its descendents.

Species and subspecies: $. (S.) altispira
(Pilsbry, 1894); $. (S.) altispira depilatum
(Pilsbry, 1895); ?S. (S.) angellum Hubricht,
1958; $. (S.) brevipila (Clapp, 1907); $. (S.)
caddoense (Archer, 1935); 75$. (S.)
calvescens Hubricht, 1961; S. (S.) florida Pils-
bry, 1940; $. (S.) magnifumosum (Pilsbry,
1900); S. (S.) maxillatum (Gould, 1848); ?S.
(S.) morosum Hubricht, 1978 (Pleistocene-
Recent fossil); S. (S.) pilula (Pilsbry, 1900); S.
(S.) spinosum (Lea, 1831); S. (S.) stenotrema
(Pfeiffer, 1842); S. (S.) unciferum (Pilsbry,
1900); ?S. (S.) waldense Archer, 1938.

Subgenus Stenotrema (Pilsbrelix)
subgen. n.

Type species: Polygyra stenotrema exodon
Pilsbry, 1900.

Description: The first Stenotrema to pos-
sess character #64, and all of its descen-
dents.

Etymology: The late Dr. Henry A. Pilsbry,
who wrote the definitive monograph on poly-
gyrids (Pilsbry, 1940), and who described the
type species; helix (Latin) “coil” or “snail.”

Species: S. (P) blandianum (Pilsbry, 1903);
$. (P) deceptum (Clapp, 1905); $. (P.) exodon
(Pilsbry, 1900); S. (P) turbinella (Clench & Ar-
cher, 1933).

Unnamed Clade Comprising Polygyrini
and Mesodontini

Definition: The first Polygyrinai to possess
characters #65, 66, and 67, and all of its de-
scendents.

Tribe POLYGYRINI s.s.

Definition: The first Polygyrinai already
having characters #65, 66, and 67, to pos-
sess character #68, and all of its descen-
dents.

Genus Giffordius Pilsbry, 1930

Type species: Giffordius pinchoti Pilsbry,
1930, by original designation.

Definition: The first Polygyrini to possess
character #69, and all of its descendents.

Species: G. corneliae Pilsbry, 1930; G. pin-
choti Pilsbry, 1930.

Unnamed Clade Comprising Polygyra,
Linisa, Lobosculum, Praticolella, Millerelix,
and Daedalochila

Definition: The first Polygyrini to possess
character #70, and all of its descendents.

Genus Polygyra Say, 1818

Type species: Polygyra septemvolva Say,
1818, by subsequent designation (Herr-
mannsen, 1847).

Definition: The first Polygyrini already hav-
ing character #70, to possess character #71,
and all of its descendents.

Species: P. caloosaensis Johnson, 1899
(Pliocene fossil); Р. cereolus (Mühlfeld, 1818);
P. paludosa (Wiegmann, 1839); P. plana
(Dunker, 1843); P. septemvolva Say, 1818.

Unnamed Clade Comprising Linisa,
Lobosculum, Praticolella, Millerelix,
and Daedalochila

Definition: The first Polygyrini already hav-
ing character #70, to possess characters
#72, 73, and 74, and all of its descendents.

Unnamed Clade Comprising Linisa,
Lobosculum, and Praticolella

Definition: The first Polygyrini already hav-
ing characters #70, 72, 73, and 74, to pos-
sess character #75, and all of its descen-
dents.

Genus Linisa Pilsbry, 1930

Type species: Helix (Polygyra) anilis Gabb,
1865, by original designation.

Synonyms (fide Pratt, 1981a): Polygyra
(Daedalochila) texasiana group (in part) (Pils-

90 ЕМВЕАТОМ

bry, 1940); Polygyra (Erymodon) Pilsbry,
1956; Polygyra (Monophysis) Pilsbry, 1956;
Polygyra (Solidens) Pilsbry, 1956; Polygyra
(Linisia) (Pratt, 1981a,b); Daedalochila (in
part) (Richardson, 1986).

Definition: The first Polygyrini already hav-
ing Characters #70, 72, 73, 74, and 75, to
possess character #76, and all of its descen-
dents.

Species: ?L. adamnis (Dall, 1890) (Upper
Oligocene fossil); L. albicostulata (Pilsbry,
1896); L. anilis (Gabb, 1865); ?L. aula-
comphala (Pilsbry & Hinkley, 1907); L. behri
(Gabb, 1865); ?L. bicruris (Pfeiffer, 1857); ?L.
cantralli (Solem, 1957); ?L. couloni (Shuttle-
worth, 1852); ?L. dissecta (Martens, 1892);
?L. dysoni (Shuttleworth, 1852); ?L. euglypta
(Pilsbry, 1896); ?L. hertleini Haas, 1961; ?L.
hindsii (Pfeiffer, 1845); ?L. idiogenes (Pilsbry,
1956); ?L. matermontana (Pilsbry, 1896); ?L.
nelsoni (Dall, 1897); L. pergrandis (Solem,
1959); ?L. plagioglossa (Pfeiffer, 1859); L.
polita (Pilsbry 4 Hinkley, 1907); ?L. ponsonbyi
(Pilsbry, 1896); [. richardsoni (Martens,
1892); ?L. suprazonata (Pilsbry, 1900); L.
tamaulipasensis (Lea, 1867); L. texasiana
(Moricand, 1833); L. ventrosula (Pfeiffer,
1845); ?L. yucatanea (Morelet, 1853).

Unnamed Clade Comprising Lobosculum
and Praticolella

Definition: The first Polygyrini already hav-
ing characters #70, 72, 73, 74, 75, and 76, to
possess characters #77 and 78, and all of its
descendents.

Genus Lobosculum Pilsbry, 1930

Type species: Helix pustula Férussac,
1822, by subsequent designation (Pilsbry,
1930b: 320).

Definition: The first Polygyrini already hav-
ing characters #72, 73, 74, 75, 76, 77, and
78, to possess character #79, and all of its
descendents.

Species: L. pustula (Férussac, 1822); L.
pustuloides (Bland, 1858).

Genus Praticolella Martens, 1892

Type species: Praticola ocampi Strebel 8
Pfeffer, 1880 (= Helix ampla Pfeiffer, 1866),
by original designation.

Definition: The first Polygyrini already hav-
ing characters #72, 73, 74, 75, 76, 77, 78, to

possess character #80, and all of its descen-
dents.

Subgenus Praticolella (Farragutia)
Vanatta, 1915

Type species: Helix mobiliana Lea, 1841,
by original designation.

Definition: The first Praticolella to possess
character #81, and all of its descendents.

Species: P. (F.) mobiliana (Lea, 1841).

Unnamed Clade Comprising Praticolella
(Eduardus), P. (Filapex), and
Р. (Praticolella) s.s.

Definition: The first Praticolella to possess
character #82, and all of its descendents.

Comment: The membership of P. (Eduar-
dus) in this clade needs to be tested by dis-
section, because Pilsbry (1936) was inexplicit
about the size of the appendix.

Subgenus Praticolella (Eduardus)
Pilsbry, 1930

Type species: Polygyra martensiana Pils-
bry, 1907, by original designation.

Definition: The first Praticolella already
having character #82, to possess character
#83, and all of its descendents.

Species: P. (E.) martensiana
1907).

(Pilsbry,

Subgenus Praticolella (Filapex) Pilsbry, 1940

Type species: Helix jejuna Say, 1821, by
original designation.

Definition: The first Praticolella already
having character #82, to possess character
#84, and all of its descendents.

Species: P. (F.) bakeri (Vanatta, 1915); P.
(F.) jejuna (Say, 1821); P. (F.) lawae (Lewis,
1874).

Subgenus Praticolella (Praticolella) s.s.

Definition: The first Praticolella already
having character #82, to possess character
#85, and all of its descendents.

Species: P. (P.) ampla (Pfeiffer, 1866); P.
(P.) berlandieriana (Moricand, 1833); P. (P.)
candida Hubricht, 1983; P. (P.) flavescens
(Pfeiffer, 1848); P. (P.) griseola (Pfeiffer,
1841); P. (P.) pachyloma (Pfeiffer, 1847); P.
(P.) strebeliana Pilsbry, 1899; P. (P.) taeniata
Pilsbry, 1940; P. (P.) trimatris Hubricht, 1983.

POLYGYRID SHELL EVOLUTION 91

Unnamed Clade Comprising Millerelix
and Daedalochila

Definition: The first Polygyrini already hav-
ing characters #70, 72, 73, and 74, to pos-
sess character #86, and all of its descen-
dents.

Genus Millerelix Pratt, 1981 (see below)

Type species: Helix mooreana W. G. Bin-
ney, 1857, by original designation.

Definition: The first Polygyrini already hav-
ing characters #70, 72, 73, 74, and 86, to
possess character #87, and all of its descen-
dents.

Subgenus Millerelix (Millerelix) s.s.
Pratt, 1981

Definition: The first Millerelix to possess
chiaracter #88, and all of its descendents
(Pratt, 19815).

Species: M. doerfeulliana (Lea, 1838); M.
gracilis (Hubricht, 1961); ?М. implicata (Mar-
tens, 1865); M. lithica (Hubricht, 1961); M.
mooreana (Binney, 1857); 2M. rhoadsi (Pils-
bry, 1900); M. tholus (Binney, 1857).

Subgenus Millerelix (Prattelix) subgen. n.

Type species: Polygyra plicata Say, 1821.

Synonyms: Polygyra plicata group (Pilsbry,
1940); Daedalochila plicata group = unnamed
subgenus (Pratt, 1981a).

Definition: The first Millerelix to possess
character #89, and all of its descendents.

Comments: The shell of the type species is
sparsely and evenly covered with long, con-
spicuous periostracal hairs, seemingly round
in cross-section and slightly curved at the tip,
that fade out toward the umbilicus (Fig. 13).
They are easily broken, hence Pilsbry’s
(1940: 626) mildly erroneous, “a few short
hairs, usually preserved only in the umbilicus
and behind the lip.” The long hairs were
found in all four of the Academy of Natural
Sciences's alcohol-preserved lots of D. pli-
cata: Alabama, Madison County (ANSP A
2423-B); Tennessee, Marion County (A 2423-
C); Tennessee, Knox County (A 2423-A, Fig.
13); and Kentucky, Barren County (A 2387-B,
most hairs broken).

Etymology: The subgenus is named in
honor of Will Pratt.

Species: deltoidea (Simpson, 1889); fatigi-
ata (Say, 1829); jacksoni (Bland, 1866); pere-
grina (Rehder, 1932); plicata (Say, 1821); simp-

soni (Pilsbry & Ferriss, 1907); troostiana (Lea,
1838).

Genus Daedalochila Beck, 1837

Type species: Helix auriculata Say, 1818,
by subsequent designation (Herrmannsen,
1847).

Definition: The first Polygyrini already hav-
ing Characters #70, 72, 73, 74, and 86, to
possess character 490, and all of its descen-
dents.

Subgenus Daedalochila (Upsilodon)
Pilsbry, 1930c

Type species: Helix hippocrepis Pfeiffer,
1848, by original designation.

Synonym: Daedalochila (Acutidens) Pils-
bry, 1956.

Definition: The first Daedalochila to pos-
sess character #91, and all of its descen-
dents.

Comment: The recurved palatal apertural
denticle of D. acutidentata also occurs in the
otherwise very different shell of D. poeyi, so
does not seem to be a reliable character for
defining a clade.

Species: ?D. (U.) acutedentata (Binney,
1858); D. (U.) burlesoni (Metcalf & Riskind,
1979); D. (U.) chisosensis (Pilsbry, 1936); D.
(U.) да! (Metcalf & Riskind, 1979); D. (U.) hip-
pocrepis (Pfeiffer, 1848); D. (U.) leporina
(Gould, 1848); D. (U.) multiplicata (Metcalf &
Riskind, 1979); ?D. (U.) poeyi (Aguayo 8
Jaume, 1947); О. (U.) sp. п. A (Pratt, 1981 a);
О. (U.) sterni (Metcalf & Riskind, 1979).

Subgenus Daedalochila (Daedalochila) s.s.

Synonym: Polygyra auriculata group (Pils-
bry, 1940).

Definition: The first Daedalochila to pos-
sess characters #92, 93, and 94, and all of its
descendents.

Comments: The downward curve on the
lower limb of the parietal apertural denticle 1$
a newly discovered synapomorphy. Pratt
(1981a) stated (without giving evidence) that
2D. ariadne and ?Mi. implicata are members
of a new genus; this needs to be investi-
gated.

Species: 2D. (D.) ariadne (Pfeiffer, 1848); D.
(D.) auriculata (Say, 1818); D. (D.) auriformis
(Bland, 1862); D. (D.) avara (Say, 1818); D. (D.)
delecta (Hubricht, 1976); D. (D.) hausmani
(Jackson, 1948); 2D. (D.) oppilata (Morelet,

92 ЕМВЕАТОМ

1849); D. (D.) peninsulae (Pilsbry, 1940); D.
(D.) postelliana (Bland, 1862); D. (D.) sub-
clausa (Pilsbry, 1899); D. (D.) uvulifera (Shut-
tleworth, 1852).

Tribe MESODONTINI Emberton, 1991

Type genus: Mesodon Férussac, 1821.

Definition: The first Polygyrinai already
having characters #65, 66, and 67, to pos-
sess characters #95, 96, and 97, and all of its
descendents.

Genus Patera Albers, 1850

Type species: Helix appressa Say, 1821,
by subsequent designation Pilsbry, 1930c:
326).

Definition: The first Mesodontini to pos-
sess character #98, and all of its descen-
dents.

Subgenus Patera (Vesperpatera)
Emberton, 1991

Type species: Polygyra binneyana Pilsbry,
1899, by original designation.

Definition: The first Patera to possess char-
acter #99, and all of its descendents.

Species: P. (V.) binneyana (Pilsbry, 1899);
P. (V.) clenchi (Rehder, 1932); P. (V.) indian-
orum (Pilsbry, 1899); P. (V.) kiowaensis (Sim-
pson, 1888); P. (V.) leatherwoodi (Pratt,
1971); P. (V.) roemeri (Pfeiffer, 1848).

Unnamed Clade Comprising P. (Patera) and
P. (Ragsdaleorbis)

Definition: The first Patera to possess char-
acter #100, and all of its descendents.

Subgenus Patera (Patera) s.s.

Definition: The first Patera already having
character #100, to possess character #101,
and all of its descendents.

Species: P. (P.) appressa (Say, 1821); P.
(P.) clarki (Lea, 1858); P. (P.) laevior (Pilsbry,
1940); P. (P.) panselena (Hubricht, 1976); P.
(P.) perigrapta (Pilsbry, 1894b); P. (P.) sargen-
tiana (Johnson & Pilsbry, 1892).

Subgenus Patera (Ragsdaleorbis)
Webb, 1954

Type species: Helix pennsylvanicus Green,
1827, by original designation.
Definition: The first Patera already having

character #100, to possess characters #102
and 103, and all of its descendents.

Species: P. (R.) pennsylvanica (Green,
1827).

Genus Inflectarius Pilsbry, 1940

Type species: Helix inflecta Say, 1821, by
original designation.

Definition: The first Mesodontini to pos-
sess character #104, and all of its descen-
dents.

Subgenus /nflectarius (Hubrichtius)
Emberton, 1991

Type species: Mesodon kalmianus Hu-
bricht, 1965, by original designation.

Definition: The first /nflectarius to possess
character #105, and all of its descendents.

Species: /. (H.) downieanus (Bland, 1861); /.
(H.) kalmianus (Hubricht, 1965)

Subgenus /nflectarius (Inflectarius) s.s.

Definition: The first /nflectarius to possess
character #106, and all of its descendents.

Species group I. (l.) edentatus: I. ((.) eden-
tatus (Sampson, 1889); I. (.) magazinensis
(Pilsbry & Ferriss, 1907).

Species group /. ((.) smithi: I. (1.) smithi
(Clapp, 1905).

Species group /. ((.) inflectus: I. (l.) арргох-
imans (Clapp, 1905); I. (l.) inflectus (Say,
1821); /. (1.) rugeli (Shuttleworth, 1852); /. ((.)
verus (Hubricht, 1954).

Species group: I. (l.) ferrissi: I. (l.) ferrissi
(Pilsbry, 1897); I. (1.) subpalliatus (Pilsbry,
1893).

Genus Fumonelix Emberton, 1991

Type species: Helix wheatleyi Bland, 1860,
by original designation.

Definition: The first Mesodontini to pos-
sess character #107, and all of its descen-
dents.

Species: F. archeri (Pilsbry, 1940); F.
christyi (Bland, 1860); F. jonesiana (Archer,
1938); F. orestes (Hubricht, 1975); F. weth-
erbyi (Bland, 1874); Е. wheatleyi (Bland,
1860).

Unnamed Clade Comprising Appalachina
and Mesodon

Definition: The first Mesodontini to pos-
sess character #108, and all of its descen-
dents.

POLYGYRID SHELL EVOLUTION 93

Genus Appalachina Pilsbry, 1940

Type species: Polygyra sayana Pilsbry,
1906, by original designation.

Definition: The first Mesodontini already
having character #108, to possess character
#109, and all of its descendents.

Species: A. chilhoweensis (Lewis, 1870); A.
sayanus (Pilsbry, in Pilsbry & Ferriss, 1906).

Genus Mesodon Férussac, 1821

Type species: Helix thyroidus Say, 1817,
by monotypy.

Definition: The first Mesodontini already
having character #108, to possess character
#110, and all of its descendents.

Subgenus Mesodon (Aphalogona)
Webb, 1954

Type species: Helix elevata Say, 1821, by
original designation.

Definition: The first Mesodon to possess
character #111, and all of its descendents.

Species: M. (Aph.) elevatus (Say, 1821); M.
(Aph.) mitchellianus (Lea, 1838); M. (Aph.) za-
letus (Binney, 1837).

Subgenus Mesodon (Akromesodon)
Emberton, 1991

Type species: Polygyra andrewsae norma-
lis Pilsbry, 1900, by original designation.

Definition: The first Mesodon to possess
characters #112 and 113, and all of its de-
scendents.

Species: M. (Akr.) altivagus (Pilsbry, 1990);
M. (Akr.) andrewsae Binney, 1879; M. (Akr.)
normalis (Pilsbry, 1900).

Subgenus Mesodon (Mesodon) s.s.

Definition: The first Mesodon to possess
characters #114 and 115, and all of its de-
scendents.

Species: M. (M.) clausus (Say, 1821); M.
(M.) sanus (Clench & Archer, 1933); M. (M.)
thyroidus (Say, 1817); M. (M.) trossulus Hu-
bricht, 1966.

Shell-Based Phylogenetic Analysis

Figures 9-11 present x-ray outlines of 57
shells representing polygyrid subgenera and
outgroups. These outlines were used for phy-
logenetic character analysis. Also included in
the character analysis were the x-rayed

shells themselves, plus four shells from the
ANSP collection representing four additional
species: Ashmunella angulata, Stenotrema
(Archerelix) barbigerum, S. (Pilsbrelix) ex-
odon, and S. (Toxotrema) hirsutum. No shell
was available of Hochbergellus hirsutus, but
its published description (Roth & Miller, 1992)
was used to score as many characters as
possible. Thus, a total of 62 species were
included, of which only five lacked x-ray
data.

Table 2 defines the 14 characters and 71
character states scored for phylogenetic
analysis, which are illustrated in Figures 12,
13, and 6. All of these characters and char-
acters states are new or newly evaluated.

Figure 14 shows the distributions of shell
characters among subgenera and species.
Figure 15 diagrams shell characters that
were excluded from phylogenetic analysis
because of Known high levels of homoplasy
within subgenera of the Triodopsini and Me-
sodontini.

Figure 16 shows the results of cladistic
analysis of the data in Figure 14. In total,
Hennig86 generated 1,529+ equally and
maximally parsimonious cladograms with a
consistency index of 0.34 and a retention in-
dex of 0.68, of which Figure 16 is the Nelson
consensus tree.

Reliability of Fossils

Comparison of shell-based and anatomy-
behavior-shell-based phylogenetic hypothe-
ses revealed conspicuous differences in res-
olution and topology. Among 54 ingroup
(polygyrid) taxa, with a maximum possible
resolution of 53 nodes, the shell hypothesis
(Fig. 16) had 10 nodes (19% of maximum)
and the anatomy-behavior-shell hypothesis
(Fig. 8) had 40 nodes (75% of maximum).
Thus, the anatomy-behavior-shell hypothesis
had four times the resolution of the shell-
based hypothesis.

Topologically, the only congruence be-
tween the two hypotheses was т the
Stenotremini, which both showed as a mono-
phyletic clade with Euchemotrema and
Stenotrema (Cohutta) at its base. Other
clades showed major discrepancies between
the two phylogenetic hypotheses. Thus, rel-
ative to the anatomy-behavior-shell hypoth-
esis/revision, the shell-based hypothesis (a)
grouped Vespericola and Appalachina, Dae-
dalocheila (Upsilodon) hippocrepis and Lo-
bosculum, and Polygyra and Millerelix (Prat-

94 ЕМВЕАТОМ
AID

IGE 9!

FIGS. 9-11. Shell ontogenies, from x-rays of adult shells, of representatives of polygyrid subgenera and of
polygyrid closest outgroup families. The representatives are type species unless otherwise indicated. All
scale bars = 3 mm. The shells are arranged by a previous classification (Webb, 1974; Richardson, 1986),
since revised (Emberton, 1994a, this paper). Dotted lines indicate portions of the x-rays that were difficult
to interpret. By chance the x-rayed shell representing Ashmunella (As) was broken internally; all characters
could be scored from this drawing, however, so the specimen was not replaced. Specimen abbreviations,
defined in alphabetical order with their catalog numbers at the Academy of Natural Sciences of Philadel-
phia, are: AIA, Allogona (Allogona) profunda, 77867; AID, Allogona (Dysemdoma) townsendiana, 100390;
App, Appalachina sayana, 139140; As, Ashmunella rhyssa, 166077; CB, Cryptomastix (Bupiogona) hena-
ersoni, 171267; CC, Cryptomastix (Cryptomastix) mullani, 171245; CM, Cryptomastix (Micranepsia) ger-
mana, 11154; DD, Daedalochila (Daedalocheila) auriculata, 57070; DU, Daedalochila (Upsilodon) hippocre-
pis, 84629; DUa, D. (U.) acutidentata, 166418; E, Euchemotrema leai, 172539; F, Fumonelix wheatleyi,
169691; G, Giffordius pinchoti, 150735; IH, Inflectarius (Hubrichtius) downieanus (non-type), 91035; Il,
Inflectarius (Inflectarius) inflectus, 91616; 11, /. (1.) ferrissi, 98085; Lb, Lobosculum pustula, 86968; Ln, Linisa
anilis, 166371; Lnb, Linisa behri, 166487; MeA, Mesodon (Aphalogona) elevatus, 81161; MeK, Mesodon
(Akromesodon) normalis, 169640; MeM, Mesodon (Mesodon) thyroidus, 71950; MiM, Millerelix (Millerelix)
mooreana, 158375; MiP, Millerelix (Prattelix) plicata, 143448; NA, Neohelix (Asamiorbis) dentifera, 78876;

(continued)

POLYGYRID SHELL EVOLUTION 95

PrE

IG 510:

NN, Neohelix (Neohelix) albolabris, 75843; NS, Neohelix (Solemorbis) solemi, 182281; OB, outgroup Brady-
baenidae: Bradybaena similaris, 174469; OC, outgroup Camaenidae Pleurodonte lynchnuchus, 32588; OH
outgroup Helminthoglyptidae: Helminthoglypta tudiculata, 112911; OS, outgroup Sagdidae: Sagda cooki-
ana, 139388; OT, outgroup Thysanophoridae: Thysanophora impura, 177310; OX, outgroup Xanthony-
chidae: Cepolis cepa, 33301; PaP, Patera (Patera) appressa, 335137; PaR, Patera (Ragsdaleorbis) penn-
sylvanica, 251512; PaV, Patera (Vesperpatera) binneyana, 176765; Po, Polygyra septemvolva, 69117; PrE,
Praticolella (Eduardus) martensiana, 98578; PrF, Praticolella (Farragutia) mobiliana, 105999; PrP, Praticolella
(Praticolella) ampla, 131749; PrX, Praticolella (Filapex) jejuna, 77035; SC, Stenotrema (Cohutta) cohuttensis,
170118; SS, Stenotrema (Stenotrema) stenotrema, 169148; SSm, S. (S.) maxillatum, 170141; SSp, S. (S.)
spinosum, 11383; ST, Stenotrema (Toxotrema) hirsutum, 11396; Tb, Trilobopsis loricata, 11149; TdB,
Triodopsis (Brooksorbis) platysayoides, 183201; Тан, Triodopsis (Haroldorbis) cragini, 186723; ТОМ, Tri-
odopsis (Macmillanorbis) rugosa, 174909; TAP, Triodopsis (Pilsbryorbis) tennesseensis, 139143; TdS, Tri-
odopsis (Shelfordorbis) vulgata, 68807; Тат, Triodopsis (Triodopsis) tridentata, 211921; TdV, Triodopsis
(Vagvolgyorbis) juxtidens, 64720; V, Vespericola columbiana, 158355; W, Webbhelix multilineata, 190168;
XW, Xolotrema (Wilcoxorbis) fosteri, 157255; XX, Xolotrema (Xolotrema) denotata, 128444.

96 ЕМВЕАТОМ

TdB

TdM

OT

E 111

telix), each pair of which has been classified
into two widely separated clades; and (b)
split Ashmunella rhyssa and A. angulata,
Daedalocheila (Upsilodon) hippocrepis and
D. (U.) acutidentata, Praticolella and Lobos-
culum, Linisa anilis and Linisa behri, Millerelix
(Millerelix) and M. (Prattelix), and Polygyra
and Giffordius, each pair of which has been
classified into a single, closely related clade.
Among outgroups, the shell-based hypothe-
sis conflicted with a previous, anatomy-
based hypothesis (Emberton, 1991c; Fig. 8)

by joining Cepolis in a clade with Pleurodonte
and Sagda.

Closest Convergences in Sympatry

Figure 1 shows the closest known, field-
validated, polygyrid convergences in sympa-
try on the globose, umbilicate, flat, and tri-
dentate shell forms. As mentioned in the
Introduction, the globose case 1$ the closest
convergence and has received detailed

POLYGYRID SHELL EVOLUTION 97

TABLE 2. Shell characters used for a separate phylogenetic analysis of the Polygyridae (Fig. 16).

1:

2.

13.

14.

Shape of the aperture's generating curve throughout late-juvenile ontogeny (Fig. 12: 1). 1a,
“egg,” “pinto bean,” or “lima bean.” 1b, “kidney bean.” 1c, “bulging kidney bean.”
Approximate number of complete adult whorls (Fig. 12: 2, which shows the shell apex
down). 2a, four. 2b, five. 2c, six. 2d, eight. 2e, nine.

Approximate ratio of apertural areas three whorls apart, beginning near the apex and
ending at least 3/4-whorl before the aperture (Fig. 12: 3). 3a, 40:1. 3b, 32:1. 3c, 15:1. 3d,
9:1. Зе, 7:1.

Columellar surface of the body whorl approximately 1/4-whorl before the adult aperture
(Fig. 12: 4). 4a, flat or slightly concave, but convex at the suture. 4b, flat or slightly concave,
flat at the suture. 4c, convex throughout.

Umbilical shape (Fig. 12: 5, which outlines the x-rayed umbilici). 5a-5h, extremely narrow
to broad (some intergradation among categories).

Umbilical sutures (Fig. 13: 6). 6a, strongly to weakly shouldered. 6b, rounded, unshoul-
dered.

Umbilical-wall whorls (Fig. 13: 7). 7a, flat. 7b, slightly flat. 7c, round.

Basal denticle(s) (Fig. 13: 8). 8a, absent. 8b, baso-columellar shoulder or knob parallel to
apertural plane. 8c, partial lamella parallel to apertural plane. 8d, basal denticle parallel to
apertural plane. 8e, denticle or lamella(e) transverse to apertural plane. 8f (= Fig. 6: char-
acter 4), complete lamella parallel to apertural plane.

Parietal denticle shape (size variable) (Fig. 13: 9). 9a, absent. 9b, linear, curved toward the
umbilicus, higher toward the aperture. 9c (= Fig. 6: character 5), linear, straight, even in
height. 9d (= Fig. 6: character 6), Triangular to spatulate. Ye, linear, curved away from the
umbilicus, higher away from the aperture.

Size (Fig. 13: 10). 10a-10i, gigantic to minute (some intergradation among categories).

. Gradually increasing outward tilt of the long axis of the aperture (= Fig. 6: character 1). 11a,

throughout ontogeny or until tilting upward slightly. 11b, until tilting upward conspicuously.

. Whorl expansion rate (= Fig. 6: character 2). 12a, constant. 12b, increasing then decreas-

ing, with successive whorls always larger. 12c, increasing then decreasing, such that
successive whorls are equal.

Lower apertural lip (= Fig. 6: character 3). 13a, separate from basal shell. 13b, joined to
basal shell as a thin callus.

Palatal denticle (= Fig. 6: character 7). 14а, absent. 146, discrete, parallel to apertural plane.
14c, non-discrete, forming a thin shelf grading into the basal denticle, parallel to apertural
plane. 14d, semidiscrete, with bottom tapering and thinning, basally slanted inward into the
apertural plane.

study, including discoveries of several sites
of sympatry (Emberton, 1994b, 1995b).

The second closest shell-form conver-
gence in sympatry is on the flat form, exhib-
ited by the mesodontin Patera (Patera) laevior
and the triodopsin Xolotrema (Wilcoxorbis)
fosteri. Fieldwork in April 1981 along the
lower Ohio River Valley, visiting all stone
bluffs reasonably accessible by road, accu-
mulated collections at 22 stations (numbered
as “H-1” through “Н-22”), all material from
which is catalogued at the Field Museum of
Natural History. Both species had been re-
ported from Grand Chain, Indiana (Pilsbry,
1940), but extensive search failed to find both
in 1981, perhaps due to recent floods. Sym-
patry between Р. laevior and X. fosteri was
discovered at only one station: on a sand-

stone wall above the Ohio River at the Fern
Cliff estate, Hawesville, Hancock County,
Kentucky, 21 April 1981.

Figure 17 illustrates the external anato-
mies, shells, and dissected reproductive
anatomies of representative specimens of
Patera laevior and Xolotrema fosteri from the
Hawesville, Kentucky, site, and summarizes
their mating-behavioral differences accord-
ing to Webb (cited in Emberton, 1994a). The
shells and external bodies were virtually iden-
tical in size and shape (Figs. 1, 17), but inter-
nally X. fosteri differed from P. laevior by its
penial sheath, penial retentor muscle, retrac-
tor-muscle attachment on the vas deferens,
and thick gametolytic-gland (= spermathecal)
duct, all correlating with its internal (vs.
external) sperm exchange (Fig. 17). Ecologi-

98 ЕМВЕАТОМ

00006
PPS BR
OOO © ©,

lamer:

А ND t=

С

FIGS. 12-13. Shell characters among polygyrid subgenera, п addition to those in Fig. 6. Definitions are
given in Table 2.

POLYGYRID SHELL EVOLUTION 99

о

LS) CD

0

NS US ——

с Ted D
Te IS e
№ Ре

eCD
{<p
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10mm
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a

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cal and conchological data on the two sym-
patric populations (Emberton, unpublished)
are archived at the Academy of Natural Sci-
ences of Philadelphia.

The third closest shell-form convergence in
sympatry 1$ on the umbilicate form, exhibited
by the mesodontin Appalachina sayana and
the allogonin Allogona (Allogona) profunda. In
the northern Midwest, these species” ranges
overlap extensively (Hubricht, 1985), but ex-
amination of Field Museum collections indi-

cated only a few cases of documented sym-
patry, of which the most conchologically
similar case was at Burnside, Pulaski County,
Kentucky (Fig. 1). Extensive search of that
region in spring of 1982 failed to yield both
species, perhaps due to degradation of rem-
nant forest. Sympatry was discovered, how-
ever, at a site on the northern slope of Pine
Mountain, Harlan County, Kentucky (station
“GS-116’’). At that site, the shell convergence
was not as extreme as in Figure 1, because

EMBERTON

100

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чо d ооочо RR RE WR TOTER RR A TR OT RR Te: TUT EA AI Ù d Ù dd TT RK OÙ DU À DU AU U U RS U U U HOH
| © © © © RI GC RU TC UT RT TR RU TI AT U TR OT чо RT AR RI vd A TA À ооо вала вая TU U U U U RO U U ©
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2, etc.).

FIG. 14. Distributions of shell characters among polygyrid subgenera. Characters are numbered as in Table

2, but character-states are converted to letters (a = 1, b

POLYGYRID SHELL EVOLUTION 101

de mer oil

FIG. 15. Shell variation discounted as characters: a, low-to-high spires are documented within Patera s.s.;
b, rounded-to-angulate-to-keeled peripheries occur within Xolotrema s.s., X. (Wilcoxorbis), and Patera s.s.;
c, covered-to-open umbilici are known within both Patera (Vesperpatera) and Mesodon s.s.; d, smooth-
to-ribbed sculpture appears within Xolotrema s.s., Triodopsis s.s., Г. (Pilsbryorbis), and T. (Haroldorbis); e,
bald-to-hirsute sculpture occurs within Xolotrema s.s., Inflectarius (Summinflectarius), and Fumonelix; f,
unicolor-to-color-banded shells are known within both Mesodon s.s. and M. (Aphalogona).

App. sayana tended to have a smaller shell
than Al. profunda. The northern Midwest was
not extensively surveyed during 1982 and
1983 fieldwork, however, so closer conver-
gences in sympatry on the umbilicate shell
form may exist in that region.

Least close—but still remarkable, espe-

cially in apertural-barrier construction—of the
four polygyrid shell-form convergences in
sympatry (Fig. 1) is on the tridentate form.
The closest field-documented case of triden-
tate convergence in sympatry comprises the
mesodontin Inflectarius (Inflectarius) inflectus
and the triodopsin Triodopsis (Triodopsis) fal-

102 ЕМВЕАТОМ

FIG. 16. Shell-based consensus tree of polygyrid subgenera resulting from cladistic analysis of data in Fig.
14.

POLYGYRID SHELL EVOLUTION 103

Lengthy courtship and mating
Intertwining of penes

External deposition of sperm |
mass on mates everted penis

|
2 <
EN

K Duct of
\ gametolytic gland
vas №7
DEFERENS () | ]

RETRACTOR

PENIS

Brief courtshipand mating
Insertion of penes

Internal deposition of sperm
mass in spermathecal duct

RETRACTOR

RETENTOR
MUSCLE

FIG. 17. Closest known convergence in sympatry on the flat shell form: Patera laevior (left) and Xolotrema
fosteri (right) from Hawesville, Hancock County, Kentucky. Center: external anatomies and shells in two
views. Bottom: dissected reproductive anatomies. Sides: mating-behavioral differences according to Webb

(cited in Emberton, 1994a).

lax, which | collected together under the
same log in dense woods, Vinton County,
Ohio, in 1979 (vouchers at Field Museum of
Natural History, Chicago). The ranges of
these two species overlap moderately in the
central Midwest, and other cases of sympa-
try are documented— sometimes as inadvert-
ently mixed lots—in the collection of the
Academy of Natural Sciences of Philadelphia
(Emberton, unpublished).

Shell Barriers and Water Loss

Table 3 gives the results of the experiment
on Triodopsis tridentata. A total of 280 “snail-
hours” were recorded. Snails from which ap-
ertural barriers had been removed lost water
by evaporation faster than those with barriers
left intact: 27% faster when inactive (re-
tracted into the shell) and 9% faster when
active (extended from shell).

104 ЕМВЕАТОМ

Snails with intact barriers were 83% more
successful than those from which barriers
had been removed in forming complete or
partial epiphragms. Epiphragms slightly re-
duced the rate of evaporative water loss in
snails with barriers intact (by 3% for both
partial and complete epiphragms). In snails
from which barriers had been removed, how-
ever, epiphragms greatly reduced the rate of
water loss: by 22% for partial epiphragms
and by 38% for complete epiphragms.

Remaining Conservation Priorities

Table 4 gives highest priorities for polygy-
rid conservation for each of four, phylogeny-
based categories. In the category of radiat-
ing, endemic clades, Fumonelix must rank
very high; the genus 1$ restricted to the
southern Appalachians, and three of its six
species are narrow-range endemics that
have been officially designated as endan-
gered or threatened: F. archeri, F. jonesianus,
and F. orestes. In the same category, al-
though Mesodon (Akromesodon) contains
the widespread and relatively common spe-
cies M. normalis, its other two species both
have narrow, high-altitude ranges: M. altiva-
gus оп summits of the Great Smoky Moun-
tains, and M. andrewsae on the summit of
Mount Rogers, Virginia.

Four species have been listed as high pri-
orities for conservation in the category of ex-
tremely autapomorphic endemics. The two-
species genus Giffordius, endemic. to
Colombia’s tiny Isla de Providencia (off the
eastern coast of Nicaragua), is unique within
the family for its ovoviviparity; Giffordius pin-
choti by far is the rarer and most endangered
of the two species (Emberton, unpublished).
Inflectarius ferrissi, endemic to high eleva-
tions of the Smoky Mountains, represents

extreme phylogenetic shifts in both shell
morphology (Emberton, 1991a, 1991b) and in
penial morphology (Emberton, 1991a). Triod-
opsis platysayoides, endemic to a few bluffs
along the New River Gorge of northeastern
West Virginia, U.S.A., also embodies extreme
phylogenetic divergences in both shell and
penis (Emberton, 1988a). Mesodon chilhow-
eensis is remarkable for its gigantic, broadly
umbilicate shell, exhibiting nearly regular,
log-spiral growth (Emberton, 1994a: fig. 1)
and for its extremely long penis, much longer
than the diameter of the shell (Emberton,
1991а).

Giffordius also deserves high priority for
conservation in another category, as relic sis-
ter-group to a major clade. According to the
general phylgenetic hypothesis (Fig. 8), Gif-
fordius is basal to the Polygyrini, hence is
sister-group to the remaining Polygyrini, a
highly diverse and speciose clade.

The conservation category of localities
with diverse sympatric convergences 15 led
by the northern slope of Pine Mountain, Har-
lan County, Kentucky. This 1$ the only known
site where convergences on all four polygyrid
shell forms (Fig. 1) coexist. There the globose
form is represented by Mesodon zaletus and
Neohelix albolabris; the umbilicate form by
Appalachina sayana and Allogona profunda
(mentioned above); the flat form by Patera
appressa and Xolotrema denotata; and the
tridentate form by /nflectarius inflectus, Tri-
odopsis vulgata, and Т. tridentata (Emberton,
19950). This site is also important as North
America's most diverse known locality for
land snails (Emberton, 19950).

Thus, Table 4 lists seven high priorities (in-
cluding a double listing of Giffordius) for poly-
gyrid conservation, based on phylogenetic
criteria. Five of these priorities are currently
under protection. The Smoky Mountains are a

TABLE 3. Effect of apertural dentition and epiphragm on the rate of
evaporative water loss in Triodopsis tridentata.

Category

Toothed, no epiphragm
Toothed, partial epiphragm
Toothed, complete epiphragm
Toothless, no epiphragm
Toothless, partial epiphragm
Toothless, complete epiphragm
Toothed, after activity
Toothless, after activity

Total “Snail-Hours”

Number of Mean Rate of
“Snail-Hours” Water Loss

70 3.86%
59 3.74%
26 3.73%
63 4.90%

Z 3.97%
20 3.06%
25 15.80%
10 17.18%

280

POLYGYRID SHELL EVOLUTION

TABLE 4. Conservation high priorities for polygyrids, based on four phylogenetic criteria.

Criterion High Priority

Fumonelix
Mesodon (Akromesodon)

Radiating, endemic clade

Radiating, endemic clade

Extremely autapomorphic
endemic

Extremely autapomorphic
endemic

Extremely autapomorphic
endemic Triodopsis platysayoides

Extremely autapomorphic Mesodon

Giffordius pinchoti

Inflectarius ferrissi

endemic chilhoweensis
Relic sister-group to

major clade Giffordius
Diverse sympatric four shell-form

convergences convergences

105
#Spp Locality Protected?
6 Southern Blue Ridge, U.S.A. Yes
3 Southern Blue Ridge, U.S.A. Yes
1 Isla de Providencia, Colombia No
1 High Smoky Mountains, U.S.A. Yes
1 New River Gorge, U.S.A. Yes
1 Smoky Mountains, U.S.A. Yes
2 Isla de Providencia, Colombia No

11 Pine Mountain, Kentucky, U.S.A. No

U.S. National Park and International Bio-
sphere Reserve, protecting four species of
Fumonelix, two species of Mesodon (Akrome-
sodon), Inflectarius ferrissi, and Mesodon
chilhoweensis. The remaining Fumonelix are
protected in U.S. National Forests, and the
remaining M. (Akromesodon) is protected by
Mount Rogers State Park. Triodopsis
platysayoides is somewhat protected in Coo-
pers Rock State Forest, West Virginia, U.S.A.

Two high-priority sites remain unprotected.
There is a “Pine Mountain State Park,” Bell
County, Kentucky, but it is about 64 km away
from and has a much lower diversity than the
Harlan-County Pine Mountain site listed in Ta-
ble 4 (Emberton & Petranka, unpublished).
Thus, the Pine Mountain site, U.S.A., is un-
protected.

Also unprotected 1$ the conservationally
important, small Isla de Providencia, Colom-
bia. In 1987, both species of Giffordius were
still surviving on the island in remnant
patches of forest, primarily at higher eleva-
tions, but deforestation had already de-
stroyed the type locality of G. pinchoti and
seemed to be rapidly advancing up the cen-
tral peak (Emberton, 1992, unpublished).

DISCUSSION
General Phylogenetic Hypothesis/Revision

This hypothesis/revision culminates the
author's 15 years of work on polygyrid sys-
tematics, and hopefully provides a replicable
data set and a fully testable hypothesis upon
which future workers may build. For allozyme

data and for more detailed phylogenetic hy-
potheses on the Triodopsini and the Meso-
dontini, see Emberton (1988a, 1991a, 1994).
For a polygyrid biogeographic/historical hy-
pothesis, see Emberton (1994a). Clearly, this
is not the final word; much remains to be
learned.

Shell-Based Phylogenetic
Analysis/Reliability of Fossils

Despite the discovery of many new char-
acters, shell-based phylogenetic analysis re-
sulted in very low resolution. Two possible
sources of additional shell characters were
neglected, however: shell-surface micro-
sculpture (Emberton, 1995b) and shell ultra-
structural layers (Boggild, 1930; Wilbur 8
Saleuddin, 1983; Roth, 1987; Watabe, 1988).
Both of these should be investigated. Quali-
tative character analysis of x-ray data failed
to detect a difference in whorl expansion rate
between the Triodopsini and the Mesodontini
suggested by quantitative analysis (Ember-
ton, 1994a). Thus, more subtle analysis of
x-rays could yield more characters for phylo-
genetic resolution. Based on current data,
however, “shells do not tell” the estimated
145 million years of phylogenetic history of
the Polygyridae in North America (Emberton,
1994a). This implies that identification of pre-
Miocene polygyrid fossils may be very diffi-
cult at best.

Thus, based on the general phylogenetic
hypothesis/revision, convergences in shell
morphology were rampant within the Poly-
gyridae. “Kidney-bean” generating curves
cropped up in one subgenus each of Poly-

106 EMBERTON

дуга and Daedalochila. Whorl-counts of
over six hypothetically evolved one or more
times each in the Triodopsini, Vespericolini,
Stenotremini, Polygyrini, and Mesodontini.
Extremely low whorl-expansion rates (< 2.7
per 360-degree rotation) seemingly evolved
independently in the Triodopsini, Stenotrem-
ini, and Polygyrini. Convergences on a col-
umellarly flattened body whorl seem to have
occurred in all polygyrid tribes except the
Ashmunellini. Expanded umbilici (Fig. 12:
char 5e-g) appeared in all tribes but the Ves-
pericolini. Rounded, unshouldered umbilical
sutures seems to be a good synapomorphy
of the genus Triodopsis, but nevertheless this
character-state was apparently reversed in
the nominal subgenus and seems to have
been converged upon in the outgroup family
Camaenidae. Slightly flattened umbilical-wall
whorls seemingly arose convergently within
one or more subgenera each of the Triod-
opsini, Allogonini, and Mesodontini. Although
transverse-to-spiral basal lamellae in Ash-
munella, Daedalochila, and Appalachina were
hypothetically plesiomorphic and hence not
necessarily convergent, all other forms of
basal denticles and lamellae seem to have
been converged upon repeatedly in the Poly-
gyridae, with the single exception о the com-
plete basal lamella, which is a hypothetical
synapomorphy of the Stenotremini. Two
types of parietal denticles—straight and tri-
angular-to-spatulate—seem to be good syn-
apomorphies, with additional phylogenetic
information, for the Stenotremini and the
Polygyrini, although the latter was apparently
lost secondarily in Practicolella; a down-
curved parietal denticle, however, seems to
have evolved independently in all other tribes
of the Polygyridae (non-type species of Ves-
pericola also have it: Pilsbry, 1940). Both ex-
tremes of shell size occur among the out-
groups, and various intermediate and small
sizes recur repeatedly among polygyrid sub-
genera, with only very slight tendencies to-
ward trends within and among tribes. Rapid
shifts in apertural tilt and expansion rate are
apparent synapomorphies for a subset of the
Polygyrini, yet both were hypothetically re-
versed in type species of some genera. An
adnate, callus-like apertural basal lip may
seem a perfect synapomorphy for four of
Stenotrema's five subgenera, but was appar-
ently reversed in S. (Archerelix) barbigerum
(Pilsbry, 1940). Regarding apertural palatal
denticles, discrete denticles (apertural barri-
ers) seem to appear sporadically in nearly all

polygyrid tribes; and semidiscrete, basally
recessed denticles seem to be only an incon-
sistent synapomorphy of the Polygyrini; but a
non-discrete ‘‘shelf” seems to be a good sy-
napomorphy of the Stenotremini.

Closest Convergences in Sympatry

Identifications and field verifications of the
closest convergences in sympatry on the flat,
umbilicate, and tridentate polygyrid iterated
shell forms (Fig. 1) provide starting points for
analyses of these naturally replicated exper-
iments in evolutionary morphology, such as
those already conducted on the globose shell
form (Emberton, 1994b, 1995a). Although
these four cases of polygyrid shell conver-
gence in sympatry (Fig. 1; Emberton, 1994a)
are the most precise known in North Amer-
ica, they are representative of numerous less
precise cases involving the same four basic
shell forms. Among and within these four
shell forms, there are other examples of con-
vergence (Emberton, 1988a, 1991a, 1994a),
the most striking (and informative for polygy-
rid evolution) of which is between /nflectarius
ferrissi and Neohelix dentifera (Emberton,
1991b).

Shell Barriers and Water Loss

The experimental results from Triodopsis
tridentata suggest that apertural barriers re-
duce the rate of evaporative water loss both
directly and indirectly, by aiding in the forma-
tion of an epiphragm. The manner in which
barriers and epiphragms retard water loss
may be counter-intuitive, judging from Ram-
say's (1935) experiments on evaporation
rates from vertical tubes. Ramsay observed
faster water loss when the evaporating sur-
face was at the bottom of the tube than at the
top, but found that the presence of a perme-
able occluder tended to diminish the differ-
ence. He hypothesized therefore that “when
[the evaporating surface] is at the bottom...
there is an upward current of moist air which
sets up a circulation in the system.” Ram-
say’s hypothesis has not been tested, to my
knowledge. Extending his hypothesis to land
snails, when a snail withdraws deeply (to es-
cape predation, for example) into the coiled
tube of its shell, its apertural barriers (and the
epiphragm they aid in forming) may function
not so much to physically block diffusing wa-
ter vapor as to interrupt the convection cur-

POLYGYRID SHELL EVOLUTION 107

rents that normally result from tubular evap-
oration.

The trend for apertural barriers in 7. triden-
tata and other triodopsins to be larger in
moister habitats (Emberton, 1988a) suggests
that their demonstrated retardation of evap-
orative water loss is not their only—or even
primary—function. Other hypothesized func-
tions are given in the Introduction.

Polygyrids provide some fascinating ex-
amples of apertural obstruction that could be
used to test these and other hypotheses. Fig-
ure 2 presents some extreme examples from
several clades, most of which have arisen by
evolutionary convergence. The most extreme
cases are in the genus Stenotrema; it is a
memorable experience to watch one of these
snails extend, its body “pouring” like molas-
ses through the slit-like, convoluted aperture.
This slit seems to be at its most narrow in $.
(Pilsbrelix) uncifera, but is augmented in S.
(Stenotrema) maxillatum (both in Fig. 2) by
recession of the basal lamella, which lacks a
notch and which 1$ overlapped by the parietal
lamella to form a three-dimensional baffle.

Species of Daedalochila have some com-
plex apertural conformations, the most bi-
zarre of which are the deeply recessed pari-
etal scoop in D. (Upsilodon) hippocrepis and
the convoluted apertural lip incorporating the
parietal scoop and augmented by recessed
denticles in D. (Daedalochila) uvulifera (both
in Fig. 2). Other notable examples occur in
Triodopsis, Lobosculum, Ashmunella, Linisa,
Lobosculum, and Inflectarius (some are Шиз-
trated in Figs. 1 and 2).

Remaining Conservation Priorities

The evaluation of conservation priorities
presented in this paper is doubtless biased
by the author's limited experience with the
western and subtropical species. Table 4 15
intended only as a preliminary guideline and
as a possible format for future, more enlight-
ened assessments.

Given these caveats, Table 4 suggests that
most phylogenetically important polygyrids
are reasonably well protected against extinc-
tion due to habitat loss. How well they and
other polygyrid species will withstand envi-
ronmental degradation due to acid rain
(Graveland et al., 1994), local extinctions of
forest-canopy tree species (Getz & Uetz,
1994), and global warming, remain to be dis-
covered, however. Unfortunately, there is no
long-term monitoring study being conducted

on any polygyrid population or community, to
my knowledge.

Two remaining conservation priorities do
demand rapid attention, however: Isla de
Providencia, Colombia, and Pine Mountain,
Harlan County, Kentucky, U.S.A. (Table 4).
The high conservation importance of these
sites almost assuredly applies to many other,

lesser-known, leaf-litter/soil invertebrates
that polygyrids represent.
ACKNOWLEDGEMENTS

Supported in part by National Science
Foundation grants BSR-8700198 and DEB-
9201060 and by Academy of Natural Sci-
ences of Philadelphia (ANSP) discretionary
funds. | am also grateful to S. Schaefer and
G. Bohlke for access to the x-ray facility, Ich-
thyology Department, ANSP; to E. Gitten-
berger for steering me to the Christelow and
Falkner papers; and especially to T. Pearce,
B. Roth, and an anonymous reviewer for very
useful comments on a previous draft of the
manuscript. | would like to take this opportu-
nity to thank fellow polygyrid workers who
have generously helped me with this and
other projects during the past 15 years: T.
Asami, K. Auffenburg, N. Babrakzai, A.
Bogan, J. Burch, R. Caldwell, H. L. Fair-
banks, H. Feinberg, Н. В. Foster, В. Fulling-
ton, G. Goodfriend, F. W. Grimm, L. Hubricht,
E. Keferl, G. Long, C. Mather, R. Maze, G.
McCracken, W. Miller, J. Murray, R. Neck, T.
Pearce, J. Petranka, W. Pratt, R. Reeder, B.
Roth, R. Selander, the late A. Solem, A.
Stiven, В. Taylor, Е. Thompson, J. Vagvolgyi,
A. Van Devender, W. Van Devender, and G.
Webb.

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MALACOLOGIA, 1995, 37(1):

111-122

GEOGRAPHIC DISTRIBUTION AND SHELL COLOUR AND BANDING
POLYMORPHISM IN MARGINAL POPULATIONS OF СЕРАЕА NEMORALIS
(GASTROPODA, HELICIDAE)

A. Honék

Department of Entomology, Research Institute of Plant Production, Ruzyné 507, 161 06
Praha 6, Czech Republic

ABSTRACT

Occurrence and colour and shell banding polymorphism were investigated in populations of
Cepaea nemoralis (L.) at the edge of the species’ geographic distribution in the Czech Re-
public. Cepaea nemoralis populations were found only at intravillan localities. The great majority
of town and village localities were occupied by C. nemoralis in three isolated areas: a northern
one integrated with the main geographic area of the species, and two isolated southern insular
areas. А few populations were scattered at localities between these areas and further south,
where C. nemoralis was replaced by Cepaea hortensis (Múller). The C. nemoralis distribution
area and abundance may have increased since 1950. | speculate that human activity encour-
aged the spread of С. nemoralis populations, and this species competitively exterminated С.
hortensis populations from the intravillan habitats. The frequency of shell colour and banding
morphs in local populations was similar to those in oceanic Western Europe (56.2 + 18.6% of
pink shell colour), and may be affected by climatic selection and random drift. There exist small
areas marked by a high frequency of 00000, 00300, 00345 and 12345 shell banding morphs.

Their edges were mostly not concordant with areas of geographic distribution.

INTRODUCTION

Cepaea nemoralis (L.) is a West-European
species distributed from southern Scandina-
via, Lithuania and the Ukraine in the east, to
Hungary and the northern Balkan Peninsula in
the south (Schilder 8 Schilder, 1953, 1957).
However, it is absent in the highlands of cen-
tral Europe, in the territories of the Czech Re-
public and Slovakia. Only Bohemia (the west-
ern half of the Czech Republic) is crossed by
the edge of the area of species’ continuous
distribution (Fig. 1). From this region Lozek
(1956) listed a number of localities scattered
mostly north of the Labe (Elbe) River where
C. nemoralis was found chiefly at intravillan
sites. However, the precise borderline of
the species distribution is impossible to trace
from the published data. The shell colour
and banding polymorphism in Bohemian pop-
ulations has never been quantitatively inves-
tigated despite the fact that variation in
marginal populations is of considerable the-
oretical interest.

Another common west European species
is Cepaea hortensis (Múller). It lives in low-
land and submontane areas of the whole ter-
ritory of the Czech Republic, at both intravil-
lan sites and in the open landscape (Lozek,

111

1956). There exist indications (see Discus-
sion) that each Cepaea species may exclude
the other from occupying the same site. A
precise delineation of the geographic distri-
bution of intravillan populations of C. horten-
sis and С. nemoralis may contribute to test-
ing this assumption. A precise study of C.
hortensis distribution in Bohemia is also not
available.

The aim of this study was: (1) mapping the
edge of the C. nemoralis distribution, (2) re-
cording shell colour and banding polymor-
phism in local populations, and (3) investigat-
ing the possible interaction of C. nemoralis
with C. hortensis populations. Distribution
and polymorphism т С. nemoralis have been
the subject of numerous studies (Jones et al.,
1977; Lamotte, 1988). The present work 1$
justified by the fact that the occurrence and
variation were never investigated in the very
eastern edge of the С. nemoralis area of dis-
tribution.

MATERIAL AND METHODS

The populations of С. nemoralis and С.
hortensis were sampled systematically at in-
travillan and open landscape habitats of

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GEOGRAPHIC DISTRIBUTION AND SHELL COLOUR 113

northern Bohemia in 1989-1991. Towns and
villages where Cepaea was found are shown
in Figure 1. At some localities, the snails were
sampled at several sites at least 100 m apart.

The shell colour was classified as pink or
yellow, although both colours varied largely
in saturation and hue. In some populations, a
large proportion of shells was very pale. п
this case, | lumped all specimens that had
even a slight trace of pink colour (usually at
the top of the shell). These were classified as
“pink,” the rest of the animals was classified
as “yellow.” For recording the shell banding
polymorphism, | used the commonly ac-
cepted notation. Visible bands are indicated
from the dorsal side by numbers 1-5, missing
bands as 0, and fusions between bands are
indicated by brackets (). The sample size
varied between 6 and 258. Proportions of col-
our and shell banding forms were calculated

for populations where at least 15 snails were
collected. The rare hyalozonate individuals
(with pale blanks instead of black bands), and
animals with bands having diluted margins
(common at some localities) were classified
as the respective band morphs. Evaluating
the geographic variation, | compared the pro-
portions of 00000, 00300, 00345, and 12345
morphs, regardless of confluences between
bands. According to the frequency of a
morph the localities were ranked into four
categories: (1) zero frequency of the morph,
(2) low frequency—quartile 1 of the series of
localities where the morph was present ar-
ranged in ascending order of the morph per-
centage, (3) medium frequency—quartile 2 of
the above series, (4) high frequency—quar-
tiles 3 and 4 of the above series.

Details of the frequencies of colour and
shell banding morphs at different localities

FIG. 1. The localities of C. nemoralis (solid circles), C. hortensis (open circles), and of mixed populations of
both species (divided circles). The areas of “continuous” distribution of С. nemoralis are delimited by heavy
lines: above—Liberec LB, left—Litomérice LT, right—Novy Bydzov NB. The area of “scattered” distribution
is fenced by a dotted line. Insert: Area shown in Fig. 1 projected onto the map of the Czech Republic. The
edge of the C. nemoralis distribution is shown by a solid line (inside the Czech Republic) and a dashed line
(after Schilder & Schilder, 1953, 1957). :

Localities of С. nemoralis (small figures): 1 Libochovany, 2 Zalhostice, 3 Litoméfice, 4 Tfeboutice, 5
LibéSice, 6 Mlékojedy, 7 Zeletice, 8 Pocaply, 9 Terezín, 10 Nové Kopisty, 11 BohuSovice, 12 Keblice, 13
Doksany, 14 Lobendava, 15 Lipová, 16 Sluknov, 17 Mikulásovicky, 18 MikulaSovice, 19 Brtniky, 20 Rum-
burk, 21 Krasná Lipa, 22 Studanka, 23 Dolní Podluzí, 24 Jifetín pod Jedlovou, 25 Rybnisté, 26 Chribska, 27
Jetfichovice, 28 Kamenicky Senov, 29 Volfartice, 30 Novy Bor, 31 Chotovice, 32 Janov, 33 Sloup, 34
Castolovice, 35 ManuSice, 36 Pise@na, 37 Ceska Lipa, 38 Staré Splavy, 39 Doksy, 40 Mafenice, 41
Marenicky, 42 Jablonné у Podjestédí, 43 Mimon, 44 Stráz pod Ralskem, 45 Hamr, 46 Brevnisté, 47 Озебпа,
48 Kfizany, 49 Ves, 50 Andélka, 51 ViSnova, 52 Minkovice, 53 Víska, 54 Kunratice, 55 Srbská, 56 Jindri-
chovice pod Smrkem, 57 Nové Mésto pod Smrkem, 58 Dolní Rasnice, 59 Krásny Les, 60 Arnoltice, 61
Frydlant у Cechách, 62 Raspenava, 63 Hejnice, 64 Bily Potok, 65 Chrastava, 66 Liberec, 67 Vratislavice, 68
Janov nad Nisou, 69 Josefüv Dül, 70 Lucany, 71 Tanvald, 72 Velké Hamry, 73 Drzkov, 74 Zásada, 75
Vrkoslavice, 76 Dalesice, 77 Pulecny, 78 Rychnov и Jablonce nad Nisou, 79 Rádlo, 80 Hodkovice nad
Mohelkou, 81 Cesky Dub, 82 Miliceves, 83 Slatina, 84 Vrbice, 85 HradiSt’ko, 86 Vysoké Veselí, 87 Cho-
mutice, 88 Ostroméf, 89 Нойсе у PodkrkonoSi, 90 Lístkovice, 91 Кпёйсе, 92 Zlunice, 93 Sekeïice, 94
Smidary, 95 Smidarská Lhota, 96 Janovice, 97 Hlusice, 98 Stary Bydzov, 99 Novy BydZov, 100 Vysoéany,
101 Prasek, 102 Mystéves, 103 Petrovice, 104 Suchá, 105 Staré Nechanice, 106 Nechanice, 107 Boharyné,
108 Skochovice, 109 Luzec, 110 Nepolisy, 111 Mlékosrby, 112 Chlumec nad Cidlinou, 113 Mnichovo
HradiSté, 114 Sedlist'ka, 115 Turnov, 116 Zelezny Brod, 117 Bozkov, 118 Sobotka, 119 Nova Рака, 120
Vrchlabi, 121 Hostinné, 122 Miletin, 123 Pardubice, 124 Libcany, 125 Hradec Králové, 126 Nedélisté, 127
Ceská Skalice, 128 Olivetin, 129 Broumov. o

Localities of С. hortensis (large figures): 1 Dolní Habartice, 2 Benesov nad Ploucnicí, 3 Zandov, 4
Stvolinky, 5 Kravaïe, 6 USték, 7 Zahradky, 8 Jestrebi, 9 Holany, 10 Dfev£ice, 11 Спит, 12 Vrchovany, 13
Dubá, 14 Pavlicky, 15 ZakSin, 16 Polepy, 17 Host'ka, 18 Snédovice, 19 Stéti, 20 Béla pod Bezdézem, 21
Klaster Hradisté, 22 Kosmonosy, 23 Mladá Boleslav, 24 Dobrovice, 25 Dolni Bousov, 26 Liban, 27 Chylice,
28 Kostelec, 29 Kopidino, 30 Cesov, 31 Chroustov, 32 Chotusice, 33 Dymokury, 34 Záhornice, 35 Méstec
Králové, 36 Lovüice, 37 Svijansky Ujezd, 38 Hubálov, 39 RadoSovice, 40 Jesenny, 41 Jablonec nad Jizerou,
42 Rovensko pod Troskami, 43 Lomnice nad Popelkou, 44 LibStät, 45 Jilemnice, 46 Horní Branná, 47 Jicín,
48 Sárovcova Lhota, 49 Lázné Bélohrad, 50 Borovnice, 51 Dolní Kalná, 52 Lanzov, 53 Dvür Králové, 54
Horicky, 55 Chvalkovice, 56 ReSetova Lhota, 57 Jaromér, 58 Semonice, 59 Cernozice, 60 Smifice, 61
Probluz, 62 Stézery, 63 Roudnice, 64 Kosicky, 65 Dobfenice, 66 Chyst', 67 Rohovládova Béla, 68 Holice,
69 Cernilov, 70 Jasenná, 71 Оробпо, 72 Tyniste nad Orlici, 73 Vamberk, 74 Praha, 75 KarlStejn.

114 HONEK

with an analysis of linkage disequilibria will be
published in a separate paper (Honek, in
prep.). The data may be also obtained on re-
quest from the author.

RESULTS
The Habitats of Cepaea populations

In Bohemia, typical habitats of C. nemora-
lis are intravillan areas. Most often the snails
were found at ancient or working cemeteries,
particularly along the south- and west-facing
walls. Typical localities were also the railway
stations surrounded by exuberant weedy
vegetation. Other sites often populated by C.
nemoralis included small gardens, weed
stands bordering the margins of quiet
streets, and intravillan shrubs on south-fac-
ing slopes. Many sites were periodically dis-
turbed by human activities: soil cultivation,
mowing, herbicide application, transport or
building activities. Despite an intensive
search, по С. nemoralis populations were
found outside the intravillan areas.

Populations of С. hortensis in towns and
villages (Fig. 1) occurred in habitats similar to
those of C. nemoralis. Cepaea hortensis gen-
erally preferred less disturbed sites, and its
populations were also frequently found in the
open landscape (localities not shown in Fig.
1). The preferred rural habitats were hedge-
rows and shrubs, south-facing slopes with
mixtures of dicotyledonous and grassy veg-
etation, and roadside ditches.

The distribution of Cepaea populations

Cepaea nemoralis was found at 129 town
and village localities of northern Bohemia
(Fig. 1). Most localities were grouped in three
areas where the species occupied nearly all
favourable intravillan sites and was present in
most towns and villages. These areas | call
“areas of continuous distribution” (Fig. 1, de-
limited by solid lines), and refer to by the
name of the principal included town. The
largest area of Liberec (LB, localities no. 14—
81) extends along the northern Bohemian
frontier and is probably a southern protrusion
of the area of continuous C. nemoralis distri-
bution (Fig. 1, insert). Two smaller “island”
areas of continuous distribution are the area
of Litomérice (LT, localities 1-13), and the
area of Novy Bydzov (NB, localities 82-112).

The boundaries of the areas of “continu-
ous” distribution could be traced with the
precision of a few kilometers, that 1$, the dis-
tance that divides the neighbouring villages
of which one is populated by C. nemoralis,
the other by C. hortensis populations. At the
edge of distribution of both species, for ex-
ample between Ceská Lípa (Fig. 1, locality
37) and Doksy (locality 39) in the LB area, or
along the entire west-south-east edge of the
NB area, there exist no prominent geo-
graphic structures which may cause the sep-
aration of the species. Other portions of the
boundary are bordered by areas generally
unfavourable for Cepaea (e.g., volcanic or
sandstone hills, or large forests). This is also
the case along the western and eastern sec-
tions of the boundary of the LB area, and the
northern sections of the LT and NB areas.

The LB and NB areas of “continuous” dis-
tribution are separated by an area of mosaic
distribution of intravillan С. nemoralis and С.
hortensis populations, which | refer to as the
area of “scattered” С. nemoralis distribution
(Fig. 1, dashed line). Sixteen villages and
towns with populations of С. nemoralis were
found on this territory. In some towns, for ex-
ample Pardubice (123) and Hradec Králové
(125), а С. nemoralis population was found at
one site, whereas many sites were occupied
by C. hortensis.

In total, | found only 14 sites where popu-
lations of С. nemoralis and С. hortensis lived
together at one place. Five mixed popula-
tions were found at the area of “scattered”
distribution, others at the margins of the LB
(two populations) and NB (seven populations)
areas of “continuous” distribution.

Shell Colour and Banding Polymorphism

The large variation in the proportions of col-
our morphs had a very localised character.
Thus, the extent of variation in the frequency
of the pink form was only slightly smaller
among 24 sites within the small town of Lito-
тёйсе (9.7-78.3%) than among 73 popula-
tions of the entire LB area (1.2-98.4%). The
average proportion of the pink form was 56.2
+ 18.3%. The figures for particular areas of
“continuous” distribution differed slightly.
The highest frequency of pink was in popu-
lations from the LB area, followed by the LT
and NB areas (Fig. 2). This difference was
correlated with greater average altitude (and
cooler climate) of the LB area (localities at

GEOGRAPHIC DISTRIBUTION AND SHELL COLOUR 115

230-580 m above sea level) than of the LT
(160-230 m a.s.l) and NB (190-240 т
a.s.l.) areas. However, within the LB area the
regression of the proportion of pink individu-
als on altitude was not significant (r° = 0.006,
p> 0-1) (Fig: 3):

The frequency of shell banding 00000,
00300, 00345 and 12345 morphs in local
populations (Fig. 2) showed typical area ef-
fects. The distribution of frequencies of the
00000 morph was right-skewed and popula-
tions with a high frequency of > 17% were
found in the LT and LB areas and also at the
eastern margin of the area of “scattered”
species distribution (Fig. 4). The populations
with high frequency > 46.5%, of the 00300
morph were aggregated in the LT and NB
areas and also in the north of LB area (Fig. 5).
The populations with high frequency of
> 6.6% of 00345 morph were aggregated in
the LT area and the north LB and west NB
areas (Fig. 6). The distribution of the 12345
morph was complementary to distribution of
other morphs. Populations with high fre-
quency of > 80.0% of this morph were ag-
gregated in the south of the LB area.

The fraction of fused band phenotypes
within the 12345 morph varied among popu-
lations. The frequencies of melanic (123)(45)
and (12345) morphs varied between 0.0-
45.2% and 0.0-28.4%, respectively, and
were not correlated (in LB area) with the alti-
tude of the locality.

The 00000 phenotype was associated with
pink colour, and yellow specimens were very
rare. By contrast, 00300, 00345, and 12345
morphs were not associated with any colour
phenotype. Details of linkage disequilibria
between colour and shell banding morphs
will be discussed elsewhere (Honék, in

prep.).

DISCUSSION
Spreading of C. nemoralis

Comparing the distribution of C. nemoralis
populations from before 1950 (Lozek, 1956)
with this study revealed that most localities
reported earlier (localities no. 3, 16, 21, 30,
37, 48, 66, 99, 115) were within the areas of
“continuous” distribution. The general pat-
tern of С. nemoralis distribution apparently
has not changed within the last 50 years, but
the species became more abundant since
Lozek (1956; personal communication) qual-

ified its abundance in the 1950's as “rare.” At
present, the species also occupies more lo-
calities than it did earlier. The difference be-
tween abundance estimates from before the
1950’s and in this study cannot be explained
by omissions by earlier authors but probably
indicates an increase of the area of species
distribution. The change involves southward
expansion of the LB area, and radial expan-
sion of LT and NB areas. On the other hand,
| did not find С. nemoralis at three localities
where it was established before the 1950's:
Karlovy Vary (Karlsbad) and Zerotín (both
outside the area shown in Fig. 1) and USt&k
(C. hortensis locality no. 6 in Fig. 1). These
localities are now populated by C. hortensis.
This indicates extinction of local C. nemoralis
populations. The causes of extinction are un-
clear, because ordinary human activity does
not endanger their survival. The plasticity of
the distribution of C. nemoralis at the eastern
edge of the species” distribution area con-
trasts with its constancy in Western Europe,
where some populations have persisted, with
little variation in morph frequency, since the
neolithic period (Cain & Cook, 1989).

The factors of fast spreading of an animal
with a limited dispersion capacity are of in-
terest. | suppose that human activity may be
an important factor in species dispersion.
This follows from a frequent occurrence of
marginal populations at two typical habitats:
railway stations and cemeteries. This distri-
bution may indicate the accidental passive
transport by man. Climbing onto and falling
off the railway coaches may disseminate the
snails. In fact, some marginal populations of
C. nemoralis (e.g. 113, 116, 119, 125) were
found at railway stations in towns otherwise
populated by С. hortensis. The occurrence of
C. nemoralis in cemeteries could be attrib-
uted to the popular exchange of potted
plants. The potted plants are usually distrib-
uted from large gardens to several surround-
ing villages. The soil infested with the eggs of
garden populations of C. nemoralis may be-
come the vehicle of dissemination.

Relationship to C. hortensis

In fact, С. nemoralis and С. hortensis rarely
occurred at the same place. In several towns
118; ВУ. 88; 112. 116.118, 123, 125. 127),€:
hortensis was found at several sites, but С.
nemoralis inhabited only one place. These
towns are at the margins of the areas of

116 НОМЁК
00000 00300 00345 12345 PINK

40 pe i a
40
15 10
20
20 5
5

30

20

30 60 30
40
20 40 20
20
10 20 10
25
60 60
40 15 40
20 20
5

0 40 80 0 40 80 0 40 80 0 40 80 0 40 80
MORPH FREQUENCY a

FIG. 2. The frequency of percentages of shell banding morphs 00000, 00300, 00345, 12345 and pink shell
coloration in populations of С. nemoralis. A—Pooled sample; B—Litomérice (LT) area of “continuous”
distribution; C—Liberec (LB) area of “continuous” distribution; D—Novy Bydzov (NB) area of “continuous”
distribution; E—area of ““scattered” distribution. The boundaries of the areas are shown in Fig. 1.

сл

PERCENT LOCALITIES

en 5 a
en a à
(==
[=>]

GEOGRAPHIC DISTRIBUTION AND SHELL COLOUR

117

100 . 4 3 я
= s
= © ña o > о о №. © e
O A AR
= 50 e e 5 ы ° e u
= e ы a e 3 e
1) e
a .° e 2 e

0 . ae

ALTITUDE

FIG. 3. The regression of the percentage of the pink colour form in populations of Liberec (LB) area on the
altitude (m) above sea level of the locality. Regression: у = —0.015x + 63.3, г? = 0.006, р >> 0.1.

“continuous” distribution or in the area of
“scattered” distribution. The reverse situa-
tion when С. nemoralis occurred all over the
inside of the town and С. hortensis populated
a few suburban sites, was established at
Novy BydZov (99), in the centre of NB area of
“continuous” distribution. Both examples
may show stages of invasion of a new locality
by C. nemoralis. At first, C. nemoralis is es-
tablished at one place and from there it
spreads, exterminating C. hortensis popula-
tions. At a given site, the transition is perhaps
quick, as mixed populations usually contain a
majority of one species, either C. hortensis
(28, 83, 85, 86) or C. nemoralis (69, 91, 120).
_ There exists some experimental evidence
for competition superiority of C. nemoralis
over C. hortensis (Cameron 8 Carter, 1979,
Tilling, 1985a, b), and competitive exclusion
has been proposed in explaining the distribu-
tion of both species (Boycott, 1934; Cain,
1983). The situation in Bohemia contrasts
with that in Western Europe, where mixed
populations may coexist for a long time (Cain
8 Currey, 1963a; Cain, 1983). This difference
may be due to the fact that in the western
part of its distribution area C. nemoralis lives
in the extravillan landscape where C. horten-

sis may resist its competition. Also, in Bohe-
mia C. hortensis populations live outside the
towns where intravillan sites are all occupied
by C. nemoralis.

The examples of invasion of a locality oc-
cupied by one Cepaea species by the other
species are few. Well documented 1$ a recent
(between 1961 and 1985) substitution of C.
nemoralis by С. hortensis on severeal sites at
Marlborough Downs, England (Cain 8 Cur-
rey, 1963b; Cowie 4 Jones, 1987). The trend
observed in this non-urban area may be a
consequence of the change of the microcli-
mate following the change of the vegetation
cover. А dense and tall vegetation may favour
the occurrence of C. hortensis, which makes
better use of solar radiation and is capable of
exploiting shade places. The microclimate
might favour also the changes in Bohemian
populations because intravillan sites where
C. nemoralis probably replaced C. hortensis
are generally warmer than the sites in the
open landscape occupied by C. hortensis.

Colour and Shell Banding Polymorphism

The study of C. nemoralis variation is a
problem that has resisted final solution de-
spite the large body of accumulated data

НОМЕК

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HONEK
Figure 6 here;

120

GEOGRAPHIC DISTRIBUTION AND SHELL COLOUR 121

(Schilder & Schilder, 1953, 1957; Jones et al.,
1977; Cain, 1983; Lamotte, 1988). Typical
features of this polymorphism are very local
differences in the proportion of morphs
(Wolda, 1969a, b), and the “area effects,”
that is high frequency of a phenotype in a
limited area (Cain 8 Currey, 1963b). Area ef-
fects have been observed for both shell co-
lour and banding polymorphism (Khemici et
al., 1989; Lamotte et al., 1989; Ratel et al.,
1989). The factors of differences in morph
proportions may be random population pro-
cesses (Lamotte, 1951, 1952) or selection by
climatic or substrate factors (Currey & Cain,
1968; Cameron et al., 1977; Khemici et al.,
1989; Mazon et al., 1988, 1989) or visual
predators (Cain & Sheppard, 1950).

Random processes have probably influ-
enced the composition of some of our iso-
lated populations. Morph frequencies may be
influenced by genetic drift when populations
were established from a small number of
founders. An example may be the population
at the railway station at Nová Paka (119),
which consisted of 95.6% of pink 00300 in-
dividuals.

Climatic effects contribute to maintaining
the frequencies of shell colour forms. Popu-
lations from areas with oceanic climate are
mostly pink in contradistinction to the mostly
yellow populations of areas with Mediterra-
nean climate (Mazon et al., 1988; Vicario et
al., 1988). Multivariate analysis of climatic
and substratum data revealed that these fac-
tors contribute also to maintaining small dif-
ferences in shell colour frequencies under
oceanic climate (Khemici et al. 1989, Ratel et
al. 1989).

The average proportion of pink individuals
in Bohemian populations (56.2%) was similar
to that in oceanic Western Europe (about
60% of the pink form) and far greater than
in Mediterranean populations (with only 20-
30% of pink form, Lamotte, 1988; Mazon et
-al., 1988). Typical for Bohemian populations
is also the absence of the yellow 00000 phe-
notype, which is the most adapted one to
warm conditions. A demonstration of climatic
effects on diversification of morph propor-
tions among Bohemian populations would
require more microclimatic and orographic
data. An indication of such effects was the
increased frequency of the pink form in hilly
LB compared to lowland LT and NB areas.

No explanation has been found for the area
effects shown by the 00000, 00300, 00345,
and 12345 morphs. The role of visual preda-

tion on maintenance of differences between
populations is difficult to evaluate. The local-
ities of C. nemoralis in Bohemia become cov-
ered mostly by sparse vegetation (weeds,
ornamental plants), which makes a rather
uniform optical background for avian preda-
tors. Under such conditions, visual predation
is not likely to create important differences in
morph proportions among local populations
of Cepaea. Anyway, the results do not con-
tradict the classical theory of the role of visual
selection in maintaining Cepaea polymor-
phism (Cain & Sheppard, 1950, 1954). The
remains of crushed shells were found at only
15 collection sites. This paucity of crushed
shells parallels the results of other studies
(e.g. Cowie & Jones, 1987) and may caused
by preference of avian predators for juveniles
(cf. Wolda, 1972; Wolda & Kreulen, 1973). A
comparison of morph frequency in adult
crushed shells and in living animals could be
made only at one locality (Arnoltice 60). There
was no significant difference in proportion of
colour and banding morphs among living an-
imals and dead shells.

ACKNOWLEDGEMENTS

| thank Prof. A.J. Cain, Prof. M. Lamotte
and Dr. V. Lozek for critical reading of my MS
and many valuable comments.

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Revised Ms. accepted 18 October 1994

MALACOLOGIA, 1995, 37(1): 123-132

KARYOTYPE ANALYSIS AND GENOME SIZE IN THREE MEDITERRANEAN
SPECIES OF PERIWINKLES (PROSOBRANCHIA: MESOGASTROPODA)

В. Vitturi', A. Libertini?, M. Panozzo” & G. Mezzapelle'

ABSTRACT

The diploid chromosome number 2n = 34 has been determined in early developing embryos
of L. saxatilis and male gonads of L. (Melaraphe) punctata both from the Mediterranean Sea.
The diploid value 2n = 33 occurred in spermatocytes of L (Melaraphe) neritoides specimens
from the Lagoon of Venice. As previously reported for L. neritoides from the Sicilian coast, a
male XO sex-determining mechanism seems to operate also in the population of the same
species here studied. Variation in the number of NOR-bearing chromosomes per cell has been
detected in both L. saxatilis and L (Melaraphe) neritoides. Flow cytometric DNA analysis indi-
cates that L. saxatilis and L. (Melaraphe) neritoides are endowed by almost equal genome sizes,
whereas L. (Melaraphe) punctata exhibits about 60% of their values.

Key words: periwinkles, karyology, genome size, Mediterranean Sea.

INTRODUCTION

Periwinkles are mesogastropod molluscs
belonging to the genus Littorina which inhabit
rocky coasts and lagoon brackish waters. п
the Mediterranean, this genus is represented
by three species: L. saxatilis (Olivi, 1792), L.
neritoides (Linnaeus, 1758) and L. punctata
(Gmelin, 1791). On the basis of their repro-
duction, embryogenesis, and morphology of
the radula and penis, the former belongs to
the subgenus Littorina, whereas the others
are grouped in the subgenus Melaraphe
(Nordsieck, 1968; Torelli, 1982).

Cytogenetically, two Mediterranean popu-
lations of L. neritoides (Thiriot-Quievreux &
Ayraud, 1982; Vitturi et al., 1988) and three
North Sea populations of L. saxatilis, from
western Sweden, northern England (Janson,
1983) and the Barentz Sea (Birstein 4 Mikhai-
lova, 1990), have been investigated. Never-
theless, the results of these studies do not
agree. More precisely, the diploid number of
34 chromosomes was proposed by Thiriot-
Quievreux 4 Ayraud (1982) for L. neritoides
males from Villefranche-sur-Mer, Provence,
France, whereas 2n = 33 was found in male
specimens from Palermo, Sicily, Italy (Vitturi
et al., 1988). In the latter population, the oc-
currence of an unpaired chromosome т
spermatocytes, along with n = 17 bivalents in
the female, made the authors hypothesise a
male XO sex-determining mechanism oper-
ating in this species.

Moreover, although Janson (1983) and
Birstein & Mikhailova (1990) agree upon the
chromosome number (34 in the diploid set) of
three different populations of L. saxatilis,
there are small differences related to the
karyotype morphology. In fact, three small
chromosome pairs were seen as subtelocen-
tric in the Swedish population (Janson, 1983),
whereas the same pairs were found to be
most likely telocentric in the strain from the
Barentz Sea (Birstein & Mlkhailova, 1990).

Karyological investigation in the present
report includes: (1) analysis of L. neritoides
male specimens from the Lagoon of Venice,
northeastern Italy, in order to verify if the
male XO sex-mechanism also occurs in this
geographical location; (2) a comparison
among the karyotypes of L. saxatilis from the
Mediterranean Sea and from the North Sea
previously described (Janson, 1983; Birstein
& Mikhailova, 1990); and (3) a preliminary cy-
togenetic characterisation of L. punctata,
which is still unknown.

Moreover, in order to better understand
karyological relationships in Littorinidae, nu-
cleolar organizer regions (NORs) of L. saxati-
lis and L. neritoides as well as genome sizes
in the three species have been investigated.

MATERIALS AND METHODS

Several specimens of the brooding L. sax-
atilis and sexually mature L. (Melaraphe) neri-
toides were collected along the dock base-

‘Institute of Zoology, University of Palermo, Via Archirafi 18-90123, Palermo, Italy.
2CNR- Institute of Marine Biology, Riva 7 Martiri 1364/A, Venice, Italy.
National Institute of Cancer Research, Biotechnological Section, Via Gattamelata 64, Padua, Italy.

124 VITTURI ЕТ AL.

TABLE 1. Counts of mitotic spreads in three Mediterranean periwinkles species.

SPECIES ORIGIN

Littorina saxatilis
Littorina (Melaraphe)

punctata Gulf of Palermo — —
Littorina (Melaraphe)
neritoides Lagoon of Venice — —

ments, in Venice, Italy, on October 1991 and
April 1992, respectively. Sexually mature
specimens of L. (Melaraphe) punctata were
sampled from July 1991 to August 1993 on
rocky shores at Sferracavallo, Palermo Dis-
trict, Italy. Identifications were made accord-
ing to the guidelines of Parenzan (1970) and
Torelli (1982), and voucher shells of five indi-
viduals per species were deposited at the
Museum of the Institute of Zoology, Univer-
sity of Palermo.

Mitotic metaphase chromosomes of L.
saxatilis were obtained from early developing
embryos extracted from 20 different females
and treated with 0.025% colchicine in 0.075
M KCI solution for 20 min according to the
air-drying technique. The same procedure
was applied to the testes of 15 specimens of
L. (Melaraphe) neritoides and 30 of L. (Mela-
raphe) punctata to obtain diakinetic bivalents
and spermatogonial metaphases. Slides
were stained т 5% Giemsa-phosphate
buffer solution (pH 6.8), observed, then de-
stained in ethanol and restained with silver
nitrate according to the colloidal 1-step
method (Howell & Black, 1980).

In order to obtain the karyogram of L. sax-
atilis and L. (Melaraphe) neritoides, the chro-
mosomes of five plates per species were cut
and paired on the basis of decreasing size
and centromere position. Chromosomes
were measured and classified by arm-ratio
(longer/short arm) following the nomencla-
ture proposed by Levan et al. (1964).

Genome size was evaluated through flow
cytometric assay. From dissected mantle of
8-20 specimens per species, a cell зизреп-
sion was obtained by a 5 min hypotonic treat-
ment with 0.075 М KCI solution. Cells were
filtered through a 30-um mesh, then fixed in
70% ethanol and centrifuged twice at 800 g
adding new fixative every time. Samples
were stored at —20°С.

A day before cytofluorimetric analysis,
cells were centrifuged again and resus-
pended into 1 ml of solution containing
0.12% sodium citrate, 0.005% propidium io-

2п =21 22 23 24 25 26 27 28 29 30 31/ 3223384 TOTAL

Lagoon of Venice 1 —

=P ======= 1 1,2906
ey nm u
Ze SSS 2 dE — 2

dide and 0.1% RNase. RNA digestion was
performed at 37°C for 30 min, while staining
lasted overnight at 4°C. Gill cells of the blue
mussel (Mytilus edulis) were prepared in the
same manner and used as standard. Control
assigned DNA content was 3.2 pg according
to Hinegardner (1974). An EPICS-C flow cy-
tometer (Coulter Electronics, Hialeah, Flor-
ida) was employed for DNA content mea-
surements.

A 488-nm argon ion laser was used for ex-
citation and total red fluorescence emission
was measured. At least three samples per
species each containing more than two thou-
sand cells were employed. Conditions of
analysis were set in order to obtain the modal
value of blue mussel at the channel 50 in a
256-channel DNA histogram.

The samples of periwinkle species were
run both with and without control cells; be-
cause data distribution of reference and test-
ing nuclei were partially overlapped, only in-
dividual histograms are shown.

RESULTS

The diploid value 2n = 34 has been found
in both L. saxatilis and L. (Melaraphe) punc-
tata from counts of 32 and 24 metaphases
respectively (Table 1). A few aneuploid
spreads displaying a chromosome number
lower than the mode were also encountered.
These might be related to technical short-
comings.

Littorina saxatilis complement (Figs. 1, 2;
Table 2) consisted of all bi-armed pairs
(M+SM) except for pairs 4 and 6, which were
mono-armed (ST). However, pairs 3, 7, and
12 were characterized by arm ratio on the
border line between SM and ST and thus,
considering their confidence limits (respec-
tively 1.53-2.01, 1.34-2.20 and 2.47-3.29),
these chromosome pairs may not be attrib-
uted to one or the other chromosome class.

NOR location and variation in number of
NOR-bearing chromosomes per cell were

KARYOTYPES OF PERIWINKLES 125

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FIG. 1. Karyogram obtained from five mitotic metaphases of L. saxatilis.

FIG. 2. Giemsa stained representative karyotype of L. saxatilis.

FIG. 3. Different NOR phenotypes (А, В, С, D) of L. saxatilis.

FIG. 4. Giemsa stained spermatocyte bivalents at diakinesis of L. (Melaraphe) punctata.

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detected in L. saxatilis by silver staining. chromosomes belonging to pairs 4 (“а
NORs were located on the short arms in three type), 6 (“‘b” type) and 15 (‘‘c”’ type). In the
different types of silver positively stained first case, NOR was in a large-sized subtelo-

VITTURI ЕТ AL.

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KARYOTYPES OF PERIWINKLES 127

centric chromosome and seemed to be ter-
minally or sub-terminally located in con-
densed (Fig. 3A) and decondensed (Fig. 3B)
chromosomes. In the second case it was ter-
minal in a medium-large subtelocentric chro-
mosome, and finally it resulted to be terminal
in а small-sized submetacentric chromo-
some. Four NOR phenotypes, differing for
the number of chromosomes involved in nu-
cleolus organization, were found: one char-
acterized by four chromosomes (а, a, b, c in
two spreads) (Fig. 3A), two by three chromo-
somes (a, a, b and a, b, c, in 7 and 5 plates,
respectively) (Fig. 3B, C) and one by two (a, b
in six spreads) (Fig. 3D).

In L. punctata, notwithstanding several at-
tempts during three reproductive seasons
(1991-1993), no slides gave spermatogonial
metaphases useful to define the karyotype
morphology of this species due to the over-
condensation of mitotic chromosomes.
Overall, the diploid number tendel tube 2n =
34 (Table 1). The haploid number was n = 17
(Fig. 4), and no heteropycnotic elements
were identified.

Counts of 21 spermatogonial metaphases
resulted in the diploid number 2n = 33 in L.
(Melaraphe) neritoides from the Lagoon of
Venice (Fig. 5). The Кагуодгат (Fig. 6, Table
2) consisted of 16 autosomal pairs, 12 of
which were bi-armed and four mono-armed
(pairs 4, 5, 6, 8), and one small unpaired sub-
metacentric element about 1 um long.

After silver staining, spermatogonial meta-
phases showed the occurrence of either three
or two NOR-bearing chromosomes per plate
referred as to two distinct NOR phenotypes.
The former consisted of two small-sized me-
tacentric chromosomes (pair 16) with terminal
NORs and a large subtelocentric (pair 6) with
telomeric NORs on the short arms designated
“a,” “a” and “b,” respectively (Fig. 7A), and
was observed in 15 spreads. The latter in-
cluded two small-sized metacentric NOR-
bearing chromosomes always similar to those
previously identified as “a” (Fig. 7B), and this
was found in 12 spreads.

Analysis of diakinetic plates showed 16
bivalents plus one small-sized submetacen-
tric. The latter being characterized by the
same morphology of the unpaired element
observed in mitotic metaphases, was inter-
preted as “univalent” (Fig. 8, arrow).

A typical fluorescence distribution histo-
gram in cell samples of the three periwinkle
species and the control are given in Fig. 9
(A-D). Each littorinid species is characterized

by two peaks. The first one represents diploid
cells while the second one—localized at a
double distance from the axes origin (tetra-
ploid peak)—is probably due to cell aggrega-
tion rather than G2-M phase cells. The refer-
ence exhibits one peak only.

Red fluorescence emission measurements
are summarized on Table 3. They indicate
that L. saxatilis and L. (Melaraphe) neritoides
are endowed by almost equal genome sizes
(estimated haploid DNA content of 1.352 and
1.376 picograms, respectively), whereas L.
(Melaraphe) punctata exhibits about 60% of
these values (0.811 pgs).

DISCUSSION

Karyological data concerning all periwin-
kles of the genus Littorina studied to date are
briefly summarized on Table 4. Interestingly,
a diploid number 2n = 33 occurs in males of
Littorina (Melaraphe) neritoides from the La-
goon of Venice. This result, and a karyotype
very similar to that observed for L. (Melara-
phe) neritoides from the Sicilian coast (Vitturi
et al., 1988), supports a male XO sex chro-
mosome system presumably operating in
this species. Moreover, as previously argued
(Vitturi et al., 1988), this result does not agree
with 2n = 34 suggested by Thiriot-Quievreux
& Ayraud (1982) for the males of L. (Melara-
phe) neritoides from Villefranche-sur-Mer,
France. Because of the hypothesis of a chro-
mosomal polymorphism in this species has
already been excluded (Vitturi et al., 1988), a
karyotype revision of the French population
would be desirable.

Male XO sex determination mechanism
has been described in some grasshoppers
(Cabrero et al., 1985; Vitturi et al., 1993) and
fishes (Chen, 1969; Le Grande, 1975). How-
ever, according to our knowledge, it has
never been reported in mesogastropods, be-
Cause neritid species, possessing this sex-
determination mechanism (Vitturi & Catalano,
1988, and authors quoted by them) belong
to Archaeogastropoda (Franc, 1968; Boss,
1982).

The same chromosome number (2n = 34)
but with small differences in karyotypes have
been found in three out of four L. saxatilis
populations analyzed so far. Observed differ-
ences are to be probably related to the fact
that some chromosome pairs may be close
to the limits between different chromosome
categories.

128 VITTURI ЕТ AL.

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FIG. 5. Giemsa stained karyotype from male gonads of L. (Melaraphe) neritoides.

FIG. 6. Karyogram obtained from five spermatogonial metaphases of L. (Melaraphe) neritoides.

FIG. 7. NOR phenotypes (A and В) of L. (Melaraphe) neritoides (д = giemsa stained and п = silver stained).
FIG. 8. Diakinetic bivalents from male gonads of L. (Melaraphe) neritoides (arrow indicates the unpaired
chromosome). Scale for figures from 1 to 8: 10 um = 26 mm

Considering NOR patterns determined for 3. at least three chromosome pairs are in-
L. (Melaraphe) neritoides and L. saxatilis, volved in nucleolus organization in L.
some considerations can be drawn: saxatilis, whereas only two operate in L.

oi (Melaraphe) neritoides. This implies a
1. intra-specific polymorphism due to dif- smaller variation of NOR patterns in the

ferent number of silver-positive chro- second species.

mosomes per cell occurs in both spe-

cies; On the whole, data from the literature doc-

2. chromosomal location of active NOR ument a wide distribution of NOR polymor-

sites differs from one species to an- phism within animal kingdom (Foresti et al.,

other; 1981; Gold & Amemiya, 1986; King et al.,

KARYOTYPES OF PERIWINKLES 129

number of cells

red fluorescence

Littorina (Melaraphe) punctata

43 C

Littorina (Melaraphe) neritoides

42 B

Littorina saxatilis

50 D

Mytilus edulis

FIG. 9. A-D. Histograms of red fluorescence emission in cell suspensions obtained from three species of
Littorinidae and Mytilus edulis (control). A: L. (Melaraphe) punctata; В: L. saxatilis; С: L. (Melaraphe) neri-
toides; D: Mytilus edulis. Red fluorescence emission and number of scored cells are expressed in arbitrary

units.

1990; Vitturi et al., 1991a). Particularly, in
mollusks, intraspecific NOR variations have
been described for two prosobranch gastro-
pods (Vitturi & Catalano, 1989, 1990), two
pulmonates (Vitturi et al., 1991b; Vitturi, 1992)
and four oysters (Thiriot-Quievreux & Insua,
1992; Insua & Thiriot-Quievreux, 1993).
Haploid genome sizes (C-value) vary from
0.8 to 1.3 pg in five littorinid species (Hine-
gardner, 1974; present paper), whereas the
diploid chromosome number is 34, except
for males of L. (Melaraphe) neritoides. Ob-
tained results allow us to speculate that large
differences in genome size of periwinkles and

loss or accumulation of DNA occur within
chromosomes without changing their num-
ber. As already reported for other organisms
(Hutchinson et al., 1980; Gold & Prince, 1985;
Olmo et al., 1989; Vitturi et al., 1993), ob-
served C-value variability ought to reflect pri-
marily gain or losses of repeated DNA se-
quences.

As in littorinid species, wide genome size
variations with no substantial change in chro-
mosome number have been already reported
among species of the same genus in culicid
mosquitoes (Diptera) (Nagesh Rao & Rai,
1990) and leaf beetles (Coleoptera, Chry-

130 VITTURI ET AL.

TABLE 3. Genome size evaluation in three periwinkle species. Relative fluorescence is referred to

Mytilus edulis

Fluorescence in

Arbitrary Units

Percentage Estimate
Number of Average Standard of Relative Haploid DNA Standard
Species Samples modal Value Deviation Fluorescence Content (pgs) Deviation
Littorina (Melaraphe)
punctata 3 25.333 0.471 50.667 0.811 0.0301
Littorina (Melaraphe)
neritoides 3 43 0.816 86 1.376 0.0522
Littorina saxatilis 4 42.250 0.829 84.500 1.352 0.0531

TABLE 4. Chromosome numbers in six species of the genus Littorina

Species n 2n Origin

L. brevicula 117 — Japan

L. strigta 1% 34 Japan

L. neritoides 17 34 Provence, France
L. neritoides 11% 33

L. neritoides 17 33

L. punctata We 34

L. saxatilis 17 34 Cornwall, England
L. saxatilis 17 34

L. saxatilis 17 34 Barents Sea

L. saxatilis — 34

L. obtusata — 34

Lagoon of Venice, Italy
Gulf of Palermo, Italy
Gulf of Palermo, Italy

Northern Sea, Sweden

Lagoon of Venice, Italy
Northern Sea, Sweden

Reference

Nishikawa, 1962
Nishikawa, 1962
Thiriot-Quievreux 4 Ayraud, 1982
Present paper

Vitturi et al., 1988

Present paper

Janson, 1983

Janson, 1983

Birstein & Mikhailova, 1990
Present paper

Janson, 1983

somelidae) (Petitpierre & al., 1993). Further-
more, no direct correlation between DNA
content and chromosome number were ar-
gued also for pleurocerid snail genus Semi-
sulcospira (Mesogastropoda) (Nakamura 4
Ojima, 1990).

Although karyological data for L. punctata
are limited to the diploid chromosome num-
ber (2n = 34) and spermatocyte bivalent mor-
phology, they allow us to exclude a XO sex-
determining mechanism for this species.
Hence, the wide karyological differences (ge-
nome size and presence of sex chromo-
somes) between L. punctata and L. neri-
toides make location of these species in the
same subgenus (Nordsieck, 1968; Torelli,
1982) unjustified. Alternatively, in accordance
with Rosewater (1970), the taxon Melaraphe
should contain only L. neritoides, which dif-
fers from all other littorinids by a pair of cusps
on the basal part of the central tooth in the
radula.

The three species of Littorina here studied
display a great variability when different cy-
togenetical parameters (i.e., karyotype mor-
phology, NOR patterns, sex chromosomes,
genome size) are considered, and, therefore,

karyology may be very useful in order to de-
fine the frequently rearranged taxonomy of
the genus (Bandel, 1974).

ACKNOWLEDGEMENTS

Financial support by the ministero per Г
Università e la Ricerca Scientifica e Tecno-
logica (60%, 1992-93), Roma, is gratefully
acknowledged.

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ridionalis (Neritidae) (Prosobranchia, Archaeo-
gastropoda). Cytologia, 53: 131-138.

VITTURI, R. 4 E. CATALANO, 1989, Spermatocyte
chromosomes and nucleolus organizer regions
(NORs) in Tricolia speciosa (Mühlfeld, 1824)
(Prosobranchia, Archaeogastropoda). Malacolo-
gia, 31(1): 211-216.

VITTURI, R. 4 E. CATALANO, 1990, Spermatocyte
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corneum(Prosobranchia: Neogastropoda): varia-
tion in silver-NOR pattern. Marine Biology, 104:
259-263.

VITTURI, В., С. MANSUETO & Е. FICARELLA DI
PAOLA, 1993, Heterochromatin variation in four
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VITTURI, R., Е. CATALANO, М. В. LO CONTE,
А. М. ALESSI, Е. P. AMICO 8 D. COLOMBERA,
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132 VITTURI ET AL.

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MALACOLOGIA, 1995, 37(1): 133-156

POPULATION GENETICS AND SYSTEMATIC STATUS ОЕ ONCOMELANIA
HUPENSIS (GASTROPODA: POMATIOPSIDAE) THROUGHOUT CHINA

George М. Davis', Zhang УР, Guo Yuan Hua’, & Christina Spolsky'

ABSTRACT

The coevolved relationships of populations considered to be Oncomelania hupensis through-
out China with Schistosoma japonicum are of interest to evolutionary biologists and consider-
able importance to medical science relative to understanding the differentiation of the parasite
throughout its range. As populations of Oncomelania dispersed and diversified in a direction
from Burma-western Yunnan, China, throughout China to Japan and the Philippines, the par-
asite has had to modify genetically with the genetically changing snail populations or become
regionally extinct. This hypothesis predicts that measures of genetic distance among snail
populations parallel genetic diversification among parasite populations. Our question here 15: to
what extent have populations of Oncomelania diverged genetically throughout China, and what
are the implications for systematic status of the populations?

Measures of population divergence for Oncomelania can be derived from shell characters,
anatomical characters, or molecular genetic distances. In this paper, we explore genetic di-
vergence based on allozyme data involving 14 populations that are widespread throughout
China, are in divergent drainage systems, and have different shell phenotypes. We find con-
siderable genetic differentiation occurring throughout China. We also find coherent discernable
patterns to the genetic differentiation. Careful examination of these patterns provides evidence
for one case of cross-country transport of snails, and for the existence of three exceptional
populations in which hybridizations between subspecies may have occurred. Excluding these
four populations, patterns of genetic differentiation are in general correlated with geographic
distribution and shell sculptural and shape characters. These patterns thus serve as the basis
for subdivision of O. hupensis into three discrete subspecies. O. h. robertsoni, О. В. tangi, and

O. h. hupensis.

INTRODUCTION

The rissoacean gastropod genus Oncomel-
ania is restricted to Asia and has two species
based on morphological criteria (reviewed in
Davis, 1994); Oncomelania hupensis poly-
typic species distributed from northern Burma
(fossil), western China to Japan, Philippines,
and Sulawesi, and O. minima in northwestern
Honshu, Japan. The genus is one of eight
genera of the Pomatiopsidae: Pomatiopsinae
deriving from a Gondwanian origin (Davis,
1979). The other genera are now found in
South Africa, South America, Australia, Ja-
pan, and the U.S.A. The genus in the U.S.A.,
Pomatiopsis, derived from Oncomelania of
Japan.

Oncomelania is of considerable interest to
very different groups of people. To the spe-
cialist in tropical medicine and parasitology,
O. hupensis is associated with the transmis-
sion of the human blood worm Schistosoma

japonicum. The two species of Oncomelania
also transmit different species of the lung
fluke Paragonimus (Davis et al., 1994). For
the systematist, variation in size and shell
sculpture throughout the range of the genus
has been the basis for debates about the
number of genera and species involved
(Bartsch, 1936a; Abbott, 1948; Davis, 1971;
Kang, 1981, 1985; Liu et al. 1981; Lou et al.
1982). The genus now plays a central role in
the debate of defining what a species is.
Does one today recognize polytypic species
or subspecies; to what extent should one rely
on measures of genetic distance to define
species (Woodruff et al., 1988; Davis, 1994)?

The coevolved relationships of populations
considered to be Oncomelania hupensis
throughout China with Schistosoma japoni-
cum are of interest to evolutionary biologists
and considerable importance to medical sci-
ence relative to understanding the differenti-
ation of the parasite throughout its range (re-

‘Academy of Natural Sciences of Philadelphia, 1900 Benjamin Franklin Parkway, Philadelphia PA 19103.
2Institute of Parasitic Diseases, Chinese Academy of Preventive Medicine, Shanghai, China.

134 DAVIS, ZHANG, GUO & SPOLSKY

viewed by Davis, 1992). As populations of
Oncomelania dispersed and diversified in a
direction from Burma-western Yunnan,
China, throughout China to Japan and the
Philippines (Davis 1979), the parasite has had
to modify genetically with the genetically
changing snail populations or become re-
gionally extinct. This hypothesis predicts that
measures of genetic distance among snail
populations parallel genetic diversification
among parasite populations.

Measures of population divergence for On-
comelania can be derived from shell charac-
ters, anatomical characters, or molecular ge-
netic distances. In this paper, we explore
genetic divergence based on allozyme data
involving 14 populations that are widespread
throughout China, are in divergent drainage
systems, and have different shell pheno-
types. We find considerable genetic differen-
tiation occurring throughout China. We also
find coherent discernable patterns to the ge-
netic differentiation. Careful examination of
these patterns provides evidence for one
case of cross-country transport of snails, and
for the existence of three exceptional popu-
lations in which hybridizations between sub-
species may have occurred. Excluding these
four populations, patterns of genetic differ-
entiation are in general correlated with geo-
graphic distribution and shell sculptural and
shape characters. These patterns thus serve
as the basis for subdivision of О. hupensis
into three discrete subspecies: О. h. robert-
soni, О. В. tangi, and O. В. hupensis.

MATERIALS AND METHODS
Specimens Studied

Two groups of populations were studied
seven months apart: all snails were alive
when brought to the United States by Dr.
Guo. They were collected by members of lo-
cal anti-epidemic stations throughout China
and sent to Dr. Guo in Shanghai. The 14 lo-
calities sampled are listed in Table 1 and are
shown in Figure 1. Group | represents seven
populations from the most geographically
separated locations in China collected in No-
vember 1984. Group II snails, collected in
November 1985 were from central to eastern
China, with populations concentrated in the
Hubei-Hunan-Anhui triangle.

Shell categories are based on the major
phenotypic classes of variants (Figs. 2-5): (1)
smooth and no varix = S; (2) smooth but with

pronounced varix = S,V; (3) smooth, with
varix but with sporadic low ribs on penulti-
mate and body whorl; these may appear as
exaggerated growth lines = S +; (4) ribbed
and with varix, the ribs are numerous (15-18
on the lower whorls) and pronounced but low
in profile = R; (5) pronounced high profile ribs,
few in number on the lower whorls (12-13),
with varix = R+. Each population consists of
one phenotypic class only.

Electrophoresis

Horizontal starch gel electrophoresis of
tissue proteins was followed by staining
for the following 30 loci: AAT-1, AAT-2 (as-
partate aminotransferase, 2.6.1.1); AK
(adenosine kinase, 2.7.1.20); AO (aldehyde
oxidase, 1.2.3.1); ACPH (acid phospha-
tase, 3.1.3.2); APH (alkaline phosphatase,
3.1.3.1); CK (creatine kinase, 2.7.3.2); EST-1,
EST-2, EST-3 (esterase, 3.1.1.1); GDH (glu-
tamate dehydrogenase, 1.4.1.2); G6PD-1,
G6PD-2 (glucose-6-phosphate dehydroge-
nase, 1.1.1.49); GPI (glucose-6-phosphate
isomerase); ISDH-1, ISDH-2, ISDH-3 (isoci-
trate dehydrogenase, 1.1.1.42); LDH (L-lac-
tate dehydrogenase, 1.1.1.42); MDH (malic
dehydrogenase, 1.1.1.37); ME-1, ME-2 (malic
enzyme, 1.1.1.40); MPI (mannose-6-phos-
phate isomerase, 5.3.1.9); NADD-1 (NADH
dehydrogenase, 1.6.99.3); 6PGD (phospho-
gluconate dehydrogenase, 1.1.1.44); PGM-1,
PGM-2 (phosphoglucomutase, 5.4.2.2); OCT
(octopine dehydrogenase, 1.5.1.11); SDH-1;
SDH-2 (sorbitol dehydrogenase, 1.15.1.1) ;
XDH (xanthine dehydrogenase, 1.1.1.204).
Procedures are those of Ayala et al. (1973) as
modified by Dillon and Davis (1980), Davis
(1983), Davis et al. (1981, 1988), and most
recently for Oncomelania, by Davis et al.
(1994).

Because the number of snails was limited,
and because the two groups were studied
some months apart, it was not possible to
obtain results for all loci for all populations. In
comparing the two groups of populations, in-
sufficient snails were available to fill in miss-
ing data for some loci and to be absolutely
certain of homology of scoring among some
populations. Accordingly, the results are
given for each group of populations sepa-
rately before giving the combined data set.
The combined data set for 14 populations
involved 25 loci. The following loci were not
included: AAT-2; ME-2; CK; GDH; ISDH-3.

Genetic parameters were calculated using
BIOSYS-1 (Swofford & Selander, 1981).

POPULATION GENETICS ОЕ ONCOMELANIA HUPENSIS

TABLE 1. Populations, localities, and catalog numbers for the fourteen populations of the
two groups. Localities are listed as they occur west to east. Latitudes and longitudes are
given. They are provided for the county town when the precise locality in the county is
not known. Shell sculptural types are as follows: Smooth; Smooth+ = smooth with a few
low riblets scattered on the last two whorls of some shells, some looking as strong growth
lines; Ribs+ = ribs very strong, few and high, about 11-13 on the body whorl; Ribbed =
ribs strong, low but pronounced, many, ¡.e. some 18-19 on the body whorl or penultimate
whorl.

Group | snails collected in October 1984:

e

Yunnan Province; Dali County
25744'43"N; 100°7’16”Е; ANSP 398317, A18327
shell: smooth; no varix

. Sichuan Province; XiChang City

2657'49"N; 10277'58“E; ANSP 398322; A18333
shell: smooth; no varix

. Hubei Province; JianLi County

29°49’N; 112°54’E; ANSP 398320; A18330
shell: ribbed; varix strong

. Jiangxi Province; PengZe County

29°54’N; 116°32’E; ANSP 398321; A18331
shell: ribs+; varix strong

. Anhui Province; GuiChi City

117°20.6’E; 30°30’N; CIPD 0356; ANSP 398319; A18329
shell: ribs+; varix strong

. Fujian Province; FuQing County

25°43’38”М; 119°24’00”Е; ANSP 398317; A18332
shell: smooth; varix strong and wide

. Fujian Province; XiaPu County

26°50/11”М; 120°E; ANSP 398323; A183333
shell: smooth; varix strong and wide

Group Il collected in November 1985:
1(8). Guangxi Province; GuiPing County

23°23’27”’N; 110°04’42”E; ANSP 375733
shell: smooth+; varix strong

2(9). Hubei Province; JiangLing County

30°20’57”М; 112°11’E; ANSP 375528
shell: ribbed; varix strong

3(10). Hunan Province; YueYang City

29722'52"N; 113°06’00”Е; ANSP 375732
shell: ribs+; varix strong

4(11). Hubei Province; HanYang County

30734'46"N; 114°01”06”Е; ANSP 375731
shell: ribs+; varix strong

5(12). Anhui Province; TongLing County/City

30°12’35”М; 116°05’27”Е; ANSP 375730
shell: ribs+; varix strong

6(13). Anhui Province; NingGuo County

30°22’23”М; 118°58’21”Е; ANSP 37573
shell: smooth; varix strong

7(14). Zhejiang Province; Chang Xing County

near Anji; 31°01’35’N; 119°54’29°E; ANSP 375729
shell: smooth; no varix

= DA, Fig. 1

= XI, Fig. 1

= Jl, Fig. 1

= PZ, Fig. 1

= GC, Fig. 1

= РО, Fig. 1

= XP, Fig. 1

= GP, Fig. 1

= JL, Fig. 1

= YY, Fig. 1

= НУ, Fig. 1

= TL, Fig. 1

= NG, Fig. 1

= СХ, Fig: 1

135

Hardy-Weinberg equilibrium was analyzed
for all polymorphic loci. Nei’s (1978) genetic
distance and Cavalli-Sforza & Edwards’
(1967) arc distance were calculated and
phenograms constructed using the UPGMA
method. Homology of alleles at certain loci
could not be rechecked because of lack of
specimens, and therefore Nei’s (1978) unbi-

ased minimum distance (mD) was used so as
not to inflate D due to possible error. Un-
rooted trees based on mD were also con-
structed using the FITCH program of PHYLIP
version 3.4 (Felsenstein, 1989). This phyloge-
netic analysis program does not assume
equal rates of evolution. Twenty repetitions of
FITCH were run with randomized input order

136 DAVIS, ZHANG, GUO 8 SPOLSKY

SHANXI
| HENAN LES te
| ANHUI
| |
|
| TL
| HY о
/
SICHUAN } an,
/ ÈS
HU | 1 GC y o
R
| | 2 RUE 5
a Г. PR & e
JL y) и PZ ZHEJIANG
Zi | YY Jl Nanchang E
| Changsha
A | JIANGXI
| ^_--ХР
| GUIZHOU HUNAN р <
eGuiyang
QEr Hai Е | Fuzhou
FUJIAN
= 25
Dali >-DA | N AS
Kunmin: ig | e | |
y 1 < =
GUANGXI |
YUNNAN GP G ANGDONG N
4 Guangzhou
Canton)
. + Jiang в. \
Nanning |
|
o 90 180 270 360km | 20°

105° 110°

FIG. 1. Мар of localities in southern China. West to east: DA = Dali; Х! = XiChang; GP = GuiPing; JL
JiangLing; Jl = JianLi; YY = YueYang; HY = HanYang; PZ = PengZe; GC = GuiChi; TL = TongLing; NG
NingGuo; FQ = FuQing; CX = ChangXing; XP = XiaPu.

and optimization by global branch rearrange-
ment.

RESULTS

Indices of genetic variability are given in
Table 2. Mean heterozygosity 1$ low, with
means ranging from 0.008 to 0.093 (1.1 to 1.6
alleles per locus). The percentage of poly-
morphic loci ranged from 4.0 to 28.0. The
lowest levels (4.0 to 8.0) involved smooth-
shelled populations from Fujian and Sichuan
provinces in the west, and from Zhejiang
Province in the east. The mean value for
smooth-shelled populations was 9.3 + 6.0
(4.0-20.0; N = 6). The highest levels (20.0 to
28.0) involved ribbed snails from Hubei, An-
hui, and Jiangxi provinces. The percentage of
polymorphic loci among ribbed snails was
20.6 + 4.7 (16-28; N = 7).

In the analysis of the seven populations in
group |, allele frequencies for 29 loci involving
67 alleles are given in Table 3. Nei’s D and
arc distances are given in Table 4. Invariant
loci and loci with fixed alternative alleles are

115° 120°

given in Table 5. Phenograms based on these
distances are given in Figure 6.

Group | populations represent highly diver-
gent locations in six different provinces from
Yunnan to Jiangxi provinces in the interior,
and from Fujian Province on the coast. Nei’s
D ranges from 0.107 (close geography and
shell type: Yunnan and Sichuan: smooth
shells) to 0.346 (Fujian smooth shell type vs.
Jiangxi ribbed shell type). The mean D was
0.241 + 0.066. The corresponding value for
arc distance is 0.457 + 0.059 (range of
0.314—0.538).

Results from group II are given in Tables 6
and 7 involving 60 alleles at 28 loci. Nei’s D
averaged 0.234 + 0.095 (range of 0.089 to
0.382); arc D averaged 0.440 + 0.087 (range
of 0.291 to 0.559). Corresponding pheno-
grams are given in Figure 7.

Nine loci were monomorphic in the first
group, 12 in the second (Table 5). One locus
in each group was invariant except for one
population; thus there were 19 and 14 infor-
mative loci, respectively. There were fixed
differences at eight loci in each group (Table
5). There were minor deviations from Hardy-

POPULATION GENETICS ОЕ ONCOMELANIA HUPENSIS 137

180 270 360 km |

105*

FIG. 2. Localities as in Fig. 1, but showing the distribution of shell types. Relative shell sizes are as shown.

Weinberg equilibrium in both groups of pop-
ulations (Table 8). In the first group, these
involved four populations, five of 25 polymor-
phic loci (summed over all populations), and
primarily reflected heterozygote deficiency.
In the second group, deviations occurred in-
five populations at three of 28 polymorphic
loci; the esterase-2 locus was involved in
three of the five populations. Grouping pop-
ulations from west to east along the Yangtze
River drainage, Nei's D averaged as follows:
Yunnan-Sichuan, 0.107 (group 1; Hubei-
Hunan, 0.089—0.102 (group II, М = 3); Anhui,
0.231 (group Il, N = 2).

For the combined data-set, the matrix of
Nei’s (1978) unbiased minimum distance is
given in Table 9. The UPGMA phenogram
based on these data is given in Figure 8. The
greatest Nei’s minimum distance was 0.453
(between Gui Ping and Fu Qing). In the phe-
nogram, the populations are grouped in two
major clusters. The upper cluster 1$, with
three exceptions, a smooth-shelled cluster
with shells having no varix (Dali, Chang Xing,
Xi Chang) at the top; and the Fujian Province
populations with smooth shells and low wide
varix (Fu Qing, Xia Pu) at the bottom. Three

populations seem out of place because they
have ribbed shells: Tong Ling, Jian Li, and
Gui Chi. Tong Ling has relatively low genetic
distances to the Yunnan (Dali) and Sichuan
(Xi Chang) populations and the lowest ge-
netic distance (mD = 0.092) to the Zhejiang
population (Table 9). The presence of ribbed
shells in these snails, which are genetically
closest to smooth-shelled populations, sug-
gests the possibility that this is a hybrid pop-
ulation. The same might be the case for the
Jian Li and Gui Chi populations, which also
have closer genetic affinity with the upper
section taxa of the phenogram.

The lower cluster, with an average pairwise
mD of 0.160 + 0.07, is one characterized by
populations with ribbed shells, except for the
Ning Guo (Anhui) and the Gui Ping (Guangxi)
smooth-shelled populations. These excep-
tions will be discussed later. In sum, popula-
tions clustered in Figure 8 appear to repre-
sent three genetically cohesive groups: the
Yunnan, Sichuan, and Zhejiang smooth-
shelled populations with no varix; the two
smooth-shelled Fujian populations with low
wide shells and a low wide varix; a cohesive
group of ribbed-shelled populations, which

DAVIS, ZHANG, GUO 8 SPOLSKY

Dune

FE
a

FIG. 3. Examples of shells from six localities: A. Dali, Yunnan Province = Oncomelania hupensis robertsoni;
В. XiChang, Sichuan Province = O. В. robertsoni; С. ChangXing, Zhejiang Province = O. h. robertsoni; D.
FuQing, Fujian Province = O. h. tangi; E. XiaPu, Fujian Province = O. В. tangi; Е. NingGuo, Anhui Province
= O. В. hupensis (fausti form). The shell at the top left is 6.0 mm; others are printed to the same scale.

POPULATION GENETICS OF ONCOMELANIA HUPENSIS 139

FIG. 4. Examples of shells from six localities: A. GuiPing, Guangxi Province = O. hupensis guangxiensis; B.
JianLi, Hubei Province = O. h. hupensis; C. JianLing, Hubei Province = O. h. hupensis; D. PengZe, Jiangxi
Province = O. h. hupensis; E. GuiChi, Anhui Province = O. h. hupensis; F. YueYang, Hunan Province = O.
h. hupensis. The shell at the top left is 6.69 mm long; others are printed to the same scale.

140 DAVIS, ZHANG, GUO & SPOLSKY

FIG. 5. Examples of shells from two localities: A. TongLing, Anhui Province = Oncomelania hupensis
hupensis; B. HanYang, Hubei Province = O. h. hupensis. The top left shell is 7.75 mm long; others are

printed to the same scale.

includes two exceptions that will be dis-
cussed later. In addition, there are three pos-
sible hybrid populations in the upper cluster.
The general topology of the UPGMA pheno-
gram is confirmed by FITCH analyses. Figure
9 gives the best unrooted FITCH tree (sum of
squares = 2.165); the 20 repetitions pro-
duced only four unique trees, with three clear
groupings in each case: Yunnan-Sichuan-
Zhejiang (Dali group), Fu Qing—Xia Pu (Fujian
group), and a mainly ribbed group of six pop-
ulations (Han Yang, Gui Ping, Yue Yang,
Peng Ze, Jiang Ling, and Ning Guo). The dif-
ferences in the four FITCH trees occurred
only in relative placement and branch lengths
of the three possibly hybrid populations; in all
cases, however, these three populations
were positioned between the two smooth-
shelled groups on the one hand and the
ribbed group on the other. The former two
groups are found in the upper cluster in the
UPGMA phenogram; the latter group corre-
sponds exactly to the lower cluster in the UP-
GMA tree. Thus, the differences between the
UPGMA and FITCH analyses lie mainly in the

placement of the three putative hybrid pop-
ulations.

Genetic Distance, Shell Sculpture and
Geographic Distance.

Pairwise comparisons among populations
on the basis of Nei’s minimum D from the
smallest value to 0.156, along with the shell
type, are given in Table 10. The value of
0.156 was arbitrarily chosen to represent the
upper limit of divergence that one might the-
oretically expect between populations based
on a number of studies of diverse taxa (re-
viewed in Ayala and Aquadro, 1982). It 1$
clear, even without doing a Mantel test (Rice,
1989), that there is no correlation between
geographic distance and genetic distance
when one considers all 14 populations. The
two farthest separated populations have the
lowest genetic distance (mD = 0.007 over
>2,000km), whereas closest neighbors have
an mD of 0.204 at 44 km, and of 0.339 at 72
km. This lack of correlation provided the pre-
liminary indication that the set of 14 popula-

POPULATION GENETICS OF ONCOMELANIA HUPENSIS 141

TABLE 2. Genetic variability at 25 loci for all 14 populations studied.

Mean Heterozygosity

Mean sample Mean no.

size per alleles per % loci direct- Hdywbg

Population locus locus polymorphic count expected
1. Dali (Yunnan) 30.0 : 12.0 0.028 0.035
(0.0) (0.1) (0.017) (0.021)
2. FuQing (Fujian) 25:0 р 8.0 0.008 0.013
(0.0) (0.1) (0.017) (0.010)
3. GuiChi (Anhui) 14.9 a 24.0 0.077 0.093
(0.1) (0.2) (0.036) (0.039)
4. JianLi (Hubei) oro : 20.0 0.045 0.082
(0.0) (0.2) (0.021) (0.039)
5. PengZhe (Jiangxi) 25.0 у 28.0 0.093 0.110
(0.0) (0.2) (0.036) (0.043)
6. XiChang (Sichuan) 16.2 . 4.0 0.008 0.022
(1.0) (0.0) (0.008) (0.022)
7. XiaPu (Fujian) 25.0 . 8.0 0.026 0.022
(0.0) (0.1) (0.020) (0.017)
8. ChangXing (Zhejiang) 25.0 4 4.0 0.010 0.018
(0.0) (0.0) (0.010) (0.018)
9. TongLing (Anhui) 2510 : 16.0 0.064 0.082
(0.0) (0.2) (0.032) (0.040)
10. JiangLing (Hubei) 25.0 р 20.0 0.064 0.082
(0.0) (0.2) (0.032) (0.040)
11. YueYang (Hunan) 25.0 ; 16.0 0.085 0.082
(0.0) (0.2) (0.041) (0.039)
12. GuiPing (GuangXi) 25.0 . 12.0 0.074 0.061
(0.0) .1) (0.041) (0.034)
13. NingGuo (Anhui) 25.0 ; 20.0 0.074 0.055
(0.0) 1) (0.044) (0.028)
14. HanYang (Hubei) 25.0 : 20.0 0.070 0.073
(0.0) .2) (0.035) (0.035)

tions is not a valid grouping, and suggested
the existence of discrete subgroups of O. hu-
pensis.

In Table 11 are given pairwise comparisons
between populations listed by an increasing
value of D from Tables 3 and 7. Crisscrossing
through southern China the values gradually
increase by increments of 0.020 + 0.01 (N =
15). There are no big gaps to suggest a
change from population groupings to dis-
crete species.

However, when populations are separated
into groups based on chonchological and
geographic criteria (Table 12), significant dif-
ferences are evident among the different
groups of populations. The Dali group of
three populations with smooth shells without
varix have a small average distance among
them (0.081). A relatively small distance is
found between the two Fujian populations
with smooth shells but with a low, wide varix
(0.154). Distances jump to >0.200 for inter-
group comparisons of smooth-shelled popu-
lations.

When all ribbed shells are compared, the

average distance is 0.204 + 0.085; one stan-
dard deviation ranges from 0.119 to 0.289.
However, if one excludes the three putative
hybrid populations, the remaining six popu-
lations form a genetically cohesive group:
this group includes two smooth-shelled pop-
ulations where shells have а varix. This pre-
dominantly ribbed-shelled set of populations
has a much lower average pairwise distance,
0.160. This distance is similar to the interpop-
ulational differences within the Fujian group
(0.154) and to the range of the Dali group
(0.007 to 0.127). Thus, there are no signifi-
cant within-group differences in distances
between the coherent ribbed-shelled group
of six and the two major smooth-shelled
groups, Dali and Fujian. There is a clear gap
between interpopulation distances within
groups (Dali, Fujian, and ribbed; mD < 0.160)
and intergroup distances between smooth
and ribbed populations (mD > 0.300). For the
three ribbed, possibly hybrid populations
(Tong Ling, Gui Chi, Jiang Li), comparisons of
each to the Dali group, the Fujian group, the

142 DAVIS, ZHANG, GUO 8 SPOLSKY

TABLE 3. Allele frequencies for seven populations of Oncomelania hupensis from throughout China
(group I). 29 loci; 67 alleles. “Same number of individuals from each population at all loci. There were
nine invariant loci; see Table 5. AAT-2 was not detected in XiChang, population 6.

POPULATION

Locus DALI FUQING GUICHI JIANLI PENGZE XICHANG XIAPU

(Ny 30 25 19 15 25 18 25
ААТ-1

А 1.0 0.0 0.97 1.0 0.0 0.0 1.0

В 0.0 1.0 0.0 0.0 1.0 1.0 0.0

С 0.0 0.0 0.03 0.0 0.0 0.0 0.0
ААТ-2

А 1.0 1.0 0.87 1.0 1.0 —= 0.74

В 0.0 0.0 0.13 0.0 0.0 — 0.26
ACPH

A 0.0 0.0 1.0 0.0 0.0 0.0 0.0

B 1.0 1.0 0.0 1.0 1.0 1.0 1.0
AK

A 0.0 0.0 1.0 0.0 0.0 0.0 0.0

B 1.0 1.0 0.0 1.0 1.0 1.0 1.0
APH

A 1.0 1.0 1.0 0.0 1.0 1.0 1.0

B 0.0 0.0 0.0 1.0 0.0 0.0 0.0
EST-1

A 1.0 0.86 1.0 1.0 0.96 0.0 0.0

B 0.0 0.0 0.0 0.0 0.02 0.50 0.0

C 0.0 0.0 0.0 0.0 0.0 0.50 0.0

D 0.0 0.14 0.0 0.0 0.02 0.0 1.0
EST-2

A 1.0 1.0 0.56 0.43 0.78 1.0 0.72

B 0.0 0.0 0.40 0.30 0.18 0.0 0.0

С 0.0 0.0 0.03 0.17 0.04 0.0 0.28

D 0.0 0.0 0.0 0.10 0.0 0.0 0.0
EST-3

A 1.0 0.0 0.92 120 1.0 1.0 1.0

B 0.0 1.0 0.0 0.0 0.0 0.0 0.0

C 0.0 0.0 0.08 0.0 0.0 0.0 0.0
G6PD-2

A 0.97 1.0 1.0 1.0 1.0 1.0 1.0

B 0.03 0.0 0.0 0.0 0.0 0.0 0.0
СР!

А 0.0 0.0 0.57 0.90 0.70 1.0 0.0

В 0.0 1.0 0.40 0.07 0.28 0.0 1.0

С 0.0 0.0 0.03 0.0 0.0 0.0 0.0

D 0.93 0.0 0.0 0.0 0.0 0.0 0.0

E 0.07 0.0 0.0 0.0 0.0 0.0 0.0

| 0.0 0.0 0.0 0.03 0.02 0.0 0.0
LDH

A 1.0 1.0 1.0 1.0 0.0 1.0 1.0

B 0.0 0.0 0.0 0.0 1.0 0.0 0.0
MDH

A 0.0 1.0 1.0 1.0 1.0 0.0 1.0

B 1.0 0.0 0.0 0.0 0.0 1.0 0.0
NADD

A 1.0 1.0 1.0 1.0 0.0 1.0 1.0

B 0.0 0.0 0.0 0.0 1.0 0.0 0.0

POPULATION GENETICS OF ONCOMELANIA HUPENSIS 143

TABLE 3. (Continued)

POPULATION
Locus DALI FUQING GUICHI JIANLI PENGZE XICHANG XIAPU
OCT
A 1.0 1.0 0.70 0.57 0.32 1.0 1.0
B 0.0 0.0 0.0 0.0 0.24 0.0 0.0
C 0.0 0.0 0.0 0.0 0.04 0.0 0.0
D 0.0 0.0 0.30 0.30 0.26 0.0 0.0
E 0.0 0.0 0.0 0.13 0.12 0.0 0.0
E 0.0 0.0 0.0 0.0 0.02 0.0 0.0
6PGD
A 1.0 1.0 0.0 1.0 1.0 1.0
B 0.0 150 1.0
PGM-1
A 0.72 0.0 0.07 0.0 0.04 1.0 0.0
B 0.28 1.0 0.53 0:77 0.62 0.0 1.0
С 0.0 0.0 0.37 0.20 0.34 0.0 0.0
D 0.0 0.0 0.03 0.03 0.0 0.0 0.0
PGM-2
A 1.0 0.0 1.0 1.0 0.90 1.0 0.08
В 0.0 1.0 0.0 0.0 0.02 0.0 0.93
С 0.0 0.0 0.0 0.08 0.0 0.0
SDH-1
A 0.82 0.94 1.0 0.0 0.72 1.0 1.0
B 0.18 0.06 0.0 0.90 0.0 0.0 0.0
C 0.0 0.0 0.0 0.10 0.0 0.0 0.0
D 0.0 0.0 0.0 0.0 0.28 0.0 0.0
XDH
A 1.0 1.0 1.0 1.0 0.0 1.0 1.0
В 0.0 0.0 0.0 1.0 0.0 0.0

TABLE 4. Pairwise genetic distances among the seven populations of group I; Nei’s (1978) D below the
diagonal; arc D above the diagonal. The lower set includes the AAT-2 locus, which was not scorable for
XiChang and therefore excluded from the upper set.

POPULATION DA РО GC Jl PZ XI XP
Dali — 0.443 0.456 0.402 0.538 0.339 0.392
FuQing 0.223 — 0.499 0.466 0.503 0.460 0.314
GuiChi 0.221 0.287 — 0.436 0.482 0.510 0.448
JianLi 0.171 0.255 0.217 — 0.513 0.483 0.417
PengZe 0.334 0.302 0.278 0.312 — 0.515 0.521
XiChang 0.107 0.227 0.292 0.244 0.301 — 0.455
XiaPu 0.176 0.108 0.233 0.200 0.346 0.230 —
POPULATION DA FQ GC Jl PZ XI XP
Dali — 0.395 0.411 0.364 0.481 — 0.358
FuQing 0.214 — 0.449 0.419 0.452 — 0.289
GuiChi 0.214 0.277 — 0.391 0.432 — 0.404
JianLi 0.164 0.245 0.210 — 0.456 — 0.381
PengZe 0.320 0.289 0.268 0.298 — — 0.473
XiChang — — — — = — —
XiaPu 0.173 0.107 0.227 0997 0.337 — =

ribbed group of five, and separately the Ning est to the Dali group (Dali group—Tong Ling,
Guo population, show that in each case, the 0.116; Dali group—Jiang Li, 0.189; Dali
putative hybrid population is genetically clos- group—Gui Chi, 0.227). These three excep-

144 DAVIS, ZHANG, GUO & SPOLSKY

TABLE 5. Loci fixed for a single allele or having
alternative alleles

Invariant loci Group 1
AO X

Group 2

>
A
|

>< << >< | <<

<
m
Sr EEE

ui
№

Invariant except population
G6PD-2
EST-1

Fixed for alternative alleles

| ©
>
| хх |

=
©
E
De SS | За |

CPR ee |

Alternative allele except population

3 (GC)

3 (GC) =
MDH = 8 (JL)

tional populations are also fairly closely re-
lated to each other (average mD 0.187), but
not clearly to any other group.

DISCUSSION
Population Genetics

Populations of Oncomelania studied here
are not unusual in having few alleles per lo-
cus (1.0 to 1.6), a low mean heterozygosity
(0.008 to 0.036), and low percentage of poly-
morphic loci (4 to 28). In a previous study of
Oncomelania from China and the Philippines,
Woodruff et al. (1988) found 16 polymorphic
loci among 21 loci studied, with 1.8 to 2.1
alleles per locus, in two populations from Gui

Chi in Anhui Province; mean heterozygosity
was 0.19 to 0.20, and the percentage of poly-
morphic loci ranged from 52 to 62. The major
difference between these two studies of On-
comelania 1$ that in this study we had results
from 29 loci. Had we studied only 21 loci, Gui
Chi of our study would have had 33% poly-
morphic loci rather than 24%. This, however,
is not the only source of discrepancy. Wood-
ruff et al. (1988) reported polymorphism at
three loci where we found none: LAP, ACP,
ME; in addition, they had a polymorphism at
PEP locus we did not study. Even with ex-
tensive additional screening of our Gui Chi
population (250 additional individuals; 38
loci), the number of polymorphic loci still re-
mains relatively low, 44%.

The results found in our study were similar
to those found in sister taxa of rissoacean
snails in the genus Hydrobia (Hydrobiidae):
Davis et al. (1988, 1989), one species, six
populations (30 loci); Haase (1993), three
species (25 loci); and in Truncatella: Rosen-
berg (1989), five species (19 loci), one spe-
cies with four populations. п the Truncatella
study, four populations had a mean heterozy-
gosity above 0.036 (0.037 to 0.077). How-
ever, the 0.077 value derived from a mean
sample size of 5.3 snails analyzed per locus.
The mean number of alleles per locus varied
from 1.1 to 1.4; mean heterozygosity varied
from 0.006 to 0.077; the percentage of poly-
morphic loci varied from 5.3 to 31.6. In sum-
mary, the Oncomelania populations studied
here are normally outbreeding rissoacean-
grade snails with an apparent usual pattern
of low heterozygosity and low percentage of
polymorphic loci.

Genetic distance and
taxonomic discrimination

Where do populations stop and higher taxa
begin? This topic was reviewed extensively
by Davis (1994) with particular reference to
Oncomelania hupensis. Large genetic dis-
tances by themselves do not serve to define
species. There is no magical cut off point be-
low which are populations and above which
are species. We have demonstrated the
gradual rise of Nei’s D to the 0.400 level in
pairwise comparisons of populations across
China, but again, this does not hold if one
excludes the three candidate hybrid popula-
tions. We documented the very great inter-
population variance in Nei's minimal D for the
ribbed populations traditionally classified as

POPULATION GENETICS OF ONCOMELANIA HUPENSIS 145

МЕ! '5 1978 D

CHANG XING, ZHEJIANG $
TONG LING, ANHUI

JIANG LING, HUBEI

НАМ YANG, HUBEI

GUI PING,

$
А+ У
R У
YUE YANG, HUNAN В+ У
R+ У
GUANGXI 5+ У
5 V

ARC D

NING GUO, ANHUI

А 1 1 == J

12 08 04 te)
CHANG XING, ZHEJIANG S NV
TONG LING, ANHUI R+ V
JIANG LING, HUBEI в у
YUE YANG, HUNAN А+ У
HAN YANG, HUBEI А+ У

GUI PING, GUANGXI S+ V

NING GUO, ANHUI 5 У

1 _ —E =i 1 2 НЕЕ
-60 .54 .48 .42 .36 .30 24

= 5. Е
18 12 .06 о

FIG. 6. UPGMA derived phenograms based on Nei’s 1978 D and Cavalli-Sforza's Arc D for group |

populations.

Oncomelania hupensis hupensis (range of
0.048 to 0.324).

Woodruff et al. (1988) showed that On-
comelania hupensis quadrasi populations
from the Philippines differed from Oncomel-
ania hupensis hupensis from China by a D of
0.62 + 0.04 and on this basis, invoking the
“evolutionary species concept,” stated that
two species rather than subspecies were in-
volved. On the other hand, Davis (1994)
noted that the land snail Cepaea nemoralis
introduced from Europe to the southern
U.S.A. differed by a D of 0.631 between con-
tinents (Johnson et al., 1984) and that a pop-
ulation near Pavia, Italy, differed from another
from Florence, Italy, by D = 0.391. This spe-
cies, used in various paradigms in evolution,
is well known for geographic variation in shell
polymorphisms and allozymes. Quoting
Johnson et al. (1984), “The decoupling of ge-
netic distance from speciation emphasizes
the limitations of viewing the process of spe-
ciation solely in genetic terms” (see also
Stine, 1989 and Murray et al., 1991).

The large genetic distances in our and the
Woodruff studies are caused by six or more
loci with alternative allele(s). Alternative alle-
les in allopatric populations are largely re-
sponsible for the results seized upon by
Woodruff et al. (1988) to defend an evolution-

ary or phylogenetic species concept. But is
this justified?

Species concepts and Oncomelania

In describing new species, and especially
in applying molecular genetic data in the pro-
cess, one must commit to a species concept
and be prepared to defend that concept.
Davis (1994) reviewed this topic in arguing for
Templeton's (1989) “cohesion” model,
which includes the biological species con-
cept whenever it applies as well as Patter-
son's (1985) recognition concept. The cohe-
sion model integrates population genetics
and ecology with standard studies of mor-
phology. The concept can be applied to all
organisms from outbreeders to syngameons
or parthogenetic organisms.

Among populations of Oncomelania hu-
pensis one finds considerable cohesion!
Given that each morphological character 1$
controlled by one to several genes, morpho-
logical distance is a measure of relative ge-
netic distance. Aside from size and a few
shell characters, the allopatric populations of
this species are qualitatively the same. Ac-
cordingly, the relative genetic divergence ex-
pressed in morphology 15 extremely low. Re-
productive cohesion is also large. Allopatric

146 DAVIS, ZHANG, GUO 8 SPOLSKY

TABLE 6. Allele frequencies for seven populations of Oncomelania hupensis from south-central China
(group II). 28 loci (no CK; GDH). М = 25 for all populations, all loci. There were 12 invariant loci; see
Table 5.

Population

Locus ChangXing TongLing JiangLing YueYang GuiPing NingGuo HanYang
AAT-1

A 1.0 1.0 0.0 1.0 1.0 0.0 1.0

B 0.0 0.0 1.0 0.0 0.0 0.0 0.0

C 0.0 0.0 0.0 0.0 0.0 1.0 0.0
AAT-2

A 1.0 1.0 0.92 0.96 1.0 1.0 1.0

B 0.0 0.0 0.02 0.0 0.0 0.0 0.0

C 0.0 0.0 0.06 0.0 0.0 0.0 0.0

D 0.0 0.0 0.0 0.04 0.0 0.0 0.0
ACPH

A 0.0 0.0 0.0 0.0 1.0 0.0 0.0

B 1.0 1.0 1.0 1.0 0.0 1.0 1.0
EST-1

A 1.0 1.0 0.44 1.0 1.0 1.0 1.0

B 0.0 0.0 0.44 0.0 0.0 0.0 0.0

С 0.0 0.0 0.0 0.0 0.0 0.0 0.0

D 0.0 0.0 0.12 0.0 0.0 0.0 0.0
EST-2

A 1.0 0.46 1.0 0.46 0.46 0.50 0.78

B 0.0 0.42 0.0 0.46 0.0 0.50 0.0

C 0.0 0.12 0.0 0.08 0.54 0.0 0.22
СР!

А 0.0 0.80 0.70 0.72 0.48 0.38 0.80

В 0.0 0.0 0.14 0.16 0.0 0.62 0.20

С 0.0 0.02 0.0 0.0 0.0 0.0 0.0

D 1.0 0.02 0.04 0.04 0.0 0.0 0.0

Е 0.0 0.10 0.02 0.02 0.0 0.0 0.0

E 0.0 0.02 0.04 0.02 0.0 0.0 0.0

G 0.0 0.02 0.06 0.04 0.52 0.0 0.0
ISDH-2

A 1.0 1.0 0.0 0.0 0.0 0.0 0.0

B 0.0 0.0 1.0 1.0 1.0 1.0 1.0
LDH

A 1.0 1.0 0.0 0.0 0.0 1.0 0.0

С 0.0 0.0 1.0 1.0 0.0 0.0 1.0

D 0.0 0.0 0.0 0.0 1.0 0.0 0.0
MDH

A 0.0 0.0 0.98 1.0 0.0 0.0 0.0

B 1.0 1.0 0.0 0.0 1.0 1.0 1.0

C 0.0 0.0 0.02 0.0 0.0 0.0 0.0
ME-2

A 1.0 О 1.0 0.0 0.0 1.0 1.0

В 0.0 0.0 0.0 1.0 1.0 0.0 0.0
МАРО

А 1.0 1.0 0.0 0.0 0.0 0.0 0.0

В 0.0 0.0 1.0 1.0 1.0 0.0 1.0

С 0.0 0.0 0.0 0.0 0.0 1.0 0.0
OCT

A 1.0 0.54 0.40 0.34 120 0.88 0.56

B 0.0 0.14 0.0 0.06 0.0 0.08 0.04

C 0.0 0.0 0.0 0.0 0.0 0.04 0.0

D 0.0 0.26 0.56 0.60 0.0 0.0 0.38

E 0.0 0.06 0.04 0.0 0.0 0.0 0.02

POPULATION GENETICS ОЕ ONCOMELANIA HUPENSIS 147

TABLE 6. (Continued)

Locus ChangXing TongLing JiangLing
6PGD
A 1.0 1.0 0.0
B 0.0 0.0 1.0
PGM-1
A 0.32 0.0 0.0
B 0.68 0.64 0.52
C 0.0 0.34 0.44
D 0.0 0.02 0.04
Е 0.0 0.0 0.0
РСМ-2
А 140 1.0 1.0
B 0.0 0.0 0.0
XDH
A 1.0 0.0 0.0
B 0.0 1.0 1.0

Population
YueYang GuiPing NingGuo HanYang
0.0 0.0 0.0 0.0
1.0 1.0 1.0 1.0
0.0 0.0 0.06 0.02
0.62 0.46 0.94 0.44
0.38 0.54 0.0 0.50
0.0 0.0 0.0 0.02
0.0 0.0 0.0 0.02
1.0 1.0 0.02 0.98
0.0 0.0 0.98 0.02
0.0 0.0 0.0 0.0
1.0 1.0 1.0 1.0

TABLE 7. Matrices of genetic distances for group Il populations. Nei's (1978) D is given above the

diagonal; arc D below the diagonal.

Population CX TL JL

ChangXing — 0.089 0.381
TongLing 0.306 — 0.293
JiangLing 0.559 0.492 —

YueYang 0.558 0.470 0.310
GuiPing 0.556 0.490 0.468
NingGuo 0.506 0.448 0.469
HanYang 0.487 0.397 0.302

populations from the Philippines, Japan,
China can readily interbreed and produce vi-
able F,, F, РГ, generations (reviewed in
Davis, 1980, 1981). Thus, there is no barrier
to reproduction; clearly, allozymic differenti-
ation does not impinge on overall genetic co-
hesiveness. It appears that gene divergence,
as seen in allozymes, can change relatively
rapidly in Oncomelania hupensis, while over-
all morphological and breeding system rec-
ognition patterns do not.

Because the fundamental niche parame-
ters for allopatric populations of the species
are the same (van der Schalie and Davis,
1968; Davis, 1971), the populations are de-
mographically exchangeable. Behavior is the
same for all populations, whether from Ja-
pan, the Philippines or China; that 1$ these
snails climb upward. In dry periods, they can
be found cemented to stone walls bounding
canals or, in the Yangtze flood plain, a meter
or more up on tree trunks. All populations lay
their eggs singly, each coated with fine mud.

YY GP NG HY
0.382 0.364 0.301 0.264
0.267 0.286 0.231 0175
0.102 0.261 0.263 0.093

— 0.155 0.287 0.089
0.373 — 0.296 0.140
0.483 0.506 = 0.193
0.291 0.360 0.403 —=

Species of Oncomelania

In contrast to the biological and ecologi-
cal cohesion among populations of On-
comelania hupensis, there is considerable
disruption between the two recognized spe-
cies of Oncomelania: O. hupensis and О.
minima (Bartsch, 1936a). Oncomelania min-
ima comes from northwestern Honshu, Ja-
pan. The type locality is the Noto Peninsula,
Ishikawa Prefecture. The species are readily
distinguished by differences in shell morphol-
ogy, genital and gill anatomy, and ecology
(Davis, 1971). Oncomelania minima can be
amphibious, as is O. hupensis, but the former
is abundant on sticks, leaves, rocks in a nar-
row mountain stream. Oncomelania minima
is also abundant on rock slabs over which
there was a great amount of water seepage-
flow. Thus, these snails live much as do many
species of the sister genus Tricula (Pomati-
opsidae: Triculinae); this mode of life is not
that of O. hupensis.

148 DAVIS, ZHANG, GUO 8 SPOLSKY

NEI'S 1978 D

DA Ll, YUNNAN $ NV
XI CHANG, SICHUAN $ NV
JIAN Li, HUBEI в У
FU QING, FUJIAN Ss У
XIA PU, FUJIAN SV
GUI CHI, ANHUI В+ У
РЕМС 2НЕ, JIANGXI В+ У
L 4 L 1 1 1 L 1 L L 1 1 4 1. L 1 4 J
40 36 32 .28 24 20 16 12 .08 04 o
ARC D
DA LI, YUNNAN $ NV
XI CHANG SICHUAN $ NV
FU QING, FUJIAN SAV,
XIA PU, FUJIAN SAN
GUI CHI, ANHUI R+ V
JIAN Li, HUBEI R У
РЕМС 2НЕ, JIANGXI В+ У
L N 1 1 1 L 1 + 1 1 it 1 = 1 = L L 1 J
-60 .54 .48 .42 .36 -30 .24 18 12 -06 o

FIG. 7. UPGMA derived phenograms based on Nei’s 1978 D and Arc D for group И populations.

TABLE 8. Deviations from Hardy-Weinberg (H. W.) equilibrium for all populations studied; Р =

probability; P level accepted = 0.05.

No. loci Locus not Probability
Population polymorphic in H.W. x pooled exact
GROUP | POPULATIONS (29 loci)
Dali 4 SDH-1 0.01 — 0.03
FuQing 2 — — — —
GuiChi 6 EST-3 0.0 — 0.04
Лапы 5 EST-2 0.01 0.02 0.03
OCT 0.0 0.02 0.03
PengZhe Uf SDH-1 0.002 — 0.01
XiChang 1 — — — —
XiaPu 2 — — — —
GROUP II POPULATIONS (28 loci)
ChangXing 1 PGM-1 0.02 — 0.03
TongLing 4 EST-2 0.001 0.02 0.04
JiangLing 5 — — — =
YueYang 5 AAT-2 0.0 — 0.02
GuiPing 3 EST-2 0.0 — 0.02
NingGuo 5 EST-2 0.0 — 0.0
HanYang 5 — — — —

Тре type of morphological discontinuity
and ecological divergence that allows recog-
nition of these two species parallels that of
the types of character-state changes seen
among species of the sister subfamily Tricu-
linae. Closely related triculine species differ
by a combination of character states, such as

given above and: penis with papilla in one
species, without in another; seminal vesicle a
coil dorsal to the gonad vs. a knot on the
posterior stomach; penis with pronounced
ejaculatory duct in the base of the penis in
one species vs. absent in another; outer mar-
ginal tooth with specialized outer cusp vs no

POPULATION GENETICS OF ONCOMELANIA HUPENSIS 149

TABLE 9. Matrix of genetic distances (mD) for the combined set of fourteen populations. R = ribbed; S

= smooth; V = marix; NV = no varix.

Population

Dali, Yunnan

FuQing, Fujian $, V —

GuiChi, Anhui В+, V —

JianLi, Hubei В, V —
PengZe, Jiangxi R+, V —
XiChang, Sichuan S, NV

XiaPu, Fujian S, V
ChangXing, Zhejiang S, NV
TongLing, Anhui В+, V
JiangLing, Hubei R, V

DA FQ GC Jl PZ

S,NV — 0.218 0.209 0.166 0.297 0.110 0.215 0.007 0.101 0.344 0.308 0.296 0.294 0.255
0.266 0.241 0.276 0.225 0.154 0.205 0.254 0.334 0.376 0.453 0.296 0.404
0.202 0.247 0.270 0.262 0.203 0.204 0.299 0.245 0.260 0.325 0.286
0.274 0.232 0.234 0.169 0.156 0.339 0.280 0.378 0.372 0.324
0.275 0.349 0.291 0.208 0.097 0.129 0.231 0.256 0.167

XI XP CX TL JL YY GP NG НУ

— 0.266 0.127 0.155 0.292 0.367 0.361 0.328 0.307
— 0.201 0.243 0.306 0.286 0.357 0.253 0.315
— 0.092 0.337 0.300 0.290 0.281 0.249

YueYang, Hunan В+, V — 0.150 0.222 0.048
GuiPing, Guangxi S+, V — 0.232 0.098
NingGuo, Anhui SV — 0.185
HanYang, Hubel В+, V —
NEIL'S MINIMUM DISTANCE
DA LI, YUNNAN $ NV

CHANG XING, ZHEJIANG S NV
TONG LING, ANHUI А+ У

XI CHANG, SICHUAN $ NV

JIAN LI, HUBEI в Vv
GUI CHI, ANHUI R+ V
FU QING, FUJIAN sv
XIA PU, FUJIAN зу
PENG РНЕ, JIANGXI R+ У
JIANG LING, HUBEI в V
YUE YANG, | HUNAN R+ У
HAN YANG, HUBEI R+ V
GUI PING, GUANGXI S+ V

NING GUO, ANHUI sv

.12 .08 .04 lo)

FIG. 8. UPGMA derived phenogram based on Nei's minimum D (Table 9) for both population groups

combined.

specialized outer cusp; and so forth. In sum,
species in the Pomatiopsinae and the sister
clade, Triculinae, regularly demonstrate mor-
phological discontinuities equatable to con-
siderable relative genetic divergence. Such
discontinuity is not found within the On-
comelania hupensis species complex.

The Oncomelania hupensis Polytypic
Species Complex

The data we have thus far for this species
indicates that overall cohesiveness—mor-
phological, genetic, and ecological—has not
been disrupted to the extent that allopatric
populations in this complex should be ac-

corded species status. Nevertheless, evolu-
tion towards species rank has progressed to
varying degrees in many allopatric popula-
tions both within and outside China.

In China, Oncomelania is distributed in 12
provinces and 347 counties (Kang, 1985). It
has been argued by Liu et al. (1981) that,
based on shell differences, there are five sub-
species in mainland China: O. hupensis hu-
pensis; O. hupensis robertsoni (Bartsch,
1946); O. hupensis tangi (Bartsch, 1936b); O.
hupensis fausti (Bartsch, 1925); and O. hu-
pensis guangxiensis (Liu et al., 1981). On the
basis of the data presented here, we are in-
clined to accept three of them.

Before discussing allozyme results and the

150 DAVIS, ZHANG, GUO & SPOLSKY

O. hupensis hupensis

Han Yang
x JiangLing

GuiChi

JianLi

O. h. robertsoni

O. h. tangi

HJ
0.10

FIG. 9. An unrooted FITCH tree based on Nei's mD for both population groups combined. Line lengths are

proportional to branch lengths.

basis for accepting three subspecies, it is im-
portant to understand the genetic basis for
shell characters. Historically, the plesiomor-
phic relevant character states are: small size,
smooth shell, no varix, colorless apex (Davis,

1979). Today, differences among populations
are seen in the increased shell size, sculpture
(smooth to heavily ribbed), varix or lack of
same, width/length ratios, and apex color.
Hybridization and schistosome susceptibility

POPULATION GENETICS ОЕ ONCOMELANIA HUPENSIS 191

TABLE 10. Comparison of Nei's mD (юг mD < 0.157), shell morphology, and geographic distance; $ =
smooth shells; R = ribbed shells (see Table 1 caption). Distances in km.

0.007 Dali, Yunnan x ChangXing, Zhejiang
0.048 HanYang, Hubei x YueYang, Hunan
0.059 JiangLing, Hubei x YueYang, Hunan
0.092 ChangXing, Zhejiang x TongLing, Anhui
0.092 JiangLing, Hubei x HanYang, Hubei
0.097 PengZe, Jiangxi x JiangLing, Hubei
0.098 GuiPing, Guangxi x HanYang, Hubei
0.101 Dali, Yunnan x TongLing, Anhui

0.110 Dali, Yunnan x XiChang, Sichuan

0.127 XiChang, Sichuan x ChangXing, Zhejiang
0.129 PengZe, Jiangxi x YueYang, Hunan
0.150 YueYang, Hunan x GuiPing, Guangxi
0.154 XiaPu, Fujian x FuQing, Fujian

0.155 XiChang, Sichuan х TongLing, Anhui
0.156 JianLi, Hubei x TongLing, Anhui

0.204 GuiChi x TongLing/ANHUI

0.339 JiangLing x JianLi/HUBEI

SxS 2,052 (most distant)
R+ x R+ 62.5

Rx R+ 141.1

SxR+ 199.0

Rx R+ 175.0

R+ xR 438.3

S+ x В+ 899.3

S x R+ 1,854

SxS 321.4

SXS 1,784

R+ x R+ 312.0

R+ x 5+ 660.0

эхо 142.3

S x В+ 1,567

Rx R+ 442.0

R+ x В+ 44.4 (closest neighbors)
RxR 72.1

TABLE 11. Pairwise comparisons of Nei's (1978) D values in a gradation from
small to large values along with the provinces involved. From Tables 4 and 7.

то Populations compared
0.089 TongLing x ChangXing
0.102 HanYang x YueYang
0.140 HanYang x GuiPing
0.155 GuiPing x YueYang
0.176 XiaPu x Dali

0.193 HanYang x NingGuo
0.221 GuiChi x Dali

0.255 JianLi x FuQing

0.261 GuiPing x JiangLing
0.287 GuiChi x FuQing
0.292 GuiChi x XiChang
0.301 XiChang x PengZe
0.334 PengZe x Dali

0.346 XiaPu x PengZe

0.364 GuiPing x ChangXing
0.382 YueYang x ChangXing

Geographic location

Anhui—Zhejiang Provinces
Hubei Province
Hubei—Guangxi Provinces
Guangxi—Hubei Provinces
Fujian—Yunnan Provinces
Hubei—Anhui Provinces
Anhui—Yunnan Provinces
Hubei—Fujian Provinces
Guangxi—Hubei Provinces
Anhui—Fujian Provinces
Anhui—Sichuan Provinces
Sichuan—Jiangxi Provinces
Jiangxi—Yunnan Provinces
Fujian—Jiangxi Provinces
Guangxi—Zhejiang Provinces
Hubei—Zhejiang Provinces

studies underscore the importance of these
sculptural aspects (Davis and Вий, 1973).
Ribbing is dominant to smooth, involving a
single gene with multiple alleles. Large size is
dominant to small as is higher whorl number.
Hybrid vigor was demonstrated, as F, snails
had 7.5 to 8.5 whorls, whereas parental
snails had 5.0 to 6.5 whorls. Susceptibility 1$
dominant to resistance. In addition to shell
characters, pigmentation is dominant to albi-
nism.

Ribbing 1$ restricted to the mainland of
China and especially to low-land flood plains
and marshy areas adjacent to the Yangtze
River or rivers flowing into the Yangtze. At
higher elevation, above the effects of annual

flooding, the shells of snails become smooth.
All snail populations in the highland areas
west of the three gorges barrier on the
Yangtze River, in Yunnan and Sichuan, are
smooth and without a varix. Ribbing seems
to be selected for relative to dispersal and
survival during rampaging floods of the
Yangtze River drainage.

The varix, the rib-like thickening at the lip
of the shell is, it seems, the last rib. Yet, there
are smooth shells that may or may not have a
varix. All shells with ribs have a varix. Reten-
tion of the varix in smooth shells seems to be
the genetic loss of all ribs except the last one.
It is also probable that the loss of ribbing with
retention of the varix has occurred indepen-

152 DAVIS, ZHANG, GUO 8 SPOLSKY

TABLE 12. Comparison of populations based on shell sculpture using Nei’s minimum D

|. Smooth x Smooth Shells
Dali x Sichuan x Zhejiang [no varix]
Fujian [strong, wide varix]
Dali groups x Fujian group
NingGuo [smooth, varix] х Dali group
NingGuo x Fujian group
GuiPing [smooth, varix] х Dali group
GuiPing x Fujian group

|. Ribbed x Ribbed Shells
All ribbed populations (incl. hybrids)
Ribbed populations (excluding hybrids)

0.081 + 0.06
0.154

0.230 + 0.024
0.301 + 0.024
0.274

0.316 + 0.036
0.405

CNL, EN EN CN BL,
вет ито
ND © D © O — w

0.204 + 0.085
0.160 + 0.092

ZZ
— N
er

a

(includes НУ, GP, YY, PZ, JL, NG = Ribbed “Group of 6”)

Smooth x Ribbed

Dali group x All ribbed populations
Dali group x Group of 6

Fujian group x All ribbed populations
Fujian group x Group of 6

NingGuo x rest of group of 6
GuiPing x rest of group of 6

IV. TongLing [ribbed, possible hybrid] x Other Groups

TongLing x Dali group
TongLing x Fujian group
TongLing x Group of 5
TongLing x NingGuo

V. GuiChi [ribbed, possible hybrid] x Other Groups

GuiChi x Dali group
GuiChi x Fujian group
GuiChi x Group of 5
GuiChi x NingGuo

VI.

JiangLi x Dali group
JiangLi x Fujian group
JiangLi x Group of 5
JiangLi x NingGuo

dently in different geographic regions and at
different times. Thus, one observes the fol-
lowing characters and character-states.
Varix: absent (0), present (1); Varix shape:
high and pronounced (0), low and wide (1).
The subspecies we accept and the basis
for accepting them are given below:
Oncomelania hupensis hupensis Gredler,
1881, is strongly ribbed and with a strong
rib-like varix. Shells are tall, large. This sub-
species occurs throughout the mid to lower
Yangtze River basin, especially Hubei,
Hunan, and Anhui provinces, and the Bei
River in Guangdong Province (Liu et al.,
1981). At first glance, our populations 3, 4, 5,
9, 10, 11, 12 (Table 1) appear to fit this de-
scription. Nei’s mD among these seven pop-
ulations is 0.204 + 0.085 (N = 21). As dis-

JiangLi [ribbed, possible hybrid] x Other Groups

0.257 + 0.077
0.304 + 0.056
0.296 + 0.053
0.334 + 0.089
0.227 + 0.036
0.182 + 0.042

225.726 FL, FL, FL CE
NL A UT NE Hall
Sn On SS 105)

MORO

0.116 + 0.032
0.249
0.212 + 0.048
0.217

CLE, PEPE
ou out

— O1 ND ©

0.227 + 0.034
0.264
0.267 + 0.027
0.325

A de
I mM

— O1 D ©

0.189 + 0.033
0.238
0.319 + 0.052
0.378

PATA PS PE,
шит
+ On Ww

cussed below, however, we also include two
smooth-shelled populations, 8 and 13, within
this grouping, but exclude ribbed popula-
tions 3, 5, and 12.

We included within O. hupensis hupensis
the nominal subspecies O. h. fausti and O. h.
guangxiensis. Oncomelania h. fausti has O. h.
hupensis-sized shells that are smooth but
with a strong varix. These smooth-shelled
snails live in uplands beyond the reach of
flooding of the Yangtze River and tributaries.
The two nominal taxa live over the same geo-
graphical regions. Lou et al. (1982) con-
cluded that they are synonymous. In Hubei
Province, one finds streams where the
smooth form is at the headwaters and the
ribbed form is along the flood plain. In inter-
mediate zones, there appears to be intergra-

POPULATION GENETICS ОЕ ONCOMELANIA HUPENSIS 153

dation of the two sculptural types. It is ap-
parent that ribbing is associated with flood
plains and the smooth shells associated with
upland areas beyond the effects of flooding.
There are no physical or reproductive barriers
between the sculptural types. The upland
habitats only reflect elevation and freedom
from annual flooding, not differences in the
microhabitat. The one population of the
“fausti” type that we studied, no. 13 from
Ning Guo, Anhui Province, differs from the
other members of the coherent group of six
ribbed populations (Table 12) by an mD of
0.227 + 0.062 (N = 13); from the two smooth-
shelled groups, Dali and Fujian, by 0.301 and
0.274, respectively. Further, in both phyloge-
netic analyses (Figs. 8, 9), this “fausti” type
clusters with the group of predominantly
ribbed-shelled taxa. We noted previously
that Nei’s mD for the latter populations is
0.160 + 0.07. Accordingly, we do not con-
sider O. h. fausti to be a distinct subspecies
but a smooth form of O. h. hupensis.

Liu et al. (1981) described a new subspe-
cies, O. hupensis guangxiensis from the
northwestern part of Guangxi along the Yu
Jiang and Hongshui river systems. The shells
were described as medium sized, rather thin,
smooth and with a weak varix. The type and
the description of the taxon puts emphasis
on deep sutures and especially rounded
body whorl. Our Guangxi population from Gui
Ping is at the confluence of the Yu Jiang and
Hongshui rivers. One of our specimens re-
sembles the figured type, but the remaining
somewhat resemble the “O. h. fausti’ type.
The shells are smooth and the varix weak in
90% of the snails. However, there are weak
scattered ribs or pronounced growth lines on
the shells of some individuals. It differs from
the two smooth-shelled groups by Nei’s тб
of 0.316 and 0.405 respectively; it differs
from the other members of the coherent
group of six ribbed populations by 0.182. As
with our “fausti” population, this population
clusters even more tightly with the coherent
group of O. h. hupensis. The faint ribs seen
on some shells and its genetic affinity for taxa
in the lower cluster indicates to us that it 1$
part of the ribbed-shell group, i.e. O. A. hu-
pensis, and not a distinct subspecies. How-
ever, topotypes must be studied to confirm
our opinion.

This then brings into question the affinity of
the three ribbed populations in the upper
cluster of Figure 8, Tong Ling, Jian Li, and
Gui Chi. Because morphologically and geo-

graphically, these populations appear to be
O. h. hupensis, but genetically and phyloge-
netically, they appear most closely allied to
the smooth-shelled Dali group, we raise the
possibility that these three populations are a
result of hybridization between smooth-
shelled taxa, such as O. h. robertsoni as dis-
cussed below (Dali, Chang Xing, and Xi
Chang), and a ribbed O. h. hupensis popula-
tion. It is particularly compelling to make this
hypothesis given: (1) that ribbing is dominant
to smooth and the populations in question
come from the flood plains; (2) an average
mD to the three O. h. robertsoni populations
of only 0.116 + 0.034; and (3) the close geo-
graphic proximity ofthe Tong Ling population
and the transported smooth-shelled popula-
tion from Chang Xing, Zhejiang Province; the
genetic distance between these two popula-
tions is only 0.092. In further discussion,
when we refer to ribbed populations, we
therefore include only the coherent group of
six populations that cluster together in phy-
logenetic analyses (Figs. 8, 9).

Oncomelania hupensis robertsoni (our
populations Dali, Xi Chang, Chang Xing) 1$
smooth and without varix. It is located in Si-
chuan and Yunnan provinces at altitudes
from 200 m up to 2,000 m (Liu et al., 1981) in
ditches along the slopes of hills, at the edges
of fields and irrigation canals in basin areas.
The geographic region is isolated from other
provinces with respect to invasion down the
Yangtze River by the long stretch of the
treacherous Yangtze Gorges west of Hubei
Province. One population assigned to this
subspecies is enigmatic in that it comes from
Zhejiang Province at the eastern end of China
(Group Il, Chang Xing), a locality that is the
farthest removed from Yunnan-Sichuan of all
populations studied, yet the least divergent
from the Yunnan population (Nei’s mD of
0.007). The presence of this population can-
not be due to dispersal by flotation down the
Yangtze, given the time involved to disperse
over such a great distance and the genetic
near-identity, but rather to a recent introduc-
tion either by man or birds. These three pop-
ulations cluster together tightly by both
methods of analysis (Figs. 8, 9; mD = 0.081).
They differ from the ribbed group (О. h. hu-
pensis) by an average mD of 0.305 + 0.071 (N
= 34). They also differ from the smooth-
shelled Fujian group (O. hupensis tangi) by
0.220 + 0.024. Thus, there is a significant dif-
ference in genetic distance between these
two smooth-shelled groups.

154 DAVIS, ZHANG, GUO 8 SPOLSKY

Oncomelania hupensis tangi (our Fu Qing,
Xia Pu populations) has smooth shells with a
low but very wide varix. The width of the
shells is greater relative to height than in O. h.
robertsoni. Oncomelania h. tangi lives along
the coast of Fujian Province in hilly environ-
ments (from 50 to 500 m altitude) as well as in
small ditches of the seaside lowlands. This
region is very isolated from other regions with
Oncomelania (Fig. 1). Our two populations
form a separate cluster (Figs. 8, 9); they differ
from all others by an mD of 0.285 + 0.066 (N
= 24); from the ribbed group by 0.334 + 0.089
(N = 10). Given an intragroup mD of 0.154,
there is a decided gap between this taxon
and О. h. hupensis or O. В. robertsoni.

There are no fixed diagnostic alleles unique
to any of the nominal subspecies. While ge-
netic distance per se is not a measure of sub-
species status, a subspecies should be co-
herent genetically as well as morphologically.
Phylogenetic clusters shown in Figs. 8 and 9
do provide evidence for genetic cohesive-
ness supporting the three subspecies we ac-
cept, if the arguments we provided concern-
ing “fausti,”” “guangxiensis,” and possible
hybrids hold up. Additional arguments to
support the subspecies are: (1) regional iso-
lation; (2) shell size/shape; (3) presence or
absence of shell ribbing, but recognizing that
hybridization in the flood plains will yield a
ribbed shell; (4) presence or absence of
varix; (5) varix, if present, low and wide (0),
weak (1), strong and rib-like (2); (6) differ-
ences in susceptibility to schistosomes (Liu
et al., 1981); (7) low interpopulation genetic
distances as well as low percentage of poly-
morphic loci within the O. h. robertsoni and
О. В. tangi smooth-shelled groups, as well as
within the coherent O. h. hupensis group.

Hope and McManus (1994) carried out
PCR-based RFLP analyses of variation in the
ITS-region of rDNA repeat unit among four
populations of Oncomelania hupensis т
China, three from the Philippines, and one
from Japan. Their data are puzzling because
the largest divergence they find is between
the Yunnan and Sichuan populations in
China, populations we classify as O. hupen-
sis robertsoni because of cohesiveness in al-
lozymes, shell sculpture, lack of varix, and
biogeographic closeness; and additionally,
because of lowest divergences among pop-
ulations in cytochrome b gene sequences
(Spolsky and Davis, unpubl.). The discrep-
ancy may be the result of the paucity of data
points (DNA fragment bands) in the RFLP

LE] ‘6

analyses and of the inability to distinguish
small differences in fragment size. We there-
fore advise caution in choice of tool to dem-
onstrate population divergence. All studies of
populations should give illustrations of the
shells to ascertain shell phenotypes. Voucher
specimens should be deposited for refer-
ence.

SUMMARY

The assortment of four shell characters to
distinct geographic regions and regional т-
fectivity patterns argue for subspecific sta-
tus. Low allozymic heterozygosity, unique
combinations of shell characters, coherent
genetic clustering and geographic isolation
indicate the usefulness of recognizing sub-
species for two smooth-shelled taxa. The
presence of a different type of varix on the
shells of these two nominal subspecies indi-
cates to us that they had independent ori-
gins. The phylogenetic analyses confirm this.

Major points to be considered are:

1. The three regional sets of populations
should be treated as subspecies. Allozyme
differences are insufficient by themselves,
but together with geographic location and
shell characters, enable recognition of each
of these population sets as subspecies.

2. In hilly areas above the Yangtze River
flood effects, ribbing becomes reduced and
lost. This does not affect subspecific status.

3. The disjunction between morphology
and genetic distance in three ribbed popula-
tions indicates possible hybridization be-
tween subspecies, in particular between O.
h. robertsoni and O. h. hupensis.

4. The considerable interpopulation ge-
netic distance between allopatric popula-
tions of the same shell type, especially in the
O. hupensis hupensis complex (that includes
all populations with ribs), shows genetic
change in the absence of anatomical change.
We predict that there would be parallel ge-
netic differences in the populations of Schis-
tosoma japonicum transmitted by these allo-
patric snail populations; such genetic
differentiation should be visible in DNA se-
quences of genes such as cytochrome b.
Molecular differences in the snails and their
parasites evolve at a different tempo and
mode than does the anatomical ground plan.
Would sufficient differences be found to de-
fine a series of morphostatic (term defined in
Davis, 1992) subspecies where differences

POPULATION GENETICS OF ONCOMELANIA HUPENSIS 159

might be driven by localized parasitism in a
tightly coevolved system as suggested by
Davis (1992)?

5. A weakness of this paper is that there was
not enough material to confirm the allele align-
ments on which the integrated data set in
Table 9 is based. Each data set by itself is
confirmed. What is now needed are: (1) a de-
tailed analysis of populations within a prov-
ince grouped along river drainage systems
from highlands to point of entry to the Yangtze
River or other primary river; (2) a comparison
between selected populations of the different
subspecies using more than 50 snails per
population in order to increase reliability in
estimates of polymorphic loci; (3) a careful
analysis of populations of O. hupensis hupen-
sis from selected streams where there is a
demonstrated change from ribbed shells on
the flood plains to smooth shells at higher
elevations.

Clearly, this baseline study has set the
stage for future, carefully targeted studies.

ACKNOWLEDGMENTS

We thank Caryl Hesterman for running the
gels; all gels were scored by Davis. Graphics
were prepared by Susan Trammell. We are
indebted to Dr. Margaret Mulvey and to Dr.
Hsiu-Ping Liu of the Savannah River Ecology
Laboratory for their independent critical re-
view of the manuscript; they made valuable
suggestions for improvement. This work was
supported by N.I.H. grant TMP Al 11373 to
Davis. The support of the Institute of Parasitic
Diseases, Chinese Academy of Preventive
Medicine is gratefully acknowledged.

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tematic implications (Caenogastropoda, Hydro-
biidae). Malacologia, 35: 389-398.

HOPE, M. & D. P. MCMANUS, 1994, Genetic vari-
ation in geographically isolated populations and
subspecies of Oncomelania hupensis deter-
mined by a PCR-based RFLP method. Acta
Tropica, 57: 75-82.

JOHNSON, М. S., O. С. STINE & J. MURRAY,
1984, Reproductive compatibility despite large-
scale genetic divergence in Cepaea nemoralis.
Heredity, 53: 655-665.

KANG, Z. B., 1981, A review of the taxonomical
history of Oncomelania. Academic Journal of
Hubei Medical College, 2: 54-55.

KANG, Z. B., 1985, Comments on some problems
of taxonomy of Oncomelania. Japanese Journal
of Parasitology, 34: 93-100.

LIU, У. Y., T. К. LOU, Y. Х. WANG & W. 2. ZHANG,
1981, Subspecific differentiation of oncomela-

niid snails. Acta Zootaxonomica Sinica, 6: 253-
266, 1 pl.

LOU, Т. Z., Y. Y. LIU, W. 2. ZHANG 8 У. X. WANG,
1982, A discussion on the classification of On-
comelania (Mollusca). Sinozoologia, 2: 97-114, 3
pls.

MURRAY, J., O. C. STINE & M. S. JOHNSON,
1991, The evolution of mitochondrial DNA in
Partula. Heredity, 66: 93-104.

NEI, M., 1978, Estimation of average heterozygos-
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PATTERSON, H. E. H., 1985, The recognition con-
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21-29. Pretoria, South Africa.

RICE, W. R., 1989, Analyzing tables of statistical
tests. Evolution, 43: 223-225.

ROSENBERG, G., 1989, Phylogeny and evolution
of terrestriality of the Atlantic Truncatellidae
(Prosobranchia: Gastropoda: Mollusca). Unpubl.
thesis; Harvard University.

STINE, O. C., 1989, Cepaea nemoralis from Lex-
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TEMPLETON, A. R., 1989, The meaning of species
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VAN DER SCHALIE, H. & G. M. DAVIS, 1968, Cul-
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WOODRUFF, D., К. С. STAUB, Е. $. UPATHAM, V.
VIYANT & Н. С. YUAN, 1988, Genetic variation in
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transmitting snails in China and the Philippines
are distinct species. Malacologia, 29: 347-361.

Revised Ms. accepted 1 June 1995

MALACOLOGIA, 1995, 37(1): 157

FENTER TO THe Е НОВ

RESPONSE TO BOUCHET 8 ВОСВОГ; “THE LOTTERY OF BIBLIOGRAPHICAL
DATABASES: A REPLY TO EDWARDS 8 THORNE”

М. A. Edwards! & М. J. Thorne?

While not wishing to protract this exchange
unnecessarily, we would like to offer a final,
brief, response to the above.

Which Names Get Omitted?

The analysis of names omitted 1$ based оп
material published mostly between 1940 and
1975, that is between 20 and 55 years ago.
During most of this period the Record was
compiled by volunteer specialists in the dif-
ferent animal groups. The work was done in
their spare time so that, while it is unfortu-
nate, it is not surprising that material, whether
from obscure or main stream publications,
was overlooked.

Bouchet 8 Rocroi say that **. . . in an era of
frequent and easy travel ... the staff of ZR
has attended only once ... an International
Malacological Congress . . .” and, atten-
dance at meetings would “. . . greatly en-

hance the efficiency of ZR...”.

There is a simple explanation for our ab-
sence at meetings and that is although travel
may be easy, it is expensive. We are occa-
sionally able to attend local meetings (hence
our presence at the 1986 congress in Edin-
burgh), but the cost of attending the annual
and other meetings of all the main zoological
disciplines in Europe, USA, Russia, or any-
where else, is quite beyond the capacity of
our budget. As referred to in our earlier re-
sponse, Zoological Record production is

heavily subsidized, and increased travel
would add considerably to the already large
overheads borne by BIOSIS.

It must also be said that while the meetings
we have attended have certainly been useful,
they have not led to the discovery of signifi-
cant numbers of new titles for indexing.

The Risks of a “List of Available Generic
Names in Zoology’? Based on Nomenclator
Zoologicus and ZR”

The policy of the Zoological Record and
the Nomenclator Zoologicus is to provide in-
formation and not to adjudicate on the status
of the names listed. The Nomenclator seeks
to give details of the first use of a name and
it is not practical to check back on previous
volumes to determine the availability of
names under the Code. A note of any rele-
vant information on errors, omissions, and
other matters is always welcomed by the ed-
itor for future action.

As mentioned above, The Zoological
Record is subsidized by BIOSIS and the No-
menclator Zoologicus is compiled on a vol-
untary basis. Each is produced as a service
to zoology and not as a financial proposition.

К is our understanding that proposals on
how a registry of names might be compiled
or made available have yet to be finalized,
therefore further discussion seems prema-
ture.

The editor-in-chief of Malacologia welcomes let-
ters that comment on vital issues of general im-
portance to the field of Malacology, or that com-
ment on the content of the journal. Publication is
dependent on discretion, space available and, in
some cases, review. Address letters to: Letter to
the Editor, Malacologia, care of the Department
of Malacology, Academy of Natural Sciences,
19th and the Parkway, Philadelphia, PA 19103.

The Zoological Society of London, Regent's Park, London, NW1 4RY, United Kingdom.
2BIOSIS, U.K., Garforth House, 54 Micklegate, York, North Yorkshire YO1 1LF, United Kingdom.

157

ЕАВАТА

Ibanez, M., E. Ponte-Lira & М. В. Alonso. 1995. EL GENERO
CANARIELLA HESSE, 1918, Y SU POSICION EN LA FAMILIA
HYGROMIIDAE (GASTROPODA, PULMONATA, HELICOIDEA).
MALACOLOGIA 36(1-2): 111-137.

Figures 12-20 are reprinted here that include the scale bar under

Figs. 19 and 20. The scale bar, and the edges of the shells at the
bottoms of figures 15 and 16 were cut out in the original printing.

158

ы , ' ' La 3

FIGS. 12-20. Concha y SEM detalles. (12) Canariella discobolus (Barranco de la Rajita, La Gomera). (13)
Canariella gomerae. Lectotipo de Helix (Gonostoma) дотегае (МНМ; es un ejemplar pequeño dentro de la
especie). (14-15) Canariella hispidula. (14) Lectotipo de Helix (Gonostoma) hispidula subhispidula (ZMZ).
(15) Lectotipo de Helix (Ciliella) lanosa (ZMZ). (16) Canariella leprosa (El Draguillo, Tenerife). (17-18) Cana-
riella eutropis. (17) Lectotipo de Helix eutropis (NMB). (18) Mandíbula de un ejemplar de Morro del Cava-
dero, Fuerteventura). (19-20) Rádula de Canariella planaria (Benijo, Tenerife). (19) Diente central y primeros
dientes laterales. (20) Dientes laterales próximos al margen radular. Escala: (12-17) 5 mm; (18) 200 um;
(19-20) 20 um.

Vol.
Vol.
Vol.
Vol.
Vol.
Vol.
Vol.
Vol.
Vol.
Vol.
Vol.
Vol.
Vol.

Publication dates

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30, No.
©: No:
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32, No.
32, No.
33, No.
34, No.
35, No.
35, No.
36, No.

19 Jan. 1988
28 June 1988
16 Dec. 1988
1 Aug. 1989
29 Dec. 1989
28 May 1990
30 Nov. 1990
7 June 1991
6 Sep. 1991
9 Sep. 1992
14 July 1993
2 Dec. 1993
8 Jan. 1995

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1 ne H y
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VOL. 37, NO. 1 MALAGQLOGIA Y "Nm Мой ae м
R | ГА. | ми и.
CONTENTS СИ | a a x
М. LAZARIDOU-DIMITRIADOU ; IS e

The Life Cycle, Demographic Analysis, том and Secondary Production of the г
Snail Helicella (Xerothracia) Papp! (Schútt, 1962) т ose Pulmonata) in E.
Macedonia (Greece) ...... dde LATE CARO OA q ER:

| HILARY PIGGOTT & GEORGES DUSSART И À
y 5 _ Egg-Laying . and Associated Behavioural : Responses of meta Peregra
EBEN (Müller) and Lymnaea Stagnalis (L.) to Calcium in their Environment coves Я Kr
- LUIZ RICARDO LOPES DE SIMONE N > y Ke Ne
= Anatomical Study оп Топпа Galea Mine, 1758) and Tonna Maculosa (Dillwyn, a
| 181 2 (Mesogastropoda, Tonnoidea, Tonnidae) from Brazilian Region. ASS DEP i 3
N Hoe D. ACUÑA & М. A. MUÑOZ e t, \ | a
"A Taxonomic Application of Multivariate Mixture Sie in Patellidae . igual Mua

aM __ № ELEUTHERIADIS & М. LAZARIDOU-DIMITRIADOU = и ee
was The Life Cycle, Population Dynamics, Growth and secondary Production of they

‘Snail Viviparus Contectus (Millet) (Gastropoda: Prosobranchia) i in the Marshes of |

< LU ‘the River Strymonas, Serres, Macedonia, Northern Greece Ll N Ba), 4
|. \ 4 KATERINE COSTIL & JACQUES DAGUZAN | У м > ANAL
a E Comparative Life Cycle and Growth of two rear ter Gastropod Species, N |

Hal m x ори Planorbarius Corneus: (L.) and Planorbis Planorbis (LT A ee Les au _ 53
Гм PE KENNETH C. EMBERTON Bu NS X Res ( 1 ei 13
IR WANN When Shells Do Not Tell: 145 Million Years of Evolution i in North M wit
UP ar у Ar УГ Polygyrid Land Snails, with a Revision and Conservation Priorities . + 9

Y y 3 AXHONEK к] 5 A > yd
N Sy 1 | GebatabhieDishtbytion and ‘Shell Colour and Banding Polymorphism, in Mar 7 $
! Hs ginal Populations of Cepaea Nemoralis (Gastropoda, Helicidae) RR 15 CR ane
he R. VITTURI, A. LIBERTINI, М. PANOZZO & G. MEZZAPELLE Ni a |
bs IA MC Karyotype Analysis and Genome Size in Three нала Species of
у TES % | winkles (Prosobranchia: Mesogastropoda) ia ere PR RT ` ño
A “GEORGE M. DAVIS, ZHANG YI, GUO YUAN HUA & CHRISTINA. SPOLSKY в.
N ln \ 4) 7 Population Genetics and Systematic Status of Oncomelania Hupensis
y an, N _ tropoda: HEURES) ее China AA Bd HE a Pine м.
И t | EEE о
| | AUTRE | LETTER TO THE EDITOR A
ем А. EDWARDS & M. “Y. THORNES : I
о у | Response to Bouchet & Rocroi; “The Lottery of Bibiographical Dép
RENE | ig ne) ly to to Edwards a AO rn a LANCE NOM Ad RE EN, ’ Pa, rl
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MALACOLOGIA ми ( \c a
Editor-in-Chief: Weg 3
GEORGE M. DAVIS DEP

N Editorial and Subscnption Offices: — ES E

7 Department of Malacology a ARA a)
The Academy of Natural Sciences of Philadelphia DAT

/ ' 1900 Benjamin Franklin Parkway An. Aa 8
Philadelphia, Pennsylvania 19103-1195, U.S.A. -

—

Co-Editors: | | N

“< х e ру

EUGENE СОАМ и | CAROL JONES |
California Academy of Sciences .. © | Denver, СО _ yy a
San Francisco, CA A Dips a CNRC > #4
4 И Assistant Managing Editor: ie | oh eee Eu т
CARYL НЕЗТЕВМАМ N 1
D Associate Editors: 3.) N
Е И | N ANNE GISMANN À
de University of Michigan ( Masti 1 AS
€ wos E , À 8 y Na
| Ann Arbor A Egypt Eu " в.
À HA "( п
à MALACOLOGIA is published Er the INSTITUTE OF MALACOLOGY, the Sponsor Members of Fe
which (also serving as o are: > у Be
RE OO
RUDIGER BIELER ‚ МАМ KOHN) ие ae
Field Museum, Chicago O: pra University of Washington, Seattle » и) 2
JOHN BURCH JAMES NYBAKKEN | Pag a
У 7 MELBOURNE R. CARRIKER, Moss Landing Marine Laboratory | A QE
A ¡NA
` President Elect y | ¡California AS
FRE Е, of Delaware, Lewes CLYDE F. E. ROPER \\ Là à
E GEORGE M. DAVIS Smithsonian Institution = в.
= Secretary and Treasurer Washington, D.C. = r ON
CAROLE $. HICKMAN, President | SHI-KUEI WU | Ч.
e ‚University of California, Berkeley ni University of SOS Museum, Bould
AR ¡IA | А # NER. y
Ви LS Fer \ а Members an N р: zZ
{ 230
А: © EDMUND GITTENBERGER JACKIE L. VAN GOETHEM il
tale Secretary, UNITAS MALACOLOGICA | y Treasurer, UNITAS MALACOLOGICA :
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AN 2% REN, re Ser
| ae ) A N | Copyright © 1996 by the Institute of Malacology. SA TAN je
) Rh: > д i
Ur à hol À 1 (9 dus | | | À a 5
|) ARE ds L | , | A ие
wg aX у | \ 4 $ VEN 4 м

J. А. ALLEN
Marine Biological Station
Millport, United Kingdom

E. E. BINDER
Muséum d'Histoire Naturelle
Genève, Switzerland

A. J. CAIN
University of Liverpool
United Kingdom

P. CALOW
University of Sheffield
United Kingdom

J. G. CARTER
University of North Carolina
Chapel Hill, U.S.A.

R. COWIE
Bishop Museum
Honolulu, HI., U.S.A.

А. Н. CLARKE, Jr.
Portland, Texas, U.S.A.

В. С. CLARKE
University of Nottingham
United Kingdom

R. DILLON
College of Charleston
SC, U.S.A.

С. J. DUNCAN
University of Liverpool
United Kingdom

D. J. EERNISSE
California State University
Fullerton, U.S.A.

E. GITTENBERGER

Rijksmuseum van Natuurlijke Historie

Leiden, Netherlands

F. GIUSTI
Universita di Siena, Italy

A. N. GOLIKOV
Zoological Institute
St. Petersburg, Russia

1996
EDITORIAL BOARD

$. J. GOULD
Harvard University
Cambridge, Mass., U.S.A.

A. V. GROSSU
Universitatea Bucuresti
Romania

T. HABE
Tokai University
Shimizu, Japan

R. HANLON
Marine Biomedical Institute
Galveston, Texas, U.S.A.

J. А. HENDRICKSON, уг.
Academy of Natural Sciences
Philadelphia, PA, U.S.A.

D. M. HILLIS
University of Texas
Austin, U.S.A.

K. E. HOAGLAND
Association of Systematics Collections
Washington, DC, U.S.A.

B. HUBENDICK
Naturhistoriska Museet
Goteborg, Sweden

S. HUNT
Lancashire
United Kingdom

R. JANSSEN
Forschungsinstitut Senckenberg,
Frankfurt am Main, Germany

R. N. KILBURN
Natal Museum
Pietermaritzburg, South Africa

M. A. KLAPPENBACH
Museo Nacional de Historia Natural
Montevideo, Uruguay

J. KNUDSEN
Zoologisk Institut & Museum
Kobenhavn, Denmark

A. LUCAS
Faculté des Sciences
Brest, France

C. MEIER-BROOK
Tropenmedizinisches Institut
Túbingen, Germany

Н. К. MIENIS
Hebrew University of Jerusalem
Israel

J. E. MORTON
The University
Auckland, New Zealand

J. J. MURRAY, Jr.
University of Virginia
Charlottesville, U.S.A.

R. NATARAJAN
Marine Biological Station
Porto Novo, India

J. VKLAND
University of Oslo
Norway

T. OKUTANI
University of Fisheries
Tokyo, Japan

W. L. PARAENSE

Instituto Oswaldo Cruz, Rio de Janeiro

Brazil

J. J. PARODIZ
Carnegie Museum
Pittsburgh, U.S.A.

J. P. POINTER
Ecole Pratique des Hautes Etudes
Perpignan Cedex, France

W. Е. PONDER
Australian Museum
Sydney

©] ZW
Academia Sinica
Qingdao, People’s Republic of China

D. G. REID
The Natural History Museum
London, United Kingdom

N. W. RUNHAM
University College of North Wales
Bangor, United Kingdom

S. G. SEGERSTRLE
Institute of Marine Research
Helsinki, Finland

A. STANCZYKOWSKA
Siedlce, Poland

F. STARMUHLNER
Zoologisches Institut der Universitat
Wien, Austria

Y. |. STAROBOGATOV
Zoological Institute
St. Petersburg, Russia

W. STREIFF
Université de Caen
France

J. STUARDO
Universidad de Chile
Valparaiso

S. TILLIER
Muséum National d’Histoire Naturelle
Paris, France

R. D. TURNER
Harvard University
Cambridge, Mass., U.S.A.

J.A.M. VAN DEN BIGGELAAR
University of Utrecht
The Netherlands

J. A. VAN EEDEN
Potchefstroom University
South Africa

N. H. VERDONK
Rijksuniversiteit
Utrecht, Netherlands

B. R. WILSON
Dept. Conservation and Land Management
Kallaroo, Western Australia

H. ZEISSLER
Leipzig, Germany

A. ZILCH
Forschungsinstitut Senckenberg
Frankfurt am Main, Germany

MALACOLOGIA, 1996, 37(2): 163-332

ANATOMY AND SYSTEMATICS OF THE WESTERN ATLANTIC ELLOBIIDAE
(GASTROPODA: PULMONATA)

António M. de Frias Martins

Departamento de Biologia, Universidade dos Açores, P-9502 Ponta Delgada Codex,
Sao Miguel, Açores, Portugal

ABSTRACT

Various conchological, radular and anatomical characters of the 18 Western Atlantic species
of the pulmonate family Ellobiidae are evaluated and used in a systematic review of the family.
The conchological features, especially protoconch, resorption of inner whorls, apertural den-
tition and radular morphology, are useful at the specific and generic levels. Features of the
radula of the Melampinae change greatly with increasing age. The youngest individuals have
strongly cusped crowns. Although the cusps usually disappear with age, some species retain
various features of the juvenile radula. The reproductive and central nervous systems are most
useful in defining subfamilial relationships. The monaulic, entirely glandular condition of the
pallial gonoducts, and the greater width of the visceral nerve ring are hereby considered
primitive. Morton’s (1955c) subfamilial division of the halophilic Ellobiidae is corroborated. The
Pythiinae have a monaulic, entirely glandular pallial gonoduct and a wide visceral nerve ring.
The Ellobiinae have a diaulic reproductive system with entirely glandular pallial gonoducts, and
a long visceral nerve ring. The Pedipedinae have a monaulic/incipient semidiaulic, partly glan-
dular pallial gonoduct, and a concentrated visceral nerve ring. The Melampinae are character-
ized by an advanced semidiaulic reproductive system with nonglandular pallial gonoducts, and
concentration of the ganglia of the visceral nerve ring.

The present work documents that Microtralia Dall, 1894, belongs in the Pedipedinae, not in
the Melampinae; that Laemodonta Philippi, 1846, belongs in the Pythiinae, not in the Pedipe-
dinae; that Leuconia succinea Pfeiffer, 1854, belongs in the Pedipedinae and in the new genus
Creedonia; that Apodosis Pilsbry £ McGinty, 1949, is synonymous with Leuconopsis Hutton,
1884; that Myosotella Monterosato, 1906, type species Myosotella payraudeaui “Shuttleworth”
Pfeiffer, 1856 [= Auricula myosotis Draparnaud, 1801], is removed from Ovatella Bivona, 1832,
and restored to generic rank; that Detracia Gray, 1840, as noted by Zilch (1959), is a subgenus
of Melampus Montfort, 1810; that Melampus monile (Bruguière, 1789) belongs in the subgenus
Detracia Gray, 1840; and that Detracia clarki Morrison, 1951, is a junior secondary homonym
and is herein renamed Melampus (Detracia) morrisoni. Leuconopsis manningi new species,
from Ascension Island, is described.

The phylogenetic relationships within the Ellobiidae are discussed, a tentative cladogram of
the family is presented, some distributional patterns are considered and reference is made to
the fossil record.

Key words: Archaeopulmonata, Ellobiidae, systematics, shell, radula, anatomy, genitalia,
nervous system, Western Atlantic, mangroves, salt marshes.

TABLE OF CONTENTS

Introduction
Materials and Methods
Abbreviations Used in Figures
Taxonomic Characters
Classification Outline, Western Atlantic Ellobiidae
Systematics
Family Ellobiidae H. & A. Adams in Pfeiffer,
1854
Subfamily Ellobiinae H. & A. Adams in Pfeiffer,
1854
Genus Ellobium Róding, 1798
Subgenus Auriculodes Strand, 1928

163

Ellobium (Auriculodes) dominicense
(Férussac, 1821)
Genus Blauneria Shuttleworth, 1854
Blauneria heteroclita (Montagu, 1808)
Subfamily Pythiinae Odhner, 1925
Genus Myosotella Monterosato, 1906
Myosotella myosotis (Draparnaud, 1801)
Genus Laemodonta Philippi, 1846
Laemodonta cubensis (Pfeiffer, 1854)
Subfamily Pedipedinae Fischer & Crosse, 1880
Genus Pedipes Scopoli, 1777
Pedipes mirabilis (Mühlfeld, 1816)

164 MARTINS

Pedipes ovalis C. B. Adams, 1849
Genus Сгеедота new genus
Creedonia succinea (Pfeiffer, 1854)
Genus Microtralia Dall, 1894
Microtralia occidentalis (Pfeiffer, 1854)
Genus Leuconopsis Hutton, 1884
Leuconopsis novimundi (Pilsbry &
McGinty, 1949)
Leuconopsis manningi new species
Leuconopsis sp.
Subfamily Melampinae Pfeiffer, 1853
Genus Melampus Montfort, 1810
Subgenus Melampus s.s.
Melampus (Меатриз) coffeus
(Linnaeus, 1758)
Melampus (Melampus) bidentatus Say,
1822
Subgenus Detracia Gray, 1840
Melampus (Detracia) bullaoides
(Montagu, 1808)
Melampus (Detracia) floridanus Pfeiffer,

1856

Melampus (Detracia) paranus (Morrison,
1951)

Melampus (Detracia) monile (Bruguiere,
1789)

Melampus (Detracia) morrisoni new

name

Genus Tralia Gray, 1840

Subgenus Тга/а $.5.
Tralia (Tralia) ovula (Bruguiere, 1789)

Conclusions

Phylogeny and Classification

Zoogeography of the Ellobiidae
Acknowledgments
Literature Cited
Appendix

INTRODUCTION

The Ellobiidae are primitive pulmonate
gastropods that characterize the malaco-
fauna of the upper and supra-littoral zones of
the mangroves of the tropical regions and
salt marshes of temperate regions. The Ello-
biidae were first assigned familial rank by
Lamarck (1809) when he included his Auric-
ula [= Ellobium Róding], along with three
other unrelated genera, within the “auricula-
cées”. Since then, several comprehensive
works have been published. The group was
illustrated in Reeve's Conchologia Systemat-
ica (1842) and Conchologia Iconica (1877).
A pictorial presentation was given in Martini
8 Chemnitz’ Conchylien-Cabinet by Küster
(1844) and Kobelt (1897-1901). Pfeiffer
(1854b) outlined a monograph of the Auricu-
lacea [= Ellobiidae] in his Synopsis and fully
developed the work in his Monographia in
1856, which he revised and completed

twenty years later. Odhner (1925) rearranged
the classification of the family on the basis
of radular morphology; Morton (1955c) in-
cluded morphology of the stomach and re-
productive organs in his review of the group.
Only a few genera have received compre-
hensive treatment. The genus Plecotrema
[= Laemodonta Philippi] was first revised by
H. 8 A. Adams (1853) and was studied by
Sykes (1895) and, more recently, by Huben-
dick (1956). The genera Ellobium Róding and
Melampus Montfort were studied by H. & A.
Adams (1854). Cox (1882) worked on the no-
menclature and distributuion of Pythia Ród-
ing, and Connolly (1915) did a similar study
on the genus Marinula King. Noteworthy are
the detailed anatomical and histological
studies on Melampus boholensis H. & A. Ad-
ams (Koslowsky, 1933), Myosotella myosotis
(Draparnaud) (Meyer, 1955; Morton, 1955b)
and Auriculinella (L.) bidentata (Montagu)
(Morton, 1955b). Marcus & Marcus (1965a, b)
discussed the anatomy of Melampus (M.
coffeus (Linnaeus), Melampus (D.) paranus
(Morrison), Ellobium (A.) dominicense (Férus-
sac) and Blauneria heteroclita (Montagu).
Giusti (1973) discussed the radula and anat-
omy of Ovatella firminii (Payraudeau), and the
shell, radula and anatomy of Myosotella my-
osotis (Draparnaud) were dealt with by Giusti
(1973, 1976) and Cesari (1973, 1976).

The Western Atlantic ellobiids were т-
cluded in the very earliest conchological re-
ports of American scientists. Say (1822), the
first New World malacologist, described the
common Melampus (M.) bidentatus. Gould
(1841) illustrated Say's species and Myoso-
tella myosotis (Draparnaud), which is thought
to have been introduced to North America
from Europe. Study of American ellobiids
was particularly influenced by Binney (1859,
1865) and Dall (1885). Binney (1859) figured
most of the common species; his later figures
(1865) were copied by subsequent workers
(Tryon, 1866; Dall, 1885, 1889; M. Smith,
1937; Abbott, 1974), sometimes without crit-
ical investigation. For example, Binney's in-
accurate representation of Melampus (D.) flo-
ridanus Pfeiffer in fact represents a dwarf
Melampus (M.) bidentatus Say.

Morrison (1946, 1951a, 1951b, 1954, 1958,
1959, 1964) addressed several aspects of
American ellobiid systematics, life history
and ecology, and Clench (1964) revised the
Western Atlantic Pedipes and Laemodonta.

The only detailed comprehensive anatom-
ical research on Western Atlantic ellobiids

WESTERN ATLANTIC ELLOBIIDAE 165

was that of Marcus 8 Marcus (1965a, b) on
the four species mentioned above. Several
aspects of the life history and anatomy of
Melampus (M.) bidentatus Say have been
investigated, almost exclusively in the New
England area. Hausman (1932), Holle 8
Dineen (1957) and Grandy (1972) focused on
various aspects of the ecology of this spe-
cies, while Apley (1970) and Russell-Hunter
et al. (1972) did extensive research on its
early life history. Additional investigations
have involved the morphology of the nervous
system (Price, 1977; Kahan 8 Moffett, 1979),
several aspects of physiology and behaviour
(Price, 1979, 1980; Hilbish, 1981; Capaldo,
1983), locomotion (Moffett, 1979) and feed-
ing (Thompson, 1984).

In the present work particular attention has
been paid to shell morphology, the radula
and internal anatomy, especially the repro-
ductive and nervous systems. This holistic
approach helps to clarify the systematic po-
sition and phylogenetic relationships of the
Western Atlantic ellobiids.

MATERIALS AND METHODS
Materials

Thousands of specimens from many local-
ities were studied to understand inter- and
intrapopulational variation in shell morphol-
ogy. To accomplish this | studied the collec-
tions at the Museum of Comparative Zool-
ogy, Harvard University, Cambridge, at the
American Museum of Natural History, New
York, at the Academy of Natural Sciences of
Philadelphia and at the United States Mu-
seum of Natural History, Washington, D. C.
Because museum collections were very poor
in material with preserved soft tissue, the ma-
jority of the internal anatomical work was
done on specimens from my collections.

Most of the Western Atlantic material was
obtained during field trips along the Atlantic
coast of the United States, to Bermuda, the
Bahamas, Puerto Rico and Venezuela. Some
specimens from R. C. Bullock's collection
were also kindly made available to me. Field
trips were very important in providing large
series of most recorded species and in allow-
ing examination of living animals in their hab-
itats. Most of this material is now in my col-
lection.

Material not from the Western Atlantic, be-
sides that in the museum collections men-

tioned above, included Azorean ellobiids
from my own collection, specimens from Ma-
laysia sent by A. Sasekumar and another se-
ries of specimens from Hong Kong sent by B.
S. Morton; all are now part of my collection.
The British Museum (Natural History) has
kindly allowed me to work on preserved
specimens of Marinula.

Most of the Western Atlantic species of el-
lobiids were first studied and described by
European scientists and much of the type
material is thought to be in European muse-
ums. Only the type material studied in brief
visits to the British Museum (Natural History)
and to the Museum National d’Histoire Na-
turelle de Paris, as well as that kindly sup-
plied by the Muséum d'Histoire Naturelle de
Genève, were incorporated in this work.
Sherborn (1940) and Dance (1966) have been
used to locate tentatively the collections that
might contain required type material.

Throughout the text, the museums and
collections in which the studied material 1$
deposited are indicated by the following ab-
breviations:

AMNH American Museum of Natural His-
tory, New York, NY, U.S.A.

ANSP Academy of Natural Sciences of
Philadelphia, PA, U.S.A.

BMNH The Natural History Museum [for-
merly British Museum (Natural His-
tory)], London, U.K.

R.B. Private collection of R. C. Bullock,
University of Rhode Island, King-
ston, RI, U.S.A.

FMNH Field Museum of Natural History,

Chicago, IL, U.S.A.
LSL Linnaean Society of London, U.K.
A.M. Private collection of A. M. F. Mar-
tins, University ofthe Azores, Ponta
Delgada, Sáo Miguel, Azores, POR-
TUGAL.

MCZ Museum of Comparative Zoology,
Harvard University, Cambridge,
MA, U.S.A.

MHNG Museum d'Histoire Naturelle de

Geneve, SWITZERLAND
MNHNP Museum National d’Histoire Na-
turelle de Paris, FRANCE

NHMB Natural History Museum of Basel,
SWITZERLAND

RAMM Royal Albert Memorial Museum,
London, U.K.

USNM National Museum of Natural History

[formerly United States National
Museum], Washington, DC, U.S.A.

166 MARTINS

Methods

Observation and Collection of Live Animals.
Observations of external morphology were
made in the field and in the laboratory. The
animals were photographed with Koda-
chrome film. Notes on the habitat were taken
during collecting.

Besides extensive search and collecting in
a variety of habitats, six transects were made
in January, 1981, in the mangroves of the
Florida Keys. Duplicate transects were made
in May, 1982, at two of the 1981 sites, one on
the previously disturbed site, another adja-
cent to it. All ellobiids found in the transects
were collected and preserved. Qualitative
analysis of this data is included in notes on
habitats of the different species.

Preservation. Most animals were immersed
directly in 70% ethanol. Some were relaxed
overnight in isotonic MgCl, (75.2 g Мас
of distilled water) and then preserved in 70%
ethanol. Some of the contracted and relaxed
animals were fixed in Bouin's solution after
the shell was cracked to allow better pene-
tration of the fixative; others were frozen in
fresh water for later dissection. This latter
method seemed quite useful, because the or-
gans maintained their original colors and
softness, allowing easier dissection many
months later.

Measurements. Various numbers of speci-
mens from different localities were selected
(Table 1, Appendix). Shells and dissected re-
productive systems were drawn using a Wild
M8 microscope with drawing tube. All mea-
surements were taken from these drawings
using a GTCO digitizer and IBM microcom-
puter. Radular teeth were counted from SEM
photographs.

Shell and Radula Preparations for Scanning
Electron Microscope. Juveniles of most spe-
cies and adults of the smaller species were
mounted for SEM observation of the entire
shell or the protoconch, or both. The shells
were cleaned in 95% ethanol in an ultrasonic
cleaner for two to ten seconds, depending on
the fragility of the specimen, and then were
mounted on a stub with double-sided tape.

The radulae were first cleaned in KOH (two
pellets/10 ml distilled water), washed in dis-
tilled water and in 70% ethanol. Ultrasonic
cleaning was reduced to two seconds for
each step. The radula was mounted on a
piece of cover slip, to which it adhered when
dry, and the cover slip was affixed to the stub

with double-sided tape. The use of 70% eth-
anol alone had the advantage of slower evap-
oration, which was preferable when small
pieces had to be manipulated at the exact
moment they dried, to ensure proper posi-
tioning and good adhesion. | found very help-
ful the use of human eyelashes attached to
dissecting needles with “superglue.” They
are fine, flexible, but sufficiently rigid for
clearing the membranes of the radula without
tearing, and for facilitating positioning while
mounting.

All specimens were first coated with a sin-
gle layer of carbon and then two layers of
gold-palladium (60:40) in a Denton DV-502
vacuum evaporator, and examined in an ISI
MSM-3 SEM.

Histology. Serial sections were made of
specimens of every species collected. Some
specimens were relaxed overnight in isotonic
MgCl, before fixation in Bouin’s solution. For
most of the specimens the shell was cracked
and the pieces removed to allow better fixa-
tion.

Whole animals, dissected reproductive or-
gans and stomach were embedded sepa-
rately. Embedding was done with an Auto-
technicon Duo, Model 2A. The specimens
were dehydrated in S-29 and embedded in
Paraplast. The blocks were refrigerated until
sectioned. Sectioning was done with a Spen-
cer 820 microtome. The thickness of the sec-
tions varied from 8 to 15 um. Best results
were achieved by keeping the block and the
blade refrigerated during sectioning. The
preparations were stained with Heidenhain's
aniline blue, following Luna (1968).

ABBREVIATIONS USED IN FIGURES

aa anterior aorta

acpn anterior cutaneous pedal nerve
ad anterior diverticulum

adgl anterior lobe of digestive gland
ад! albumen gland

al aperture length

aln anterior labial nerve

alpn anterolateral pedal nerve

ато! anterior mucous gland

ampn anteromedial pedal nerve

an aortic nerve

angl anal gland

ann anal nerve

aoen anterior esophageal nerve

au auricle

WESTERN ATLANTIC ELLOBIIDAE 167

anterior vas deferens
aperture width

buccal bulb

buccal commissure
buccal ganglion

bursa

bursa duct

body whorl

body whorl length

central tooth

gastric caecum

cardiac region of stomach
cerebrobuccal connective
cerebral commissure
cerebral ganglion

cutaneous-lateral pleural nerve

columellar muscle
columellar muscle nerve
cerebropedal connective
cerebropleural connective
crop

columellar tooth
columellar tooth width
penial diverticulum

eye

external pallial nerve
elbow of vagina

foot

female genital opening
fertilization pouch
genital nerve
hermaphroditic duct
hypobranchial gland
intestine

inner lip

internal pallial nerve
kidney

kidney pore

lateral teeth

lower pneumostomal gland
marginal teeth

mantle skirt

mantle skirt artery nerve
muscular band

mucous gland

mantle lappet

medial lip nerve

mantle organ

medial pallial nerve
muscular strand of stomach
mantle skirt vein

nuchal nerve
esophagus

osphradial ganglion
outer lip

ocular nerve

open spermatic groove

ovotestis

propodium

posterior artery

pedal commissure

posterior cutaneous pedal nerve
posterior cutaneous visceral nerve
posterior diverticulum
posterior lobe of digestive gland
penis

pericardium

penial nerve

penial retractor muscle

pedal ganglion

pallial gland

pharyngeal retractor muscle nerve
pharyngeal nerve

pleural ganglion

posterior lip nerve
pleuropedal connective
posterior lateral pedal nerve
pleuroparietal connective
posterior mucous gland
posteromedial pedal nerve
pneumostome

pneumostomal nerve
posterior esophageal nerve
first posterior pedal nerve
second posterior pedal nerve
prostate gland
parietocutaneous nerve
parietal ganglion

anterior left parietal ganglion
posterior left parietal ganglion
parietovisceral connective
anterior parietal tooth
posterior parietal tooth
peritentacular nerve

width of posterior parietal tooth
pulmonary vein

posterior vas deferens

pyloric region of stomach
rectum

riblets

roof of mantle cavity
stomach

salivary gland

salivary gland nerve

shoulder of body whorl

shell length

subpedal commissure
spermoviduct

spire

spire length

statocyst

statocyst nerve

seminal vesicle

shell width

168 MARTINS

18 transitional teeth

t tentacle

tem tentacular control muscle
tn tentacular nerve

upe unwrapped penis

ира! upper pneumostomal gland
V vagina

ve ventricle

vg visceral ganglion

wpe wrapped penis

TAXONOMIC CHARACTERS

Мауг (1969: 121) stated, “А taxonomic
character is any attribute of a member of a
taxon by which it differs or may differ from a
member of a different taxon.” Application of
this definition cannot be uniform and gener-
alized. Although there is consensus that a ho-
listic approach is essential to sound classifi-
cation (Mayr, 1969; Solem, 1978), one must
be aware of the difference between charac-
ters used at the species level or even for ge-
neric grouping and those used for higher taxa.
Characters that stress differences are used to
define lower taxa, whereas those characters
sensitive to convergence and seemingly less
affected by environmental factors are used to
define phylogenetic relationships among
higher taxa. For example, the pattern of the
spiral grooves on the shell is useful in sepa-
rating Melampus (M.) coffeus from Melampus
(M.) bidentatus, whereas the arrangements of
the nervous and reproductive systems are the
most consistent characters in defining the
subfamilies of the Ellobiidae. In the Ellobiidae
the shell and radular morphology are useful
mostly at the generic or specific level. Harry
(1951) and Hubendick (1978) pointed out the
value of anatomical studies for clarifying tax-
onomic relationships within the group. Morton
(19550), followed by Marcus (1965) and Mar-
cus & Marcus (1965a, b), adopted this holistic
approach by including analyses of the stom-
ach and reproductive organs; this approach
led to somewhat surprising results, such as
inclusion of Auriculinella and Blauneria within
the Ellobiinae.

In the present study conchological, radular
and anatomical characters are used. Each of
these different characters will now be con-
sidered in more detail.

FIG. 1. Conchological characters. A, Shell termi-
nology; B, Morphometry.

Conchological Characters (Fig. 1)

The shell, more than other molluscan
structures, has the obvious advantages of
permanence and ease of study. Traditionally
it has been the important basis for distinc-
tion of most taxa (Zilch, 1959). Application of
mathematical models and statistical analyses
has provided tools for the interpretation of
shell morphometry with accuracy and preci-
sion (Sokal & Sneath, 1963). Mathematical
analysis of geometry of shell coiling has also
been used (Raup, 1961, 1962, 1966, 1967;
Raup 4 Michelson 1965; Rex & Boss, 1976;
Warburton, 1979; Harasewych, 1981). This
method aims at providing an opportunity for
interpretation of evolutionary changes in shell
morphology in functional terms and as an in-
dication of strategies of adaptation to differ-
ent habitats (Vermeij, 1971). Such interpreta-
tion has been challenged recently (Gould,
1984). In spite of modern refinements in anal-
yses of conchological characters, it remains
true that some, such as shell shape, have
limited weight in assessing phylogenetic re-
lationships because shell morphology is of-
ten strongly influenced by diverse environ-
mental parameters (Hubendick, 1978; Solem,
1978).

Other shell characters, such as the proto-
conch, resorption of the inner whorls and ap-
ertural dentition, demonstrate more nearly
constant patterns and are useful at the spe-
cific and even generic level.

The gastropod protoconch indicates the
type of larval development that organisms in
the different groups have undergone (Dall,
1924; Lutz et al., 1984). It might also have
other very distinctive features that make it a

WESTERN ATLANTIC ELLOBIIDAE 169

useful taxonomic character (Walter, 1962;
Bouchet 8 Warén, 1980; Turner & Lutz,
1984). Study of the protoconch has been
aided greatly by the use of the scanning elec-
tron microscope, an increasingly important
tool in malacology (Solem, 1970; Calloway 8
Turner, 1978). The SEM has been used to
examine small and juvenile specimens, and
to study the external morphology of the shell,
radula and larvae. In this study the SEM was
used to examine the protoconchs and radu-
lae of most of the Western Atlantic ellobiids,
and to provide photomicrographs of small
and juvenile specimens.

Ellobiid protoconch morphology proved a
very useful taxonomic character in most
cases. The Melampinae, for example, have
only one type of heterostrophic protoconch,
which shows one-half of each nuclear whorl.
This feature might reflect the fact that, as far
as is known, all have free-swimming larvae.
Indeed, a similar type of protoconch occurs
in Pyramidellidae having larvae with a long
pelagic phase (Haszprunar, 1985). The mor-
phology of the protoconch in the other sub-
families of the Ellobiidae does not show an
exclusive subfamilial pattern. For example, all
the Pythiinae, Ellobiinae and the pedipedin-
ian genera Pedipes and Creedonia have a
bulbous protoconch with an umbilicus and a
laterally facing aperture. The protoconch of
the pedipedinian genera Microtralia and
Pseudomelampus sits atop the teleoconch
with the aperture facing the columellar axis,
as in the Melampinae, rather than laterally as
in all pythiinians and ellobiinians. Particular
anatomical features indicate that these two
genera belong to the Pedipedinae, however.
The protoconch is very uniform within a spe-
cies and, in the case of the West Indian Pe-
dipes, it was the only consistent diagnostic
conchological character that allowed clear
separation of species.

Shell resorption, as seen in the Ellobiidae,
also occurs in the Neritacea, Helicinidae and
Conidae. It was first noted by Montagu (1803:
235) in his Voluta denticulata [= Myosotella
myosotis (Draparnaud)] and was reported for
most members of the Ellobiidae by Gray
(1840: 220-221). Crosse & Fischer (1879,
1882), however, studied the phenomenon in
more detail and are usually credited with its
discovery. Resorption of the inner whorls pro-
vides a larger cavity in which the organs of the
visceral mass can be rearranged with spatial
economy. For example, in Melampus and Mi-
crotralia, which show a high degree of resorp-

tion, the conspicuous ovotestis has moved to
an apical position and displaced the posterior
lobe of the digestive gland; in Pedipes and
Creedonia, which do not гезо the inner
whorls, the ovotestis lies embedded in the
apical, conspicuous posterior lobe of the di-
gestive gland. This character varies within the
different subfamilies of the Ellobiidae, but can
be useful at lower taxonomic levels. In Melam-
pus s.s., for example, the partition о the inner
whorls occupies only one-fourth of the body
whorl (Figs. 225, 267), whereas in the subge-
nus Detracia it occupies at least three-fourths
of the body whorl (Figs. 302, 316). The ex-
treme case of variation within one subfamily
occurs in the Pedipedinae, in which Pedipes
and Creedonia completely retain the inner
whorls (Figs. 106, 128, 153), whereas in Mi-
crotralia resorption reaches the most ad-
vanced stage in the Ellobiidae with less than
a quarter of the interior partitions left (Fig.
178). In conjunction with other features, the
lack of shell resorption justified the creation of
the new genus Creedonia. The degree of re-
sorption also supported the separation of De-
tracia as a Subgenus of Melampus s.l. and was
helpful in the interpretation of some anatom-
ical differences observed between that sub-
genus and Melampus s.s.

Apertural dentition, an important character
in gastropod classification, is а conspicuous
feature of the Ellobiidae. One of the most
commonly accepted functions of the aper-
tural dentition is that of constituting a barrier
against predators. From my observations on
the disposition of the various branches of the
columellar muscles along the conspicuous
internal lamellae of Melampus (D.) bullaoides
(Fig. 302), | think that this feature also helps
in positioning the shell during locomotion.
Although variable, there are some general
patterns of apertural dentition. On this basis
one can characterize broadly the different
subfamilies as follows: Ellobiinae with bipli-
cate inner lip, with columellar and parietal
teeth very close together; Pythiinae with
evenly spaced triplicate inner lip with first pa-
rietal tooth strongest; Pedipedinae with two
columellar teeth and strong parietal tooth;
Melampinae with inner lip dentition restricted
to anterior half, columellar and posterior pa-
rietal teeth conspicuous, outer lip dentate.
There are exceptions to these patterns, how-
ever, forthe species in the melampinian genus
Tralia have an inner lip structure very similar to
that of the pythiinian genus Myosotella. The
inner lip dentition of Microtralia deviates from

170 MARTINS

FIG. 2. Terminology for radular teeth of Melampus
(M.) coffeus. A, Top view of central tooth; B, Top
view of first lateral tooth; C, Lateral view of first
lateral tooth; D, Top view of tenth marginal tooth;
E, Top view of 20th marginal tooth.

the pattern of the Pedipedinae in having only
one columellar tooth. The apertural structure
of the pythiinian genera Cylindrotis and Au-
riculastra resembles that of the Ellobiinae and
Ellobium (E.) aurisjudae has a conspicuous
posterior parietal tooth.

Radular characters

The molluscan radula is a valuable charac-
ter in the classification of higher taxa and is
the basis of phylogenies proposed for the
Gastropoda (Gray, 1853; Troschel, 1856-
1893; Mörch, 1867). Some authors have stud-
ied ellobiid radulae in an attempt to divide the
Ellobiidae into subfamilies. Classifications of
the ellobiids by Odhner (1925) and Thiele
(1931) were based mainly on radular morphol-
ogy, but these authors differed in their sub-
familial division. Odhner adopted six subfam-
Шез whereas Thiele recognized only three.
Observation of the radula with a light micro-
scope provides only limited information on the
intricate articulation of the different teeth with
one another (Figs. 250, 395). The SEM opened
new vistas in the study of radular morphology
and function (Solem, 1972b, 1974).

The terminology used in this study follows
that of Fretter & Graham (1962) and Ober-
holzer et al. (1970) (Fig. 2). The radula of the
ellobiids characteristically has many teeth in
each row; the central tooth usually has a
small crown. In most species the transition
from the lateral teeth, which have a strong

mesocone, to the pectinate marginal teeth is
gradual. Morphology of the radula in the
Melampinae and Ellobiinae 1$ rather uniform,
but it varies in the Pedipedinae and Pythiinae.
The radula of the Melampinae undergoes a
series of morphological changes with age.
The deeply indented crown of the lateral
teeth of very young individuals becomes the
unicuspid, triangular crown of adults. Some
species, however, seem to have a radula with
neotenic features, for example Melampus (D.)
floridanus and Melampus (D.) paranus, which,
as adults, have a conspicuous ectocone on
the lateral teeth. This structure, present in the
radula of the juveniles of some species of the
Melampinae (Figs. 243-249, 370, 371), disap-
pears with age. Marcus 8 Marcus (1963, fig.
8) observed the same morphological change
in the radula of Ellobium (A.) dominicense.
Their illustration of the radula of a very young
specimen of that species shows a striking
resemblance to the radula of an adult Blaune-
па heteroclita.

The Pedipedinae and Pythiinae display
great radular diversity with as many as three
radular types in each subfamily. Radulae of
some Pedipedinae, such as Microtralia, re-
semble that of the Melampinae, whereas in
the Pythiinae the strong mesocone on the
radular teeth of Cassidula and Pythia resem-
bles that in adult Ellobium.

The radula of the ellobiids is a much more
useful character at the generic level than at
the specific level. The minute differences in
the radulae developed by analysis of closely
related pairs such as Melampus (M.) cof-
feus—Melampus (M.) bidentatus, Pedipes
mirabilis—Pedipes ovalis and Leuconopsis
novimundi—Leuconopsis manningi failed to
provide morphological evidence useful in the
separation of these species pairs. On the
other hand, the different genera, mainly within
the subfamilies Pythiinae and Pedipedinae,
are readily distinguished on the basis of their
radular morphology.

Anatomical characters

A series of anatomical characters com-
monly used in devising classifications was
listed by Solem (1978). Because all charac-
ters do not have the same taxonomic value
weighting always must be applied. Those
characters having greater influence on the
cohesion of the group should be used in phy-
logenetic studies. Those same characters
should be the least affected by nongenetic

WESTERN ATLANTIC ELLOBIIDAE 171

factors, such as environmental and com-
petitive pressures, exemplified by habitat
and food. Thus the reproductive and the ner-
vous systems ought to be considered prime
taxonomic characters for the interpretation
of phylogenetic relationships among higher
taxa.

Stomach: Graham's comprehensive studies
(1939, 1949) of the functional morphology of
the molluscan stomach showed an evolution-
ary trend toward the disappearance of the
crystalline style and simplification of the sort-
ing areas in conjunction with the adoption of
a macrophagous carnivorous diet. He ob-
served the forward migration of the cardiac
opening, with consequent reduction of the
stomach to a blind sac into which the diges-
tive gland discharges, and the increase in the
muscularity of the mid-section to form a giz-
zard.

К is generally recognized that the basom-
matophoran stomach originated from the
prosobranch condition and it appears to me
that it evolved along two different lines. The
lower basommatophoran or archaeopulmo-
nate stomach shows a tendency toward a
forward migration of the cardiac opening.
Otina otis (Turton), a primitive marine pulmo-
nate, retains a vestigial style sac and has a
rudimentary gizzard (Morton, 1955a). In the
higher, limnic basommatophorans the stom-
ach remains open-ended, with esophageal
and intestinal openings at opposite ends, as
shown in Carriker (1946) and Morton (19550).
In this group the simplest stomach occurs in
Acroloxus, which lacks musculature, has a
well-developed caecum similar to a style sac
and a structure similar to a crystalline style
(Hubendick, 1978). Morton (1952, 1953) also
investigated the functional morphology of the
gastropod stomach and, on the basis of the
disposition of the internal ciliary patches and
of the tendency toward stronger muscularity
of the mid-section, used it as a character in
the classification of the ellobiids (Morton,
19550).

In this study only the external appearance
of the stomach was noted. Without an under-
standing of the functional morphology of the
internal parts, phylogenetic inferences and
use in classification would be unwarranted
and possibly misleading.

Reproductive System: Traditionally the mol-
luscan reproductive system has been ac-
corded special value in the understanding of
the phylogenetic relationships among higher

taxa (Duncan, 1960a, b; Visser, 1977, 1981;
Gosliner, 1981; Haszprunar, 1985, 1988; Sal-
vini-Plawén 8 Haszprunar, 1987). The impor-
tance of the reproductive system in gastro-
pod evolutionary studies is corroborated by
my studies.

A basic plan of the gastropod reproduc-
tive system consists of a posteriorly located
gonad, a middle glandular section and an
anterior duct associated with the pallial re-
gion, primitively glandular owing to its prob-
able origin from the hypobranchial gland
(Fretter, 1984). This simple tube becomes in-
creasingly complex with the appearance of
specialized evaginations and of the herma-
phroditic condition (Ghiselin, 1966). In proso-
branchs gonochorism is the rule, a condition
currently considered primitive (Haszprunar,
1988). Cases of protandry and of simulta-
neous hermaphroditism exist in archaeo-
gastropods and mesogastropods, however
(Fretter, 1946). Simroth (1907) and Huben-
dick (1945) both thought that hermaphrodit-
ism was the original condition among the gas-
tropods. Krull (1935, fide Fretter & Graham,
1962) also shared Simroth's view and, based
on the fact that the pallial oviduct of the
prosobranch hydrobiids is divided in a man-
ner similar to that of the monaulic pulmo-
nates (species with one bisexual duct), he
suggested that the hydrobiids were the most
primitive gastropods. This view has not been
accepted by later authors. In the euthy-
neurans (opisthobranchs and pulmonates),
once commonly thought to have evolved from
the archaeogastropods (Pelseneer, 1894a;
Hubendick, 1945; Morton, 1955c), but more
probably from the mesogastropods (Fretter,
1946; Boettger, 1954; Duncan, 1960a; Gos-
liner, 1981) or Apogastropoda (Salvini-Plawén
8 Haszprunar, 1987; Haszprunar, 1988), her-
maphroditism 1$ the universal condition (Ghis-
elin, 1969). Opinions also differ as to which of
the two hermaphroditic conditions appeared
first, monaulic (one bisexual duct) or diaulic
(two separate sexual ducts). Pelseneer's sug-
gestion (1894b: 19) that hermaphroditism in
mollusks arose by the secondary addition or
grafting of a male system to the female indi-
vidual has led to the view that топацу 1$ the
primitive condition (Ghiselin, 1966; Marcus 4
Marcus, 1965b; Visser, 1977, 1981; Huben-
dick, 1978). Solem (1972a, 1978) considered
diauly the primitive condition, however, and
stated that partial or total fusion of the male
and female reproductive tracts has evolved
independently in several groups. The choice

172 MARTINS

of one or another hypothesis has obvious
phylogenetic implications for the use of the
reproductive system. Visser (1981) rejected
Solem's opinion because there is no evidence
of two separate gonads with two separate
gonoducts in primitive gastropods. Visser, in
contrast to Pelseneer, stated that hermaph-
roditism in the Basommatophora, unlike that
of the Stylommatophora, was derived from a
male prosobranch. As evidence he cited the
consistency ofthe penial structure throughout
the basommatophorans (see also Hubendick,
1978).

My work has led me to support the most
commonly held view, namely that monauly
and a glandular pallial gonoduct represent
the primitive condition. The tendency toward
reduction of the glandular elements of the re-
productive system to the proximal, nonpallial
portion is hereby taken as a derived trend.
Supporting this view 1$ the presence of glan-
dular pallial gonoducts among littorinids and
the primitive opisthobranchs (Gosliner, 1981).
Existence of such glandular ducts in groups
otherwise clearly primitive (Pythiinae, Ellobii-
nae) also is taken as supportive circumstan-
cial evidence for the case.

The terminology used in this study follows
that of Duncan (1975), Visser (1977), Berry
(1977) and Tompa (1984). Histological stud-
ies were carried out to clarify some critical
features of basic morphology, such as the
extent of the mucous and prostate glands
and to establish the aulic condition, the site
of separation of male and female ducts. No
distinction was made between the different
components of the penial complex (penial
sheath, preputium and penis) and this entire
structure is herein called the penis. The de-
gree of adhesion of the anterior vas deferens
to the penis is also considered, the free con-
dition being interpreted as derived.

Central Nervous System. Use of the pulmo-
nate central nervous system as a primary tax-
onomic character has become increasingly
accepted (Bargmann 1930; Van Mol, 1967;
Bishop, 1978, Haszprunar, 1985, 1988; Sal-
vini-Plawén 8 Haszprunar, 1987). Morton
(1955c) and Regondeau et al. (1976) agreed
that within the gastropods concentration of
the ganglia is a derived character, but Morton
shared Fretter 8 Graham's concern (1949)
that sole reliance on characters of the ner-
vous system to establish phylogenetic rela-
tionships can be misleading. Haszprunar
(1985, 1988) emphasized the possibility that

concentration of ganglia and consequent eu-
thyneury could be associated with small size
in some cases.

The degree of concentration of the ganglia
of the central nervous system is considered
important because the complexity of an en-
tire system 15 generally unaffected by envi-
ronmental pressures. Any major change in
the arrangement of the ganglia probably
would mean a greater rearrangement at most
levels of anatomical organization. For this
reason the morphology of the central nervous
system is considered herein to be a taxo-
nomic character useful at higher levels of
classification.

A detailed treatment of the ellobiid central
nervous system 1$ provided for Ellobium (A.)
dominicense (Fig. 21) and Melampus (М.)
coffeus (Fig. 255). The terminology adopted
here is from several sources (Simroth 1912,
1925-1928; Bargmann, 1930; Carriker, 1946;
Brisson, 1963; Price, 1977). For most species
only the relative concentration of the ganglia
seemed important, but the nerves were
found to approximate the pattern in Melam-
pus (M.) coffeus.

CLASSIFICATION OUTLINE, WESTERN
ATLANTIC ELLOBIIDAE

Family Ellobiidae H. & A. Adams in Pfeiffer, 1854
Subfamily Ellobiinae H. & A. Adams in Pfeiffer,
1854
Genus Ellobium Róding, 1798
Subgenus Auriculodes Strand, 1928
Ellobium (A.) dominicense (Férussac,
1821)
Genus Blauneria Shuttleworth, 1854
Blauneria heteroclita (Montagu, 1808)
Subfamily Pythiinae Odhner, 1925
Genus Myosotella Monterosato, 1906
Myosotella myosotis (Draparnaud, 1801)
Genus Laemodonta Philippi, 1846
Laemodonta cubensis (Pfeiffer, 1854)
Subfamily Pedipedinae Fischer & Crosse, 1880
Genus Pedipes Scopoli, 1777
Pedipes mirabilis (Mühlfeld, 1816)
Pedipes ovalis C. B. Adams, 1849
Genus Creedonia new genus
Creedonia succinea (Pfeiffer, 1854)
Genus Microtralia Dall, 1894
Microtralia occidentalis (Pfeiffer, 1854)
Genus Leuconopsis Hutton, 1884
Leuconopsis novimundi (Pilsbry &
McGinty, 1949)
Leuconopsis manningi new species
Leuconopsis sp.
Subfamily Melampinae Stimpson, 1851

WESTERN ATLANTIC ELLOBIIDAE 173

Genus Melampus Montfort, 1810
Subgenus Melampus $.5.

Melampus (M.) coffeus (Linnaeus, 1758)
Melampus (M.) bidentatus Say, 1822
Subgenus Detracia Gray т Turton, 1840
Melampus (D.) bullaoides (Montagu,

1808)
Melampus (D.) floridanus (Pfeiffer, 1856)
Melampus (D.) paranus (Morrison, 1951)
Melampus (D.) monile (Bruguiere, 1789)
Melampus (D.) morrisoni new name
Genus Tralia Gray in Turton, 1840
Subgenus Тга/а s.s.
Tralia (T.) ovula (Bruguière, 1789)

SYSTEMATICS

Family Ellobiidae H. & A. Adams in Pfeiffer,
1854

Auriculidae Lamarck, 1809: 321 [corrected
from “Les Auriculacées” by Gray, 1840].

Auriculae Lamarck. Férussac, 1821: 32.

Auriculadae Lamarck. Gray, 1824: 107.

Auriculacea Lamarck. Blainville, 1824: 245.

Auriculaceae Lamarck. Menke, 1828: 19.

Auriculoidea Lamarck. Cristofori & Jan,
1832: 6.

Auriculidea Lamarck. Beck, 1837: 101.

Auriculata Lamarck. Sismonda, 1842: 26.

Auriculiadae Lamarck. De Kay, 1843: 57.

Auriculina Lamarck. Agassiz, 1847: 41 [cor-
rection for Auriculacea]. Non Grateloup,
1838, nec Gray, 1847a.

Carychiadae (Leach MS) Gray, 1847b: 269.

Auriculae’inae Lamarck. Strobel, 1850: 32.

Conovulidae Clark, 1850: 444.

Melampidae Stimpson, 1851: 51.

Ellobiidae H. & A. Adams in Pfeiffer, 1854b:
146 [in synonymy with Auriculacea La-
marck].

Description: Shell spiral, dextral (except in
Blauneria), oval-conic, sometimes umbili-
cate, smooth or with spiral sculpture, cov-
ered with brownish periostracum; aperture
elongate, round at base, angulate posteriorly,
with strong folds on inner lip, outer lip sharp
or weakly reflected, often dentate. Inner
whorls resorbed (except in Pedipes and
Creedonia). Protoconch heterostrophic.
Animal completely retractable into shell.
Head separated from foot by transverse
groove, into which a large mucous gland
opens. Operculum present in embryos, ab-
sent in adults. Mouth T-shaped; horny upper
jaw sometimes with folded extremities lining
lateral lips; one pair of subcylindric, contrac-

tile or subretractile tentacles; eyes sessile,
medial to base of tentacles; foot long, ante-
riorly blunt, sometimes transversely divided,
posteriorly tapered and entire or bifid; pneu-
mostome on right side, medial to anal aper-
ture.

Radula broad, elongate; teeth numerous;
central tooth equilateral; lateral teeth inequi-
lateral, becoming shorter toward outer edges
of radula, abruptly or gradually changing into
marginal teeth.

Digestive system moderately long; salivary
glands usually elongate; esophagus long,
thin walled, longitudinally grooved, opening
posteriorly into wide crop; stomach generally
tripartite with thin-walled cardiac region,
muscular medial and pyloric regions and
thin-walled, smaller posterior caecum; diges-
tive gland usually bilobed, emptying anteri-
orly at crop, posteriorly at gastric caecum.

Reproductive system hermaphroditic; ovo-
testis acinose and embedded in digestive
gland or leaf-like and covering part of stom-
ach; hermaphroditic duct with generally con-
voluted seminal vesicle; glandular complex
composed of whitish albumen gland, convo-
luted posterior mucous gland, straight ante-
rior mucous gland and prostate gland cover-
ing pallial ducts (except in Melampinae);
fertilization chamber follows posterior mu-
cous gland and gives rise to oviduct and
spermiduct, which might or might not be
completely separate for their entire length;
bursa duct and bursa arising from vagina at
variable distances from aperture; female ap-
erture medial to pneumostome, anterior to
union of mantle with neck; male aperture on
right corner of cephalic groove, under right
tentacle; a fold of skin (sperm groove) runs
from near female to male aperture, functional
only in Pythia; in all others the vas deferens
lies embedded in neck skin; it separates from
skin inward near male aperture and enters
penis at posterior end; penial complex (penis
and penial sheath) lying over buccal bulb and
cerebral ganglia.

The hypoathroid, pentaganglionate central
nervous system is of the basommatophoran
type (Bargmann, 1930; Haszprunar, 1985),
composed of 11 discrete ganglia, joined by
connectives of various lengths: paired cere-
bral, buccal, pleural, parietal and pedal gan-
glia, and an unpaired visceral ganglion.

Remarks: The Ellobiidae were first assigned
familial rank by Lamarck (1809) under the
vernacular Les Auriculacées. The group т-

174 MARTINS

cluded Lamarck's Auricula and three other
genera (Melanopsis, Melania and Limnaea)
that were assigned subsequently by other
authors to different families. Many incorrect
Latinizations of Lamarck's vernacular name
followed; Blainville's (1824) Auriculacea be-
came well established and was used in major
monographs on the family (Reeve, 1842;
Küster, 1844; Pfeiffer, 1856a, 1876; Kobelt,
1898).

The correct Latin designation, Auriculidae,
was first used by Gray (1840) and was widely
accepted until the 1920s, when the names
Ellobiidae H. & A. Adams т Pfeiffer, 1854,
and Melampidae Stimpson, 1851, replaced
Lamarck's name.

According to the International Code of
Zoological Nomenclature Art. 11 (e) the name
Auriculidae has priority because, when first
published, it was based upon the name then
valid for the contained genus Auricula La-
marck. Odhner (1925), however, preferred
the name Ellobiidae H. 4 A. Adams because
the type genus, Auricula Lamarck, 1799, is a
synonym of Ellobium Róding, 1798. The
name Ellobiidae has been in general use
since that time. Works dealing exclusively
with the family, such as those of Odhner
(1925), Morton (1955b, c), Hubendick (1956),
Clench (1964), Marcus (1965), Marcus & Mar-
cus (1965a, b), Cesari (1973, 1976), or gen-
eral ones, such as those of Thiele (1931),
Zilch (1959), Hyman (1967), Franc (1968),
Fretter (1975), Jones (1975), Runham (1975),
Berry (1977), Hubendick (1978), Solem (1978,
1985), Boss (1982) and Haszprunar (1985,
1988), and even popular books, such as
those of Morris (1973), Humphrey (1975),
Emerson & Jacobson (1976) and Rehder
(1981), are the most obvious examples of the
acceptance of the name Ellobiidae.

Recently the name Melampidae Stimpson,
1851, has appeared in some influential mal-
acological works, such as those of Keen
(1971), Abbott (1974), Rios (1975) and Kay
(1979). Morrison's reintroduction (1964) of
the name Melampidae was unfortunate in
several ways. It was an unnecessary distur-
bance of taxonomic stability, because the
name Ellobiidae had already been universally
accepted. It also required a change to a dif-
ferent type genus for the family. The appear-
ance of the term Melampidae in influential
malacological works obviously was leading
to widespread use and consequent renewed
taxonomic confusion.

Strict application of the law of priority to

family-group names would upset general use
of the name Ellobiidae. In accordance with
Art. 23 (d) of the ICZN, a petition should be
submitted to the International Commission
on Zoological Nomenclature to place the
name Ellobiidae on the Official List of Family-
Group Names in Zoology, and to place the
names Auriculidae and Melampidae on the
Official List of Rejected Names.

Credit usually is given to H. & A. Adams
(1855b) for the introduction of the name El-
lobiidae. However, Pfeiffer (1854b), who had
access to the Adams brothers’ manuscript,
referred to the to-be-proposed family name,
but continued to use the name Auriculacea.
For this reason the name Ellobiidae, which
should be credited to H. & A. Adams, must
take the date 1854, when it was first pub-
lished by Pfeiffer as a synonym.

The family Ellobiidae varies greatly in mor-
phology and anatomy, but it is nevertheless
readily identifiable as a group at the familial
level. Starobogatov (1976) exaggerated the
differences in the reproductive system and
raised the family name to ordinal status and
considered existing subfamilies separate
families. This view has not gained accep-
tance and | have concluded that the differ-
ences within the ellobiids are reconcilable
within a single family.

Odhner (1925), using radular characters,
and Zilch (1959), using shell morphology,
recognized six subfamilies, Carychiinae, Mel-
ampinae, Pedipedinae, Pythiinae, Cassiduli-
nae and Ellobiinae. Morton (1955c) and Hu-
bendick (1978) merged the Cassidulinae with
the Pythiinae on the basis of the similarities of
their reproductive systems, and assigned the
latter name to the group. My studies support
Morton’s conclusions and | have followed his
scheme of classification for the division of the
Ellobiidae into subfamilies.

Zilch (1959) recognized 20 genera of living
halophilic ellobiids. Zilch’s classification is
accepted here with certain modifications and
21 genera are recognized in this paper. Zilch
considered Sarnia H. & A. Adams a subgenus
of Tralia Gray, but Marincovich (1973), on the
basis of radular morphology, placed it in the
Ellobiinae. Sarnia (Fig. 181) shows strong
conchological similarity to Pseudomelampus
and Microtralia, and for that reason | include
it in the Pedipedinae. Further information on
the reproductive and nervous systems is
needed to confirm the systematic position of
this genus, however. | have synonymized
herein Apodosis Pilsbry & McGinty with Leu-

WESTERN ATLANTIC ELLOBIIDAE 175

ELLOBIINAE CARYCHIINAE PEDIPEDINAE

0002000 a A, Lan

à À

AMIA

FIG. 3. Pictorial review of subfamilies of Ellobiidae, from most primitive to most advanced. A, Pythia (P.)
scarabaeus (Linnaeus), radula; B, Pythia (P.) scarabaeus, reproductive system; C, Pythia (P.) plicata (Fér-
ussac), central nervous system; D, Pythia (P.) scarabaeus; E, Рута (Trigonopythia) trigona (Troschel); Е,
Ophicardelus australis (Quoy & Gaimard); G, Myosotella myosotis (Draparnaud); H, Ovatella firminii Pay-
raudeau; |, Laemodonta octanfracta (Jonas); J, Allochroa bronni (Philippi); К, Cassidula (C.) aurisfelis (Bru-
guière); L, Cassidula (Cassidulta) doliolum (Petit); M, Cylindrotis quadrasi Móllendorff; М, Auriculastra subula
(Quoy & Gaimard); O, Ellobium (E.) aurismidae (Linnaeus), radula; P, Ellobium (E.) aurismidae, reproductive
system; Q, Ellobium (E.) aurismidae, central nervous system; R, Ellobium (E.) aurismidae; S, Ellobium
(Auriculodes) gangeticum (Pfeiffer); T, Auriculinella (Leucophytia) bidentata (Montagu); U, Blauneria hetero-
clita (Montagu); V, Carychium tridentatum (Risso), radula; W, Carychium tridentatum, reproductive system;
X, Carychium tridentatum, central nervous system; Y, Carychium minimum Müller; 2, Zospeum spelaeum
(Rossmássler); AA, Pedipes mirabilis (Mühlfeld), radula; BB, Pedipes pedipes (Bruguiere), reproductive
system; CC, Pedipes pedipes, central nervous system; DD, Pedipes pedipes; EE, Marinula pepita King; FF,
Creedonia succinea (Pfeiffer); GG, Pseudomelampus exiguus (Lowe); HH, Sarnia frumentum (Petit); Il,
Microtralia occidentalis (Pfeiffer); JJ, Leuconopsis obsoleta (Hutton); KK, Melampus (M.) coffeus (Linnaeus),
radula; LL, Melampus (M.) coffeus, reproductive system; MM, Melampus (M.) coffeus, central nervous
system; NN, Melampus (M.) coffeus; 00, Melampus (Micromelampus) nucleolus Martens; PP, Melampus
(Detracia) bullaoides (Montagu); QQ, Melampus (Signia) granifer (Mousson); ВВ, Tralia (T.) ovula (Bruguière);
SS, Tralia (Persa) costata (Quoy & Gaimara).

conopsis Hutton, and have created the genus
Creedonia. Myosotella Monterosato, treated
by Zilch as a subgenus of Ovatella Bivona, is
given herein generic status.

A pictorial review of the subfamilies 1$ pre-
sented in Figure 3. The radula, nervous sys-
tem and reproductive system of the type or of
a representative species of each subfamily
are shown, as well as the shells of all the type
species of the genera and subgenera herein
recognized. The reproductive and nervous
systems provided the most consistent basis
for the separation of the subfamilies.

Detailed descriptions of general anatomy
and of the nervous system are provided un-
der Melampus (M.) coffeus; these descrip-
tions will be used as standards of compari-
son in discussions of other species.

Habitat: Ellobiids are mainly tropical. They
commonly occur around the high-tide mark in
mangrove areas, under rocks or pieces of de-
caying wood. In extratropical regions they live
in eurhyaline environments of salt marshes or
in upper littoral rocky areas.

Morton (1955c) divided the ellobiids into

176 MARTINS

four groups according to habitat. The upper-
tidal marine ellobiids, such as Melampus,
Myosotella, Ophicardelus, Cassidula and El-
lobium, prefer the high-tidal fringe of man-
groves and marshes, never venturing far from
the reach of the highest spring tides. The in-
tertidal and crevice-dwelling species include
the minute ellobiids of the genera Auricu-
linella, Leuconopsis, Pedipes, Microtralia and
Marinula, to which Creedonia, Laemodonta
and Blauneria might be added, which live
buried at different depths in the sediment or
under partly buried rocks, roots and fallen
branches in the upper intertidal zone. Pythia
is the only coastal terrestrial ellobiid; it always
frequents moist places near the shore, al-
though out of reach of the highest tides. The
inland terrestrial ellobiids are Carychium and
Zospeum, which live in very humid environ-
ments, frequently under forest leaf litter or in
caves.

The ellobiids are commonly thought to have
evolved from an estuarine ancestor. Con-
quest of the terrestrial habitat brought about
modifications in the structure of the larval
stages. Such modifications, however, are not
exclusively related to distance from the sea
and a single feature, such as suppression of
a free-swimming veliger, can exist in intertidal
and terrestrial species. A more or less mod-
ified veliger stage 1$ present throughout the
family. The Melampinae have a free-swim-
ming veliger larva. Apley (1970) recorded two
to six weeks of planktonic life for Melampus
(M.) bidentatus, while Marcus & Marcus
(1965a) suggested equally long periods for
the veligers of Melampus (M.) coffeus and
Melampus (D.) paranus. The veliger stage of
the other subfamilies passes inside the egg
and the embryo crawls immediately after
hatching. Larvae of Ellobium (A.) dominicense
have а ciliate velum and can swim for very
short periods of time (Ewald, 1963). That
same ciliated structure was found in Blauneria
heteroclita by Marcus & Marcus (1963). Mor-
ton (1955c) observed that the velum of the
larvae of Myosotella myosotis and of Auricu-
linella (L.) bidentata is reduced and lacks cilia.

Another feature of larval ellobiids is the
widespread presence of an operculum, which
is lost at an early age. Blauneria heteroclita,
which normally reaches 7 mm in length, sheds
the operculum at a shell length of about 0.7
mm (Marcus & Marcus, 1963). The reduced
operculum of Myosotella myosotis and of Au-
riculinella (L.) bidentata helps to break the
shell during hatching (Morton 1955b).

Range: The family Ellobiidae has worldwide
distribution, but appears to have three main
centers, a large Indo-Pacific center, charac-
terized by Ellobium, Cassidula and Pythia; a
smaller West Indian center, characterized by
Melampus; and a much poorer Mediterra-
nean region, characterized by Myosotella and
Ovatella.

The fossil record of the Ellobiidae 1$ rela-
tively poor and 1$ insufficient for the determi-
nation of evolutionary lineages. The presence
of the Indo-Pacific genera Ellobium and Cas-
sidula in Europe during the Eocene and
Miocene (Zilch, 1959) suggests that the Eu-
ropean shores were connected with the Indo-
Pacific region. This is consistent with the ex-
istence of the Tethys Sea which, in various
ways, extended longitudinally from Australia
through Europe and northern Africa to the
tropical West Indies and eastern Pacific.
Existence of this seaway is indicated by the
distribution of several groups of inverte-
brates, and has been more extensively stud-
ied for the Mediterranean region. Evidence
of a Tethyan distribution in American faunas
was found in Foraminifera (C.G. Adams,
1967), Ostracoda (McKenzie, 1967) and in bi-
valves and gastropods (Palmer, 1967). The
ellobiid fossil record does not provide any
new information about Tethyan relationships
between Europe and America. The ellobiid
fossils of North America are represented by
the melampinine genera Rhytophorus and
Melampoides from the Cretaceous of Wyo-
ming (White, 1895; Henderson, 1935) and by
the more recent Melampus, Marinula, Tralia
and Pedipes, from the Eocene, Miocene and
Pleistocene (Conrad, 1862; Dall, 1912; Wood-
ring, 1928; Gibson-Smith & Gibson-Smith,
1979, 1982, 1985). The Mesozoic genera
seem not to have European counterparts,
but Rhytophorus was recorded from the
Lower Cretaceous of China (Zhu, 1980).
Present records are too sparse to allow elab-
oration of the meaning of such an occur-
rence. The Cenozoic genera represent only
the Recent ellobiid fauna of the West Indian
region.

Subfamily Ellobiinae H. & A. Adams т Pfeif-
fer, 1854

Auriculea Pfeiffer, 1853a: 9.

Ellobiinae “Н. & A. Adams” Pfeiffer, 1854b:
146.

Auriculinae H. & A. Adams, 1855a: 30 [emen-
dation of Auriculea Pfeiffer, 1853].

WESTERN ATLANTIC ELLOBIIDAE 177

Description: Shell very small and thin (Auri-
culinella, Blauneria) to large and thick (Ello-
bium), dextral except in Blauneria. Spire low
to high, with very faint to marked and granu-
lar spiral lines. Body whorl 60-80% of shell
length, smooth or sculptured like spire. Ap-
erture 70-80% of length of body whorl, oval-
elongate; columellar tooth small, very oblique;
anterior parietal tooth stronger, perpendicular
to (Auriculinella) or weakly oblique to columel-
lar axis; smaller posterior parietal teeth some-
times present; outer lip thin and sharp to thick
and weakly reflected, smooth internally. Pro-
toconch smooth, prominent, with umbilicus-
like slit in apex.

Radula with central tooth small, triangular;
lateral teeth bicuspid, with endocone smaller
than mesocone; marginal teeth similar to lat-
eral teeth but smaller.

Animal whitish; eyes often concealed by
thick skin; tentacles short, subcylindric or
with dilated tips; foot entire (Ellobium) or
transversely divided. Mandible corneous,
semilunate. Stomach tripartite; mid-section
very muscular. Spermiduct separates from
oviduct before the latter leaves posterior
glandular complex; anterior mucous gland
covers entire length of vagina; spermiduct
surrounded by prostate gland and might
communicate with base of bursa duct near
vaginal opening. Penis large and complex to
small and simple (Blauneria); associated vas
deferens adheres to penis. Visceral nerve
ring long, with evidence of streptoneury in
Ellobium; right parietovisceral connective
very short.

Remarks: There has been confusion in defin-
ing the limits of the subfamily Ellobiinae.
Pfeiffer (1853a) was the first to try to group
the genera of the Ellobiidae into higher taxa.
On the basis of the absence or presence of a
reflected outer lip he recognized the subfam-
ilies Melampea and Auriculea. He assigned
Pythia, Auricula [= Ellobium] and Carychium
to the latter group. The two subfamilial
names were emended to Melampinae and
Auriculinae by H. & A. Adams (1855a). Pfeiffer
(1854b), after seeing the unpublished manu-
script of the Genera of Recent Mollusca by H.
& A. Adams (1855), mentioned some of the
Adams’ conclusions, including the names
Melampinae and Ellobiinae, and it is H. & A.
Adams (in Pfeiffer) who should be credited
with the introduction of the latter name (see
the remarks under the Ellobiidae). Pfeiffer
(1854b) continued to use his previous names

and to the existing list of the Auriculea he
added Plecotrema [= Laemodonta], Cassid-
ula, Alexia [= Myosotella] and Blauneria. Two
years later in his Monografia Pfeiffer (1856a)
tentatively included the genus Leuconia [=
Auriculinella] in this subfamily.

Odhner (1925) noted the peculiar radula of
Ellobium and he admitted only this genus to
the subfamily. He wrongly stated that Ello-
bium (E.) aurismidae lacks the central radular
tooth (Fig. 17). Zilch (1959), who used con-
chological characters, also considered the
Ellobiinae monotypic. Thiele (1931), on the
basis of radular morphology, reached quite
different conclusions and he included in the
Ellobiinae the subfamilies Melampinae, Pythi-
inae and Cassidulinae.

Studies of the comparative anatomy of the
group are essential to an understanding of
the taxonomic relationships within the Ellobi-
inae and of the entire family as well. Morton
(1955с) noticed the similarity of the reproduc-
tive tracts of Auriculinella and Ellobium, and
placed those two genera in the subfamily El-
lobiinae. Likewise on the basis of reproduc-
tive structures Marcus (1965) and Marcus 4
Marcus (1965b) added В/аипепа. In spite of
the sinistrality of В/аипепа and the fact that
Blauneria and Auriculinella are much smaller
than Ellobium, the dentition of the inner lip
shows a constant pattern in all three genera
of the subfamily. This conchological similar-
ity, corroborating the evidence shown by the
reproductive system, makes these features
useful phylogenetic characters. | therefore
concur with the inclusion of Auriculinella and
Blauneria in the subfamily Ellobiinae.

Habitat: The various genera of the Ellobiinae
live in somewhat different habitats. Ellobium
is common on the muddy surface of Indo-
Pacific mangroves, just below the high-tide
mark, around roots and decaying wood
(Berry et al. 1967). Blauneria lives buried in
the black sediment, and under rocks and rot-
ting vegetable material at the high-tide mark
(Marcus & Marcus 19656; Martins, personal
observation). Auriculinella lives closer to the
low-tide mark than the other two genera; in
the Azores it lives under rocks buried in
gravel, sometimes into the intertidal zone
(Martins, 1980).

Range: The Ellobiinae have a worldwide dis-
tribution, with only partial overlap of the dif-
ferent genera. Ellobium, which is character-
istic of the Indo-West Pacific mangroves, has
a single representative in the tropical Eastern

178 MARTINS

Pacific and another in the tropical Western
Atlantic. Blauneria occurs only in the Western
Indo-Pacific and in the Western Atlantic. Au-
riculinella is restricted to the Mediterranean,
the eastern North Atlantic and Macaronesian
Islands.

The subfamily seems to have had a
Tethyan distribution, which is shown by the
present distribution of Ellobium and by the
presence of Ellobium and the Blauneria-like
Stolidoma Deshayes in the Jurassic and Oli-
gocene deposits of Europe (Degrange-
Touzin, 1893; Zilch, 1959; Huckriede, 1967).

Genus Ellobium Röding, 1798

Ellobium Röding, 1798: 105. Type species by
subsequent designation of Gray (184 7a):
Ellobium midae Röding, 1798 [= Bulla
aurismidae Linnaeus, 1758].

Auricula Lamarck, 1799: 76. Type species by
monotypy: Auricula midae (Röding,
1798) [= Bulla aurismidae Linnaeus,
1758].

Auriculus Montfort, 1810: 310. Type species
by monotypy: Auriculus judae Montfort,
1810 [= Bulla aurisjudae Linnaeus, 1758].

Marsyas Oken, 1815: 305 [new name for El-
lobium Röding].

Geovula Swainson, 1840: 344 [new name for
Ellobium Röding].

Description: Shell moderately large and thin
(25 mm) to large and thick (100 mm) and cov-
ered with рае brown periostracum. Spire low
to moderately high, sculptured with granular
spiral lines crossed by more or less conspic-
uous axial cords. Body whorl about 80%
shell length, with same sculpture as spire,
sometimes weakly depressed dorsoventrally.
Aperture about 80% length of body whorl;
small, very oblique, twisted columellar tooth;
stronger anterior parietal tooth; posterior pa-
rietal tooth sometimes present; outer lip thin
to thick, sharp to weakly reflected.

Radula with central tooth small, very nar-
row, without mesocone, with ectocones
curved inwards; lateral teeth with very wide,
bicuspid crown; marginal teeth similar to lat-
eral teeth, but smaller.

Remarks: The name Ellobium Róding, 1798,
was ignored for a long time in favor of its
junior synonym Auricula Lamarck, 1799.
Most probably the reason for maintaining the
junior name was the acceptance of La-
marck's work and ignorance of the Bolten
Catalogue published by Róding in 1798 (Fis-

cher, 1857; Dall, 1915). The vernacular name
Auricule was first published by Lamarck in
the Actes de la Société d'Histoire Naturelle
de Paris in 1795 or 1796 (fide Férussac, 1821:
95), but the Latinized name Auricula first ap-
peared in Lamarck's (1799) Prodrome, pub-
lished in the Mémoires of the same society.

Montfort (1810) pointed out that Lamarck
(1799) had confused Auricula midae and Au-
ricula judae by including in the references Ar-
genville’s (1757: 226, pl. 10 [13], fig. G) “oreille
de Midas,” which Montfort identified with Au-
ricula judae. Montfort, then, renamed La-
marck’s genus Auriculus and selected for its
type species Auriculus judae [= Ellobium (E.)
aurisjudae (Linnaeus)]. Pfeiffer (1876) pre-
ferred Auriculus Montfort to Ellobium Róding
and Auricula Lamarck, both of which he dis-
missed as vague, owing to the heterogeneous
assemblage of species that they included.

The genus Ellobium is conchologically well
Characterized by its auriform shape, by a
finely reticulate sculpture and by the conspic-
uous straw-colored to dark brown репо-
stracum (Figs. 4-9). The central tooth of the
radula is greatly reduced but not lost in Ello-
bium (E.) aurismidae, as Odhner (1925) erro-
neously reported (Fig. 17). The mesocone of
the central tooth has been lost and the ecto-
cones curl inwards and resemble a pair of
pincers (Figs. 13, 14).

Zilch (1959), on the basis of conchological
characters, recognized the subgenera Ello-
bium s.s. and Auriculodes Strand. Ellobium
was Characterized as having a large, thick
shell with a thick, reflected outer lip (Fig. 9),
whereas Auriculodes had a smaller, thinner
shell, with the outer lip sharp and weakly re-
flected (Figs. 4-8). Some scattered informa-
tion on the reproductive system of species
belonging to both subgenera (see remarks
under Auriculodes) indicates that the penis
and vagina are usually more complex in Ello-
bium s.s. More detailed research on a greater
number of species of both subgenera is
needed, however, to clarify the relative taxo-
nomic positions of Ellobium s.s. and Auricu-
lodes. Pending additional information, on the
basis of shell thickness, | concur with Zilch
(1959) in the recognition of these subgenera.

Subgenus Auriculodes Strand, 1928

Autonoe Guppy, 1868: 244. Type species by
monotypy: Autonoe riparia Guppy, 1868
[= Auricula dominicense Ferussac,
1821]. Non Leach, 1852.

WESTERN ATLANTIC ELLOBIIDAE 179

Auriculina Kobelt, 1898: 77. Type species by
original designation: Auricula (Auriculina)
дапдейса Pfeiffer, 1855. Non Grateloup,
1838, nec Agassiz, 1847.

Auricella Móllendorff, 1898: 160. Type spe-
cies by original designation: Auricula
(Auricella) auricella Férussac, 1821 [=
Bulimus auricula Bruguière, 1789]. Non
Jurine, 1817.

Auriculodes Strand, 1928: 64 [new name for
Auriculina Kobelt, 1898].

Autonoella Wenz, 1947: 36 [new name for
Autonoe Guppy, 1868].

Description: Shell to 25 mm long, thin to
somewhat solid. Spire with fine spiral lines,
sometimes granular and crossed by axial
granular cords. Body whorl not flattened dor-
soventrally, smooth and shiny or with granular
appearance as in spire. Inner lip of aperture
with very oblique, twisted columellar tooth
and somewhat stronger, weakly oblique pa-
rietal tooth; outer lip sharp, sometimes slightly
thick and somewhat sinuous at mid-length,
slightly reflected in gerontic specimens.

Animal with portion of vagina anterior to
confluence with bursa duct straight, very
short; associated vas deferens adhering to
anterior vagina; penis moderately long,
straight; associated vas deferens adhering to
penis.

Remarks: Guppy (1868), on the basis of a
single beach specimen, introduced Autonoe
[= Autonoella Wenz], which he considered al-
lied to Melampus and Laimodonta [= Laem-
odonta]. Later, in the revised list of the spe-
cies of Trinidad, Guppy (1872: 7) observed
under Synonyms, etc., “Comp. Auricula pel-
lucens [= Ellobium (A.) dominicense (Férus-
sac)].” Thiele (1931) considered Autonoe a
subgenus of Melampus, as did Zilch (1959)
for Autonoella Wenz, a replacement name for
the preoccupied Autonoe Guppy. From the
original description of Autonoella riparia
(Guppy), and from Guppy’s illustration (1871:
pl. 17, fig. 1), it seems that the specimen con-
sidered was a juvenile of Ellobium (A.) domin-
icense (Férussac). In view of this, | consider
Autonoella Wenz a junior synonym of Auric-
ulodes Strand.

Kobelt (1898) proposed Auriculina at the
same time as Móllendorff (1898) introduced
Auricella for the smaller and thinner-shelled
forms of Ellobium s.l. Because both names
were preoccupied, Strand (1928) introduced
the substitute name Auriculodes for Kobelt's
Auriculina.

Only two species of Auriculodes have been
investigated anatomically, and they appar-
ently differ greatly from each other in their
pallial gonoducts. According to Marcus 4
Marcus (1965b) and Martins (this study) Ello-
bium (A.) dominicense has a very short,
straight vaginal section anterior to the con-
fluence with the bursa duct, and a moder-
ately long, straight penis. Knipper & Meyer
(1956) briefly described the reproductive sys-
tem of Ellobium (A.) gaziense (Preston, 1913)
and they mentioned the lack of separation
between male and female ducts. This feature
is not typical of the subfamily and could lead
to removal of Ellobium (А.) gaziense, a spe-
cies with typical Auriculodes shell (Fig. 8),
from the Ellobiinae. Knipper 8 Meyer's rep-
resentation of the nervous system 1$ so sim-
ilar to that of Ellobium (A.) dominicense (Mar-
tins, this paper), however, that the accuracy
of their report on the reproductive system
should be questioned instead. Apparently
there are variations in the penial structure of
Ellobium s.s. as well. The highly coiled penis
of Ellobium (E.) aurisjudae (Linnaeus) is typi-
cal ofthe nominate subgenus (Morton, 1955b;
Berry et al., 1967). Sumikawa 8 Miura (1978)
observed a thick, straight penis in Ellobium
(E.) chinense (Pfeiffer) although this species
retains the characteristic long, coiled anterior
vagina. Odhner (1925: pl. 1, fig. 10), on the
other hand, represented a small, somewhat
thickened, straight penis, and an equally
straight vagina for Ellobium (E.) subnodosum
(Metcalfe, 1851). All of these scattered ana-
tomical observations on the genus hardly al-
low conclusions to be drawn concerning the
correlation between conchological and ana-
tomical characters of these two subgenera,
but | find the conchological characters suffi-
cient to justify the separation of Auriculodes
from Ellobium s.s.

Habitat: Species of the subgenus Auricu-
lodes prefer to live above the high-tide mark
of mangrove swamps, gathering wherever
there is rotten wood (Morrison, 1946; Marcus
& Marcus, 1965b; Keen, 1971; Martins, per-
sonal observation).

Range: The subgenus Auriculodes is known
from the eastern coast of Africa (Knipper &
Meyer, 1956) and throughout the Indo-Pa-
cific region. It is represented along the west-
ern coast of Central America by Ellobium (A.)
stagnale (Orbigny, 1835) and in the West In-
dian region to Brazil by the closely related
Ellobium (A.) dominicense (Férussac).

180 MARTINS

Ellobium (Auriculodes) dominicense
(Férussac, 1821)
Figs. 4-7, 10-16, 18-22

Auricula dominicensis Férussac, 1821: 103
[Santo Domingo Island (Hispaniola); lec-
totype herein selected MNHNP (Fig. 4));
Beck, 1837: 103; Beau, 1858: 15.

Auricula pellucens Menke, 1828: 78 [Demer-
ara (Guyana), South America; location of
type unknown]; Menke, 1830: 36, 131;
Kúster, 1844: 17, pl. 2, figs. 16, 17; Pfeif-
fer, 1854b: 151; Pfeiffer, 1856a: 137; Bin-
ney 8 Bland, 1870: 87; Simpson, 1889:
68.

Conovulus pellucens (Menke). Voigt, 1834:
hl

Ellobium pellucens (Menke). H. & A. Adams,
1855b: 237; Morrison, 1951b: 10; Perry
& Schwengel, 1955: 197, pl. 39, fig. 185;
Morrison, 1958: 123; Marcus, 1965:
124-128 [taxonomy]; Marcus & Marcus,
1965b: 426-438, pl. 1, figs. 1-7, pl. 2,
figs. 8-11, pl. 3, figs. 12-16 [anatomy,
ecology, taxonomy]; Rios, 1970: 139;
Abbott, 1974: 334, fig. 4106 [illustration
from Dall (1885)]; Rios, 1975: 159, pl. 48,
fig. 769; Altena, 1975: 88; Vokes &
Vokes, 1983: 60, pl. 22, fig. 18.

Autonoe riparia Guppy, 1868: 244 [Mayaro
Point, Trinidad; type presumed to be in
Victoria Institute, Trinidad, destroyed by
fire in 1920 (Sherborn, 1940); Guppy,
1871: 306, pl. 17, fig. 1 [type figured];
Guppy, 1872: 7.

Melampus riparius (Guppy). Pfeiffer, 1876:
STE

Auriculus pellucens (Menke). Pfeiffer, 1876:
359.

Auricula (Auriculastrum) pellucens Menke.
Dall, 1885: 275, pl. 18, fig. 8; Dall, 1889:
90, pl. 47, fig. 8; Maury, 1922: 54.

Auriculastra pellucens (Menke). Kobelt, 1898:
101, pl. 15, figs. 5, 6; Haas, 1950: 197;
Ewald, 1963: 11-14 [larval history].

Melampus (Autonoe) riparius (Guppy). Ko-
belt, 1898: 213, pl. 25, figs. 5, 6; Thiele,
1931: 467.

Auriculastrum pellucens (Menke). С.М.
Johnson, 1934: 158; M. Smith, 1937, pl.
67, fig. 8 [plate from Dall (1885)]; Webb,
1942, pl. 11, fig. 21; M. Smith, 1951: 145,
pl. 55, fig. 2, pl. 67, fig. 8; Coomans,
1958: 103.

Melampus (Autonoella) riparius
Zilch, 1959: 66, fig. 210.

(Guppy).

Description: Shell (Figs. 4-7, 10, 11) to 27
mm long, oval-elongate, somewhat solid,
whitish-yellowish, covered with brownish pe-
riostracum. Spire with as many as eight
weakly convex whorls; sculpture as in sub-
genus. Body whorl about 85% of shell length,
subcylindric, smooth or with same sculpture
as spire. Aperture about 80% length of body
whorl. Inner partition of whorls occupying
one-third of body whorl (Fig. 6). Protoconch
smooth, prominent, with about one whorl vis-
ible; lip weakly reflected at base, forming um-
bilicus-like perforation in apex (Figs. 10, 11).

Animal white; tentacles partly retractable,
moderately long, subcylindrical, with swollen
tip; eyes inside base of tentacles, deep in
integumentum, barely visible; foot entire;
mantle skirt white; anal opening continued by
fold of mantle skirt forming longitudinally split
tubular extension. Kidney long, narrow, whit-
ish.

Radula (Figs. 13-16, 18) with formula
(26+1+26)x70. Central tooth small; base
roughly rhombic; posterior portion elongate,
emarginate at anterior quarter, where crown
of next tooth seems to articulate; crown very
small; mesocone lacking; endocones thin,
sometimes curled inwards. Lateral and mar-
ginal teeth not sharply distinct, here de-
scribed always as lateral teeth; first seven to
12 with base short and wide, weakly pro-
jected lateroanteriorly, with median anterior
notch with which posterior process of crown
of next tooth articulates; crown wide, roughly
triangular, bicuspid, with conspicuous poste-
rior process; mesocone wide, somewhat
rounded anteriorly; endocone sometimes
barely defined, mainly in adult specimens;
gradual narrowing of crown and somewhat
sharper definition of endocone marks teeth
12 to 21; base shorter and narrower than that
of first group of lateral teeth, with lateral pro-
jection resembling basal ectocone; crown
somewhat narrow, elongate; endocone first
very rudimentary, then absent; no clearly de-
fined ectocone.

Digestive system with mandible solid,
crescentic, with concave, sharp anterior
edge and tapered ends (Fig. 12). Salivary
glands fusiform, separated from each other,
attaching to whitish esophagus by thin liga-
ments. Stomach tripartite (Fig. 19); anterior
portion membranous, comprising cardiac
and pyloric regions; mid-portion (gizzard)
very muscular, subcylindric; gastric caecum
thin, receiving posterior diverticulum anteri-

WESTERN ATLANTIC ELLOBIIDAE 181

FIGS. 4-11. Ellobium. (4) E. (A.) dominicense (Férussac), lectotype (MNHNP), Santo Domingo [Hispaniola],
sl 16.2 mm. (5) E. (A.) dominicense, Demerara, Guyana (ANSP 22251), sl 22.3 mm. (6) E. (А.) dominicense,
Big Torch Key, Florida, sl 20.6 mm. (7) E. (А.) dominicense, Big Torch Key, Florida, sl 23.0 mm. (8) E. (А.)
gaziense (Preston), syntype (BMNH 1969103), Gazi, British East Africa [Kenya], sl 18.2 mm. (9) E. (E.)
aurismidae (Linnaeus), Malaysia, sl 90.4 mm. (10) Е. (A.) dominicense, lateral view of spire and protoconch,
Big Torch Key, Florida. (11) E. (А.) dominicense, top view of spire and protoconch, Big Torch Key, Florida.
Scale 1 mm.

orly. Digestive gland bilobed, brownish; pos-
terior lobe conic, partly covering ovotestis.
Reproductive system (Fig. 20) with ovotes-
tis follicular, covering posterior portion of
stomach, beneath posterior lobe of digestive
gland; hermaphroditic duct thin, straight;
separation of male and female ducts just an-

terior to fertilization chamber; secondary
connection of posterior vas deferens with an-
terior end of bursa duct; bursa duct as long
as pallial gonoducts, emptying into oviduct a
short distance from female aperture; anterior
mucous gland covers oviduct as far as con-
fluence with bursa duct. Penis moderately

182 MARTINS

FIGS. 12-17. Ellobium, mandible and radular teeth. (12) E. (А.) dominicense, mandible, Big Torch Key,
Florida; scale 1 mm. (13) Е. (А.) dominicense, radula of young specimen, Big Torch Key, Florida, sl 5.4 mm;
scale 100 um. (14) E. (А.) dominicense, radula of young specimen, Big Torch Key, Florida, sl 5.4 mm; scale
100 um. (15) E. (А.) dominicense, radula of young specimen, Big Torch Key, Florida, sl 5.4 mm; scale 200
um. (16) E. (4.) dominicense, radula of young specimen, Big Torch Key, Florida, sl 5.4 mm; scale 200 um.
(17) E. (E.) aurismidae, radula, Malaysia, sl 90.4 mm; scale 200 um.

long, somewhat thin, simple; ramifications of
right tentacular retractor muscle attach to pe-
nis near male aperture; associated vas defe-
rens adhering to penis; penial retractor short,
attaching to nuchal region.

Nervous system (Fig. 21) with ganglia

wrapped in thick connective tissue; cerebral
commissure twice length of cerebral gan-
glion; pedal commissure very short; left ce-
rebropedal and cerebropleural connectives
somewhat longer than right ones, about as
long as cerebral commissure; pleuroparietal

WESTERN ATLANTIC ELLOBIIDAE 183

Cc 1L 2L 7L 8L

17L 20L 24L

(AP

9L 10L yi 12L
я |
ON (
=

FIG. 18. Ellobium (A.) dominicense, radula, Big
Torch Key, Florida. Scale 10 um.

FIG. 19. Ellobium (A.) dominicense, stomach, Big
Torch Key, Florida. Scale 1 mm.

connectives very long; left parietovisceral
connective shorter than right one, beneath
branch of columellar muscle coming from be-
hind right tentacle; right parietovisceral con-
nective crossing over left one before insertion
in visceral ganglion (rudiment of chiasto-
neury); cerebral ganglia as large as pedal
ganglia; left parietal ganglion double; anterior
portion of left parietal ganglion giving off
nerve to artery in mantle skirt, posterior por-

Е
hd в У
= RE,
per $ su A EN

FIG. 20. Ellobium (A.) dominicense, reproductive
system, Big Torch Key, Florida. Scale 1 mm.

tion giving off pallial internal and parietal cu-
taneous nerves; osphradial ganglion present
on pneumostomal nerve; tentacular nerves
split at their origin; penial nerve coming off
median lip nerve.

Remarks: Ellobium (A.) dominicense (Férus-
sac, 1821) has been considered a synonym
of Ellobium (А.) pellucens (Мепке, 1828) al-
though Dall (1885: 276) stated that Férus-
sac's species was described in such a way
as to be unidentifiable. Férussac's reference
(1821: 103) to the sculpture and size of the
shell and comparison with Auricula auricella,
which he had just introduced, constitute
enough indication to recognize the species.
Contention might arise owing to the fact that
Férussac's Auricula auricella was not de-
scribed (Pfeiffer, 1856a: 134, footnote), but
the author referred to Bulimus auricula Bru-
guière, 1789, Lister (1770: pl. 577, fig. 326)
[error for 32b] and Gualtieri (1742: pl. 55, fig.
F). Férussac's description contains enough
information to allow identification of the spe-
cies and his name has priority over that of
Menke (1828).

Emerson & Jacobson (1976) considered
Ellobium auricula (Bruguière) to be the earli-
est name for the West Indian Ellobium. Bru-
guiere (1789: 342) provided a description of a
Bulimus aurícula that indeed could apply to

184 MARTINS

gn ст ргсп тап

a prg1

plprc pmpn *
‘ x \

`

E A
и ‘ и 4
ppni / / plg
и
LA

0
‘ Ц

р!рп iu

D----
+
2

st ampn pen

FIG. 21. Ellobium (A.) dominicense, central nervous system, Big Torch Key, Florida. Scale 1 mm.

the West Indian shells. No locality was given
in the original description, and reference was
made to the Gualtieri and Lister illustrations
just cited. Férussac, as stated above, intro-
duced without description Auricula auricella
from Baie des Chiens Marins, New Cale-
donia. He mentioned in his synonymy of Bru-
guière’s name the same synonymic refer-
ences given by that author. The fact that
some Indo-Pacific species are conchologi-
cally very similar to the West Indian species
contributed to this confusion. Ellobium (A.)
dominicense has been stated erroneously to
live in Natal, East Africa (Krauss, 1848), prob-
ably the result of a misidentification of Ello-
bium (A.) gaziense (Preston, 1913), and
Pfeiffer (1856a) considered the Indian Ello-
bium ceylanicum H. 8 A. Adams, 1854, a jun-
ior synonym of Auricula pellucens Menke,
1828 [= Ellobium (A.) dominicense (Ferussac,
1821)]. All this circumstantial evidence indi-
cates that Bruguiere (1789) had described an
Indo-Pacific shell, which was deposited at
the Museum d’Histoire Naturelle de Geneve
(Mermod & Binder, 1963). I therefore dis-
agree with Emerson & Jacobson (1976), who
misidentified Bruguiere’s name for the West
Indian species.

Ellobium (A.) dominicense has been placed

wrongly in Auriculastra [Auriculastrum is an
unjustified emendation (Marcus & Marcus,
1965b)] by Dall (1885) and others. Martens
(1880) created Auriculastra as a subgenus of
Marinula for those species similar to Ellobium
s.s., but with visible eyes and knobbed ten-
tacle tips. Ellobium (A.) dominicense has
these characteristics, a fact which might ex-
plain Dall’s decision. However, Auriculastra
elongata (Parreyss, 1845), also originally
listed by Martens and very similar to the type
species, Auriculastra subula (Quoy & Gaim-
ard, 1832), has a very different radula (Odh-
ner, 1925) and appears to belong in the
Pythiinae.

The nervous system and the radula of the
specimens of Ellobium (A.) dominicense here
examined, collected on Big Torch Key, Flor-
ida, differ from those of animals from Brazil
studied by Marcus & Marcus (1965b). The
central nervous system of the Floridian spec-
imens is very similar to that of Ellobium (A.)
gaziense (Preston), illustrated by Knipper &
Meyer (1956: 106, fig. 6), differing only in that
the left parietal ganglion is double in Ellobium
(A.) dominicense. In the Brazilian specimens
(Marcus 8 Marcus, 1965b: 431, pl. 3, fig. 13)
there is only one left parietal ganglion. The
latter authors did not refer to the osphradial

WESTERN ATLANTIC ELLOBIIDAE 185

ganglion and their figures indicate that the
pleuroparietal connectives are shorter than
the cerebropleural connectives. In the Florid-
ian specimens the pleuroparietal connectives
are three times longer than the cerebropleu-
ral connectives. The radula of the specimens
from Florida is very similar to those of Ello-
bium (E.) aurismidae (Fig. 17) and Ellobium
(E.) aurisjudae, both from Malaysia. Marcus &
Marcus (1965b: 433, pl. 2, fig. 8) described
and figured a tricuspid central tooth with a
small, triangular mesocone and rudimentary
ectocones. In the specimens from Florida the
mesocone is lacking and the slender ecto-
cones are sometimes curved inwards, re-
sembling small fangs. Preserved material
from northern South America was not avail-
able for comparative anatomical study; how-
ever, intrapopulational variability in shell
shape and intensity of sculpture is seen
throughout the range of the species, al-
though the sculpture seems to be more
marked in northern South American speci-
mens (Fig. 5). | am unsure about the phylo-
genetic significance of the anatomical differ-
ences observed in the Brazilian specimens.
Should further comparative anatomical re-
search establish that the South American
specimens are a separate taxon, Menke’s
name pellucens is available.

Deposited in the Muséum National d’His-
toire Naturelle de Paris are two syntypes of
Auricula dominicensis Férussac, from which
a lectotype is herein selected and figured
(Fig. 4).

Habitat: Ellobium (A.) dominicense lives in
protected embayments in which mangrove
growth is thin, buried in the soft black humic
sediment or under rotting logs seldom cov-
ered by high tide. It seems to be an oppor-
tunistic species, usually found in colonies
and apparently with very limited movement
once established. Great numbers of shells
clustered in a small area, indicative of former
colonies, are often found without living ani-
mals in the immediate vicinity. lt seems that
the colonies are destroyed by lack of food or
by some environmental change, even though
apparently suitable habitats exist a few
meters away (Ewald, 1963; Marcus & Mar-
cus, 1965b; Martins, personal observation).

Range: Florida, from Miami to Cedar Key
(Dall, 1885); Dominican Republic (Férussac,
1821); Haiti; Guadeloupe (Beau, 1858); Trin-
idad (Guppy, 1868); Yucatan, Mexico, to

90 75 60 45 30

FIG. 22. Ellobium (A.) dominicense, geographic
distribution. Open circle, locality from literature.

Cananeia, Brazil (Marcus & Marcus, 1965b).
(Fig. 22).

Specimens Examined: FLORIDA: Golden
Beach (MCZ 157854); Miami (ANSP 77056;
MCZ 104943); Virginia Key (USNM 338303);
Key Biscayne (ANSP 345210; USNM 700836,
700911); Coconut Grove (MCZ 201646); El-
liot Key (MCZ 110206); Key Largo (ANSP
192837; MCZ 243979; USNM 590644,
701421); Card Sound, Key Largo (A.M.); Rab-
bit Key (ANSP 88136); Big Pine Key (ANSP
106384); Big Torch Key (USNM 61046,
492482, 492484; A.M.); Middle Torch Key
(USNM 663960); Oyster Bay (USNM 37596);
Lossman Key (MCZ 291093); Cape Sable
(MCZ 291085, 292564; USNM 525156); Rog-
ers River (MCZ 3981); 2.5 km E of Chokolos-
kee Key (MCZ 58955); Harris Island, Ten
Thousand Islands (USNM 381326); Blue Hill
Island, near Goodland Point (ANSP 82742); $
of Cape Romano (ANSP 62833); Marco (MCZ
292565); Bonita Springs (MCZ 291088); Carl
E. Johnson Park, near Little Carlos Pass
(A.M.); Fort Myers (ANSP 66963, 140799;
USNM 87733, 492483); Punta Rassa (MCZ
291091, 291094, 292566; USNM 39804);
Punta Gorda (USNM 592297); Sanibel Island
(ANSP 170650; MCZ 13721, 291089,

186 MARTINS

291090, 292563); Bokeelia (MCZ 291087);
Wulfert (ANSP 219866). HAITI: lle-a-Vache
(USNM 403877, 404948). MEXICO: Silam,
Yucatán (ANSP 62656). VENEZUELA: М of
Sinamaica, Zulia (USNM 536129). GUYANA:
Demerara (ANSP 2225, 22241; MCZ 146522;
USNM 31572, 58857, 119552). FRENCH
GUIANA: Cayenne (MCZ 102934; USNM
126413). SURINAME: Saramacca (USNM
635276).

Genus Blauneria Shuttleworth, 1854

Blauneria Shuttleworth, 1854a: 148. Type
species by monotypy: Blauneria cuben-
sis (Pfeiffer, 1841) [= Voluta heteroclita
Montagu, 1808].

Blanneria Shuttleworth. Dall, 1885: 287 [in
synonymy; error for Blauneria].

Blaumeria Shuttleworth. Verrill, 1901: 35 [er-
ror for Blauneria].

Description: Shell to 8 mm long, elongate,
fragile, translucent, whitish, sinistral. Spire
with as many as nine flattened whorls. Body
whorl about 60% of shell length. Umbilicus
absent. Aperture about 70% of length of body
whorl, oval-elongate; inner lip with very small
columellar tooth; outer lip sharp, smooth. Pro-
toconch prominent, smooth, with about one
and one-half whorls visible.

Radula having central tooth with wide, tri-
angular, emarginate base; crown small, uni-
cuspid. Lateral teeth gradually becoming
smaller toward margin of radula, bicuspid,
with strong mesocone and much smaller en-
docone; no morphological distinction be-
tween lateral teeth and marginal teeth.

Animal whitish, translucent, with short,
cylindrical tentacles and very conspicuous
black eyes. Foot transversely divided. Ar-
rangement of organs sinistral. Separation of
male and female ducts just anterior to fertili-
zation chamber, before oviduct enters pos-
terior glandular complex; posterior vas defe-
rens secondarily communicates with anterior
end of bursa duct. Penis small, simple; asso-
ciated vas deferens adhering to penis. Con-
nectives of visceral nerve ring long.

Remarks: The genus Blauneria is readily
identifiable because it is the only sinistral el-
lobiid taxon. The history of this once enig-
matic small group, before it was placed tim-
idly in a separate genus by Shuttleworth
(1854a), is connected with that of the type
species Blauneria heteroclita (Montagu), and

will be discussed in the remarks under that
species.

Once it was discovered to be a member of
the Ellobiidae, the genus Blauneria was
placed in different subfamilies, depending
upon which character assumed greater im-
portance in the classification scheme of the
particular malacologist. Fischer & Crosse
(1880) included Blauneria and other “marine”
ellobiids with an elongated spire in the Auri-
culinae [= Ellobiinae]. Odhner (1925), on the
basis of radular characters, considered the
genus to belong to the Cassidulinae. Thiele
(1931), who based his classification largely
upon Odhner's radular studies, did not rec-
ognize the subfamilies Pythiinae and Cassid-
ulinae, and placed Blauneria, together with
many other genera, in the heterogeneous
subfamily Ellobiinae. Zilch (1959), probably
on the basis of conchological similarities with
the dextral Cylindrotis Móllendorff, 1895, re-
moved Blauneria to the Pythiinae. Finally,
Marcus (1965) and Marcus & Marcus (1965b),
followed by Hubendick (1978), included
Blauneria in the Ellobiinae owing to similari-
ties of the reproductive system with those of
Ellobium and Auriculinella. My anatomical
studies confirm the taxonomic conclusions of
these latter authors.

Habitat: Blauneria commonly lives in man-
groves at the high-tide mark in the sediment
under rocks and decaying branches. Pease
(1869: 60) reported that his Blauneria gracilis
from Hawaii lives in the same habitat as Pe-
dipes, in the crevices of stones covered at
high tide. He observed that Blauneria never
crawls on the sides or tops of the rocks dur-
ing low tide, but only around the base, which
was always wet.

Range: The genus is known from the warm
regions of the Indo-Pacific and from the trop-
ical Western Atlantic. There is no known fossil
record of this sinistral genus, but the concho-
logically closely related, dextral Stolidoma
Deshayes, 1863, has been recorded from
strata as old as the Paleocene of Europe (De-
grange-Touzin, 1893; Zilch, 1959). Zhu (1980)
described a Blauneria ? elliptiformis from the
Cretaceous of northeastern China.

Blauneria heteroclita (Montagu, 1808)
Figs. 23-40

Voluta heteroclita Montagu, 1808: 169 [Dun-
bar, Scotland (error), herein corrected to
Matanzas, Cuba; location of type un-

WESTERN ATLANTIC ELLOBIIDAE 187

known]; Laskey, 1811: 398, pl. 81, figs.
1, 2; Turton, 1819: 254.

Acteon heteoclita (Montagu). Fleming, 1828:
>37

Achatina (?) pellucida Pfeiffer, 1840: 252
[Cuba; location of type unknown].

Tornatellina cubensis Pfeiffer, 1841: 130
[Cuba; location of type unknown].

Auricula heteroclita (Montagu). Thorpe, 1844:
146.

Tornatella heteroclita (Montagu). Forbes &
Hanley, 1852: 526.

Blauneria cubensis (Pfeiffer). Shuttleworth,
1854a: 148; Franc, 1968: 525.

Blauneria pellucida (Pfeiffer). Pfeiffer, 1854b:
152; Pfeiffer, 1856a: 153; H. 8 A. Adams,
1858: 643, pl. 138, fig. 8; Binney, 1859:
175, pl. 53, fig. 2; Binney, 1860: 4; Binney,
1865: 21, text fig. 22; Mórch, 1878: 5.

Oleacina (Stobilus) cubensis (Pfeiffer). H. 8 A.
Adams, 1855a: 136.

Odostomia (Tornatellina) cubensis (Pfeiffer).
Shuttleworth, 1858: 73.

? Odostomia cubensis (Pfeiffer). Poey, 1866:
394.

Blauneria heteroclita (Montagu). Pfeiffer, 1876:
368; Arango y Molina, 1880: 60; Fischer
8 Crosse, 1880: 9, pl. 34, figs. 14, 14a,
14b [anatomy, radula, taxonomy]; Dall,
1885: 287, pl. 17, fig. 6; Dall, 1889: 92,
pl. 47, fig. 14; Simpson, 1889: 60;
Crosse, 1890: 259; Kobelt, 1900: 260, pl.
31, figs. 19, 20; Dall & Simpson, 1901:
369; Davis, 1904: 126; Peile, 1926: 88;
Thiele, 1931: 466; Bequaert & Clench,
1933: 538; C.W. Johnson, 1934: 160; M.
Smith, 1937: 147, pl. 67, fig. 14 [plate
from Dall (1885)]; Morrison, 1951b: 10;
Coomans, 1958: 104; Nowell-Usticke,
1959: 88; Zilch, 1959: 74, fig. 241;
Warmke & Abbott, 1961: 152; Marcus,
1965: 124-128 [taxonomy]; Marcus &
Marcus, 1965b: 438-446, pl. 4, figs.
25-29 [anatomy, taxonomy, habitat];
Rios, 1970: 139; Abbott, 1974: 334, fig.
4104 [illustration from Binney (1859)];
Altena, 1975: 87, fig. 42; Rios, 1975: 159,
pl. 48, fig. 768; Hubendick, 1978; 20, fig.
164, 24, fig. 176 [nervous and reproduc-
tive systems redrawn from Marcus &
Marcus (1965b)]; Vokes & Vokes, 1983:
60, pl. 31, fig. 19; Jensen & Clark, 1986:
458, pl. 153.

Blanneria pellucida (Pfeiffer). Dall, 1885: 287
[error for Blauneria; in synonymy].

Blaumeria heteroclita (Montagu).
1901: 35 [error for Blauneria].

Verrill,

Description: Shell (Figs. 23-32) with length
to 7 mm, elongate, fragile, transparent to
translucent, shiny, whitish. Spire with as
many as nine flat or weakly convex whorls;
very faint spiral lines on teleoconch, crossed
by irregular growth lines. Body whorl about
60% shell length in gerontic specimens, 70-
75% in young individuals. Aperture about
70% body whorl length, oval-elongate; inner
lip weakly canaliculate at base, with small,
very oblique columellar tooth, stronger, ob-
lique parietal tooth at mid-length of aperture;
outer lip sharp, smooth inside. Partition of
inner whorls occupying about three-quarters
of the body whorl (Fig. 26). Protoconch
smooth, well developed, with one and one-
half whorls visible, leaving umbilicus-like per-
foration on apex (Figs. 30-32).

Radula (Figs. 33-36) having formula (17 + 1
+ 17) x 70. Base of central tooth wide, tri-
angular, deeply emarginate anteriorly; crown
very small, narrow, unicuspid. Lateral teeth 15
to 18; base quadrangular, anteriorly oblique
away from central tooth, with small notch on
anterior edge; crown wider and longer than
base, bicuspid; mesocone strong, long; en-
docone less than half the length of mesocone;
from about sixth lateral tooth outward a pro-
cess develops on posterolateral edge of
crown, which articulates with notch in base of
next tooth. Marginal teeth not morphologi-
cally distinct from lateral teeth except in grad-
ual decrease in size.

Animal has external anatomy as in genus.
Stomach (Fig. 37) with thin, somewhat di-
lated cardiac region, and smaller, slightly
thicker pyloric region; gizzard very muscular,
barrel-shaped; gastric caecum invaginable,
without posterior diverticulum.

Reproductive system (Fig. 38) with ovotes-
tis apical, granular, orange; hermaphroditic
duct simple, with some pouch-like dilations
(seminal vesicle) as it approaches albumen
gland; male and female ducts separating just
anterior to fertilization chamber; spermiduct
thick, covered with prostatic tissue, commu-
nicating with bursa duct where the latter
opens into vagina; anterior mucous gland
covers oviduct until confluence with bursa
duct. Penis small, simple; associated vas
deferens adhering to penis; penial retractor
very short, attaching to nuchal region.

Nervous system (Fig. 39) with cerebral
ganglia largest; cerebral commissure as long
as width of cerebral ganglion; pedal commis-
sure very short; right cerebropedal and cere-
bropleural connectives longer than left coun-

188 MARTINS

FIGS. 23-35. Blauneria heteroclita (Montagu). (23) Hungry Bay, Bermuda, sl 6.7 mm. (24) Hungry Bay,
Bermuda, sl 5.2 mm. (25) Hungry Bay, Bermuda, sl 4.3 mm. (26) Hungry Bay, Bermuda, sl 6.3 mm. (27)
Plantation Key, Florida, sl 3.5 mm. (28) Matanzas, Cuba (MCZ 131769), sl 3.7 mm. (29) Isla Mujeres,
Yucatán, Mexico (R.B.), sl 3.5 mm. (30) Lateral view of spire and protoconch, Big Pine Key, Florida. (31) Top
view of spire and protoconch, Crawl Key, Florida. (32) Top view of spire and protoconch, West Summerland
Key, Florida. (33) Lateral and central teeth of radula, Hungry Bay, Bermuda, sl 4.5 mm. (34) Lateral and
central teeth of radula, Hungry Bay, Bermuda, sl 4.5 mm. (35) Lateral teeth of radula, Hungry Bay, Bermuda,
si 4.5 mm. Scale, Figs. 30-32, 1 mm; Figs. 33-35, 100 um.

WESTERN ATLANTIC ELLOBIIDAE 189

C IL 2b 3L

A\ } 7 №
À | }
A A = ES

8L 9L 13L 14L 15L

ES == AS der
uy | |
YA | и \ V

FIG. 36. Blauneria heteroclita, radula, Hungry Bay,
Bermuda. Scale 10 um.

FIG. 37. Blauneria heteroclita, stomach, Hungry
Bay, Bermuda. Scale 1 mm.

terparts; left pleuroparietal and right parie-
tovisceral connectives very long, the latter
somewhat shorter than the former; right pleu-
roparietal and left parietovisceral connectives
about same size, about half length of cerebral
commissure.

Remarks: Ваипепа heteroclita (Montagu)
was originally thought to belong to the En-
glish malacofauna. The appearance of this
Western Atlantic shell on the shores of Dun-
bar, Scotland, can be attributed to the dump-
ing of ballast of ships from the West Indies.

FIG. 38. Blauneria heteroclita, reproductive sys-
tem, Hungry Bay, Bermuda. A-C, transverse sec-
tions and their locations. Scale 1 mm.

FIG. 39. Blauneria heteroclita, central nervous sys-
tem, Hungry Bay, Bermuda. Scale 1 mm.

This little, fragile and elegant shell puzzled
the European naturalists for some time.
Pfeiffer, within 12 months, introduced the
names Achatina ? pellucida (1840) and Tor-
natellina cubensis (1841) for specimens from
Cuba. H. 8 A. Adams (1855b, 1858) treated
those two names as referring to species in
very different groups. They assigned Torna-
tellina cubensis to the terrestrial Oleacina,
and they followed Pfeiffer (1854b) in allocat-
ing Achatina ? pellucida to Blauneria. The
species in question was placed in seven dif-
ferent genera before Shuttleworth (1854a)

190 MARTINS

hesitantly proposed that “Odostomia cuben-
sis’’ probably should belong to a separate
genus. Shuttleworth, in a presentation made
at the Lyceum of New York a month before
the appearance of that paper but published
four years later, had considered the species
to be marine on the word of the naturalist
Blauner. Pfeiffer (1854b), upon receiving a
communication from Gundlach that the ani-
mal in question had conspicuous eyes at the
base of the tentacles (Pfeiffer 1856a: 153),
immediately adopted Shuttleworth's name
Blauneria and placed the genus within the
Auriculidae [= Ellobiidae].

| have found some discrepancies between
the specimens | studied and those from Bra-
zil examined by Marcus 8 Marcus (1965b).
The Marcuses stated (p. 443) that the en-
docone of the radular teeth is basal. The SEM
photographs of my Bermudian specimens
clearly show the endocone as part of the
crown, not of the base (Figs. 33-35). Another
discrepancy is found in the lengths of the
pleuroparietal connectives of the visceral
nerve ring (pl. 5, fig. 28). Based upon my ob-
servations in the current study | suspect that
the Marcuses reversed the right and left con-
nectives.

Blauneria differs from all other ellobiids in
its sinistrality. Gerontic specimens have a
very elongate and slender shell (Fig. 23).
Most commonly, however, the body whorl of
the shell is longer and wider than the spire
(Figs. 27-29). This form has been the one
commonly illustrated, represented by Binney
(1865) and copied by Dall (1885), M. Smith
(1937) and Abbott (1974).

Habitat: Blauneria heteroclita lives in man-
groves above the high-tide mark, where it is
usually deeply buried in the soft sediment un-
der rocks, rotten wood or on the roots of the
propagules, where it occurs with Laemo-
donta, Creedonia and Microtralia. Marcus 4
Marcus (1965b) stated that these animals are
common in decaying banana trees washed
ashore in Cananeia, Brazil.

Range: Bermuda; Florida to Texas and Yu-
catán, Mexico; West Indies; Panama (Olsson
8 McGinty, 1958); Suriname (Altena, 1975);
Brazil (Fig. 40).

Binney (1859: 176) stated, “Dr. Foreman
collected a few specimens in a garden of
Washington city. He believes them to have
been brought on plants from Charleston,
S.C.” Both places are distant from the range

90 75 60° 45 30

FIG. 40. Blauneria heteroclita, geographic distribu-
tion. Open circle, locality from literature.

of the species and, because there has been
no confirmation of either record, | do not in-
clude them in the range of the species.

Specimens Examined. BERMUDA: Fairyland
(ANSP 99076); Old Road, Shelly Bay (A.M.);
Cooper's Island (ANSP 131647); Hungry Bay,
S of Ely's Harbour (both A.M.). FLORIDA
(USNM 39843, 67953): St. Augustine (USNM
663064); Rose Bay, N of New Smyrna Beach
(A.M.); Miami (MCZ uncatalogued); Barnes
Sound (ANSP 196748; MCZ 291100); Key
Largo (MCZ uncatalogued; USNM 597460);
Tavernier Key (USNM 492513); S of Ocean
Dr., Plantation Key (A.M.); Lignumvitae Key
(ANSP 156648; MCZ 294648); Lower Mate-
cumbe Key (USNM 492521); Long Key
(A.M.); Grassy Key (ANSP 397277; MCZ
291102; A.M.); Crawl Key (A.M.); Big Pine
Key (ANSP 104106); end of Long Beach Drive
and W of Kohen Avenue, both Big Pine Key
(both A.M.); Sugarloaf Key (ANSP 88804,
104107); Boca Chica Key (USNM 270352);
Cape Sable (MCZ 291099, 291101); Marco
(ANSP 22470; USNM 37615, 37616); Semi-
nole Point (ANSP 105422); Starvation Key
(ANSP 130061); Fort Myers (USNM 492512);
E of St. James, Pine Island (ANSP 93432);
Captiva Island (ANSP 149907); Sarasota Bay
(USNM 30626); Mullet Key (USNM 652410,

WESTERN ATLANTIC ELLOBIIDAE 191

653108; A.M.); Tampa Bay (USNM 37614);
Boca Ciega Bay (ANSP 9570); Shell Key
(USNM 466212); Clearwater Island (ANSP
9350). ALABAMA: Coden Beach (USNM
422371). TEXAS: Galveston (MCZ 227843);
Port la Vaca (MCZ 223050); N end of Padre
Island, 45 km S of Рой Aransas (MCZ
228745). MEXICO: Isla Mujeres, Quintana
Roo, Yucatán (В.В.). BAHAMA ISLANDS:
GRAND BAHAMA ISLAND: North Hawksbill
Creek (ANSP 370564); South Hawksbill
Creek (ANSP 371810); GREAT ABACO IS-
LAND (ANSP 299496); ANDROS ISLAND:
South Mastic Point (A.M.); Stafford Lake
(ANSP 151931); Mangrove Key (USNM
180672, 269947, 270198); Smith's Place,
South Bight (USNM 257569, 269649); Linder
Key (USNM 270224); NEW PROVIDENCE IS-
LAND: Nassau (MCZ uncatalogued); SE
shore of Lake Cunningham (ANSP 299720);
Bonefish Pond (A.M.); ROYAL ISLAND
(USNM 468124); AKLINS ISLAND: between
Pleasant Point and Claret Cove (MCZ
225524). CUBA (ANSP 22471; MCZ uncata-
logued; USNM 39842, 57726, 492511): Ha-
bana (ANSP 130745, 326340; MCZ 233993);
Salt Works, Hicacos Peninsula (ANSP
157338); La Chorrera (MCZ 128256, 167956);
Сало (MCZ 167955); Matanzas (MCZ
131769; USNM 492510); Batabanó (ANSP
93730; MCZ 167957). JAMAICA (ANSP
16705, 22472; USNM 94765): Green Island
Harbor (USNM 440791); Montego Bay (ANSP
329122); Port Morant (USNM 423688); King-
ston (USNM 427130, 467555); Hunt's Bay
(USNM 427117). HAITI: lle-a-Vache (USNM
403701, 403859, 403872, 404947); Landep-
rie Bay (USNM 383264); between Vieux
Bourg and Baie des Flamands (USNM
402467); Aquin (USNM 403149); Bizoton
(USNM 403324). PUERTO RICO: Punta Are-
nas, N of Joyuda (A.M.); Puerto Real (A.M.).
VIRGIN ISLANDS: ST. THOMAS (ANSP
22473). CARIBBEAN ISLANDS: GRAND
CAYMAN ISLAND (ANSP 209768). BRAZIL:
Cananeia (ANSP 305213; USNM 699448).

Subfamily Pythiinae Odhner, 1925

Scarabinae Fischer & Crosse, 1880: 5.
Pythiinae Odhner, 1925: 14.

Description: Shell variable in size. Aperture
usually heavily dentate; one columellar tooth;
one to three, commonly two parietal teeth,
anterior one strongest; outer lip generally in-
ternally dentate.

Radula with mesocone of lateral teeth tri-
angular, usually pointed; marginal teeth be-
coming smaller toward margin, with as many
as three subequal cusps.

Animal with rudimentary anterior tentacles
sometimes present; foot entire. Pallial gono-
duct entirely hermaphroditic; anterior mu-
cous gland and prostate gland covering
spermoviduct along entire length; bursa duct
emptying near vaginal opening; spermatic
groove open in Pythia; penis simple; vas def-
erens adhering to penis externally or free in
haemocoel. Ganglionic connectives of vis-
ceral nerve ring long, leaving pedal ganglia
mid-way between cerebral ganglia and vis-
ceral ganglion; right parietovisceral connec-
tive longer than left one.

Remarks: Fischer & Crosse (1880) created
the subfamily Scarabinae for Scarabus Mont-
fort, 1810 [= Pythia Róding, 1798] on account
of its oddly shaped, dorsoventrally flattened
shell. Odhner (1925) used the name Pythiinae
because by that time Scarabus Montfort was
recognized as a junior synonym of Pythia
Roding; he included Alexia [= Myosotella] and
Blauneria on the basis of radular characters.
Cassidula and Ophicardelus were added by
Morton (1955c), who merged Odhner's Cas-
sidulinae with the Pythiinae. Zilch (1959) re-
verted to Odhner’s division and included in
the Pythiinae the Recent genera Pythia, Ova-
tella, Cylindrotis and Blauneria and removed
Ophicardelus and Cassidula to the Cassiduli-
nae. Marcus (1965) and Marcus 8 Marcus
(1965b) noted that in Blauneria the spermi-
duct and oviduct separate before the her-
maphroditic duct enters the glandular com-
plex, and so removed this genus to the
Ellobiinae. In consideration of shell and ana-
tomical features, | have concluded that Lae-
modonta must be included in the Pythiinae.

Dall (1885) included Sayella within the El-
lobiidae and Zilch (1959) listed it, with a ques-
tion mark, within the Pythiinae. Morrison
(1939), however, showed that Sayella Dall is
not an ellobiid, but a pyramidellid opistho-
branch.

Separation of the Cassidulinae from the
Pythiinae, as Odhner (1925) proposed and
Zilch (1959) supported, 15 not justifiable. The
two groups are similar in the basic pattern of
the inner lip teeth of the shell aperture but
their radular morphology shows too much di-
versity and overlap to constitute a useful tax-
onomic character at the subfamilial level.
Both groups have a similar plan of the ner-

192 MARTINS

vous system and, for this reason, Morton
(1955c) regarded Odhner's Cassidulinae as
superfluous. The nervous system of the Cas-
sidulinae indeed shows the elongate right pa-
rietovisceral connective, characteristic of the
Pythiinae. Morton erroneously stated that the
pallial gonoduct of Cassidula is very similar to
that of Myosotella in that it remains hermaph-
roditic until the vaginal aperture. According
to Berry et al. (1967), Berry (1977) and Mar-
tins (personal observation) the vas deferens
of Cassidula aurisfelis (Bruguière) separates
from the oviduct some distance before the
vaginal opening, and runs free until entering
the neck skin to follow the spermatic groove.
This feature can be considered secondary to
the general pattern of the reproductive sys-
tem, however, for the bursa duct opens at the
same position relative to the separation of
the vas deferens in Cassidula as it does in the
other Pythiinae. The same arrangement oc-
curs in the Ellobiinae. Ellobium (E.) aurisjudae
also has a long, nonglandular vagina, which
is in accordance with the highly specialized
penial complex of the species. Ellobium (E.)
aurismidae, on the other hand, has a less
specialized penis and lacks the long, non-
glandular vagina (Morton, 1955c; Berry et al.,
1967; Martins, personal observation). In both
species, however, the bursa duct opens at
the anterior end of the glandular portion of
the oviduct.

In view of the similarities of the repro-
ductive and nervous systems of the two
groups, as well as their similar patterns of
apertural dentition, Morton's decision (1955c)
to merge the Cassidulinae with the Pythiinae
is hereby followed.

Habitat: The Pythiinae contain very primi-
tive ellobiids such as Pythia, Myosotella,
Ophicardelus and Cassidula. These groups
have left the proximity of the sea and are less
dependent upon that element than all other
halophilic ellobiids. Руа has acquired a
semiterrestrial habitat, and Myosotella, Oph-
icardelus and Cassidula were placed by
Morton (1955c) among the “supratidal and
estuarine ellobiids.” Laemodonta lives in
rocky areas at the high-tide mark, with Pe-
dipes, and in the mangroves at or just below
the high-tide mark, under rocks and fallen
branches.

Range: The Pythiinae have a worldwide
distribution. Pythia, Cassidula and Ophicar-
delus are characteristic of the tropical Indo-

Pacific; Laemodonta, also common in the
Indo-Pacific, is represented in the West Indies
by one species. Ovatella and Myosotella are
represented in the Mediterranean, but the
latter has been introduced to eastern North
America (Binney, 1859; Verrill, 1880), Califor-
nia (Hanna, 1939), western South America,
South Africa and Australia (Climo, 1982).

Genus Mysotella Monterosato, 1906

Phytia Róding, 1798. Gray, 1821: 231 [mis-
spelling of Pythia].

Phitia Gray. Blainville, 1824: 246 [misspelling
of Gray’s misspelling of Ру].

Phythya Gray. Deshayes, 1832: 762 [mis-
spelling of Gray’s misspelling of Руа].

Jaminia Brown, 1827, pl. 51. Type species by
subsequent designation of Gray (1847a):
Jaminia denticulata (Montagu, 1803) [ =
Auricula myosotis Draparnaud, 1801].
Non Risso, 1826.

Alexia “Leach” Gray, 1847a: 179. Type spe-
cies by monotypy: Alexia denticulata
(Montagu, 1803) [= Auricula myosotis
Draparnaud, 1801]. Non Stephens,
1835.

Kochia Pallary, 1900: 239. Type species by
subsequent designation of Monterosato
(1906): Alexia (Kochia) oranica Pallary,
1900 [= Auricula myosotis Draparnaud,
1801]. Non Frech, 1891.

Myosotella Monterosato, 1906: 126. Type
species by original designation: Myoso-
tella payraudeaui “Shuttleworth” Pfeif-
fer, 1856a [= Auricula myosotis Drapar-
naud, 1801].

Nealexia Wenz, 1920: 190 [new name for A/-
exia Gray, 1847, non Stephens, 1835].

Description: Shell to 10 mm long, fragile to
somewhat solid, pale yellow to purplish red.
Spire high, with as many as eight weakly con-
vex, spirally striated whorls; only one spiral
row of hairs in juveniles. Aperture oval-elon-
gate; inner lip with small, very oblique col-
umellar tooth, strong anterior parietal tooth
and usually one, sometimes more, parietal
teeth becoming smaller posteriorly; outer lip
sharp, weakly reflected, commonly with one
or more inner tubercles. Protoconch smooth,
large, with one and one-half protruding
whorls, leaving umbilicus-like slit in apex of
shell (Figs. 76, 77).

Radula with base of central tooth wide,
emarginate half of its length; crown of mar-
ginal teeth pointing medially, mesocone
stronger than endocone.

WESTERN ATLANTIC ELLOBIIDAE 193

Animal grayish-white; neck and tentacles
sometimes darkly pigmented. Hermaphro-
ditic duct convolute; pallial gonoduct her-
maphroditic as far as the vaginal арейиге;
anterior mucous gland and prostate gland
cover entire length of spermoviduct; bursa
duct emptying near vaginal aperture; penis
short, thick; vas deferens adhering to penis.
Ganglia of visceral nerve ring widely spaced;
osphradial ganglion present.

Remarks: The majority of modern literature
has treated Myosotella Monterosato, 1906,
as a subgenus of Ovatella Bivona, 1832. The
anatomy of the type species of Myosotella,
Myosotella myosotis (Draparnaud), has been
studied extensively (Meyer, 1955; Morton,
1955b) and Giusti (1973) looked briefly into
the anatomy of the type species of Ovatella,
Ovatella firminii (Payraudeau, 1826). A study
of the anatomy of Ovatella aequalis (Lowe,
1832) from the Azores (Martins, personal ob-
servation) revealed the presence of a pallial
gland, not noted by Giusti (1973) for Ovatella
firminii, similar to that in Carychium tridenta-
tum (Müller) (Morton, 1955b), Руа scara-
beus (Gmelin, 1791) (Plate, 1897), Cassidula
labrella (Deshayes, 1830) (Renault, 1966) and
Laemodonta cubensis (Pfeiffer, 1854) (Mar-
tins, this study). In another work (Martins,
1980) Ovatella aequalis was shown to have
a tripartite mandible with tapering ends,
whereas that of Myosotella myosotis 1$ entire
and quadrangular. These two characteristics,
corroborated by differences in the proto-
conch, justify the attribution of generic rank
to Myosotella.

Some modern authors, following Kennard
8 Woodward (1919), treat Myosotella as a
junior synonym of Gray’s misspelling “Phy-
па’ (Morrison, 1951а; M. Smith, 1951; Мс-
Millan, 1968; Keen, 1971; Climo, 1982). The
word “Phytia” appeared in Gray (1821) and is
obviously a misspelling of Pythia Róding,
1798, for two reasons. First, as Watson
(1943) pointed out, the family Ellobiidae was,
at the time of Gray’s publication, divided into
very few genera, and Carychium Müller,
1774, Рута Roding, 1798, and Auricula La-
marck, 1799, all had been established many
years earlier. Pythia had been introduced for
Pythia helicina Róding [= Helix scarabeus
Gmelin], a species which has a row of tuber-
cles inside the outer lip. Group b of Gray's
“Order 1. Adelopneumona” included the am-
phibious Auricula, Carychium and “Phytia.”
Gray's only example of “Ррува’” was Voluta

denticulata Montagu, a form of Myosotella
myosotis that also has two or more tubercles
inside the ощег lip. It can be assumed, there-
fore, that Gray was including Voluta denticu-
lata Montagu within the already known genus
Pythia Róding on the basis of the dentition of
the outer lip. Second, Gray's publication is
notorious for the number of misspellings it
contains. For example, in the first nine lines
of page 231, on which “Phytia” appears in
the fifth line, one can read: Clauselia [= Clau-
silia], Ancillus [= Ancilus] and Phaneropneu-
mana [= Phaneropneumona] and, near the
bottom of the page, Neritino [= Neritina]. Fur-
thermore, Gray (1847a) corrected “Phytia” to
Pythia. In view of the above, “Phytia” of Gray
must be treated as a misspelling of Pythia
Rôding, in accordance with Articles 19 and
32 ii of the ICZN, and as such it lacks taxo-
nomic standing. Сгау’$ misspelling was later
misspelled by Blainville (1824) and Deshayes
(1832).

Gray (1847a) also introduced Alexia for Vo-
luta denticulata Montagu [= Аипсша myoso-
tis Draparnaud]. Stephens (1835) had used
the same name for a genus of Coleoptera,
however, rendering Gray's name preoccu-
pied. This fact prompted Wenz (1920) to pro-
pose Nealexia as a new name for Alexia Gray,
but Myosotella Monterosato, 1906, has pre-
cedence over Wenz' name.

In two more instances Gray made mistakes
concerning Ovatella [= sensu Myosotella]. In
1840 he used Ovatella Bivona as a subgenus
of Conovulus Lamarck for Voluta denticulata
Montagu; later (1847a) he included ‘‘Ovatella
Gray non Bivona” in the synonymy of his A/-
exia. Because Voluta denticulata was not in-
cluded in Bivona's (1832) original species,
“Ovatella 'Bivona' Gray” must be treated as
a misuse of Ovatella Bivona. Gray (1847a)
also included in the synonymy of his Alexia
the name Jaminia Brown, 1827, but the latter
name was preoccupied by Jaminia Risso
(1826).

Pallary (1900) proposed Kochia as a sub-
genus of A/exia Gray and he included, among
other species, Alexia (K.) denticulata (Mon-
tagu) and Alexia (K.) oranica Pallary [both
junior synonyms of Myosotella myosotis
(Draparnaud)]. The latter species was se-
lected as type species by Monterosato
(1906). Pallary (1921), unaware of Montero-
sato’s selection, proposed Alexia (K.) dentic-
ulata as the type species of Kochia, noting at
the same time that this name was preoccu-
pied by Kochia Frech (1891).

194 MARTINS

Monterosato (1906) considered Montagu's
Voluta denticulata and Draparnaud's Auricula
myosotis not only as being different species,
but as belonging to different genera. Leaving
the former within Gray's Alexia, he included
the latter within his genus Myosotella, which
he created for a group of species under
Pfeiffer’s Alexia #2 (1856a: 147); he desig-
nated Myosotella payraudeaui ('“Shuttle-
worth” Pfeiffer, 1856) as the type species. On
the basis of Pfeiffer's description, | consider
Myosotella раугаиаеаш conspecific with My-
osotella myosotis (Draparnaud). Monterosa-
to's name, then, 1$ the earliest available name
for the subgenus that includes Myosotella
myosotis (Draparnaud).

Habitat: Myosotella lives mainly above the
high-tide mark, sometimes even away from
the influence of spring tides (Morton, 1955b).

Range: Although it has a worldwide distribu-
tion Myosotella is generally absent from the
tropics.

Myosotella myosotis (Draparnaud, 1801)
Figs. 41-84

Auricula myosotis Draparnaud, 1801: 53
[Mediterranean coast; type probably in
Vienna (Locard, 1895)]; Draparnaud,
1805: 56, pl. 3, figs. 16, 17; Férussac,
1821: 103; Lamarck, 1822, 6: 140; Blain-
ville, 1824: 246; Blainville, 1825: 453, pl.
37 bis, fig. 6; Gould, 1833: 67; Griffith &
Pidgeon, 1834: 36; Küster, 1844: 19, pl.
1, figs. 15-17; Moquin-Tandon, 1851:
348-351 [апаюту].

Voluta denticulata Montagu, 1803: 234, pl.
20, fig. 5 [Devon, England; lectotype
herein selected RAMM 4100 (Fig. 41);
paralectotypes RAMM 4100]; Dillwyn,
1817: 506; Wood, 1825: 90, pl. 19, fig.
18.

Voluta ringens Turton, 1819: 250 [England;
lectotype herein selected USNM 85901 1
(Fig. 42); paralectotype USNM 55351].

Voluta reflexa Turton, 1819: 251 [Exmouth,
England; holotype USNM 55370 (Fig.
44)].

Phytia denticulata (Montagu). Gray, 1821:
132; Gardiner, 1923: 64; Germain, 1931:
561, text fig. 597.

Auricula veneta Von Martens, 1824: 433
[Venice: location of type unknown (fide
Cesari, 1976)].

Jaminia denticulata (Montagu). Brown, 1827,
ВТ. tig. 6.

Jaminia quinquedens Brown 1827, pl. 51, fig.
11 [Prestonpans, England; type probably
at Manchester (Sherborn, 1940)].

Acteon denticulatus (Montagu). Fleming,
1828: 337.

Auricula tenella Menke, 1828: 36 [Type local-
ity herein designated to be Norderney Is-
land; location of type unknown]; Menke,
1830: 131; Küster, 1844: 57.

Carychium personatum Michaud, 1831: 73,
pl. 15, figs. 42, 43 [Bretagne, France;
lectotype herein selected MNHNP (Fig.
45)].

Melampus borealis Conrad, 1832: 345 [New-
port, Rhode Island; type material pre-
sumed lost (Baker, 1964)]; Jay, 1839: 59;
Н. & A. Adams, 1854: 10.

Melampus gracilis Lowe, 1832: 288 [Madeira;
location of type unknown].

Auricula myosotis Lamarck. Orbigny, 1835:
28:

Руша denticulata (Montagu) Gray. Beck,
1837: 103:

Руа myosotis (Draparnaud). Beck, 1837:
104.

Auricula reflexilabris Orbigny, 1837: 326, pl.
42, figs. 1-3 [Lima, Peru; lectotype
herein selected ВММН 1854.12.4.242
(Fig. 46)].

Auricula (Auricula) myosotis (Draparnaud).
Anton, 1839: 48.

Auricula denticulata (Montagu). Gould, 1841:
199, fig. 129; De Kay, 1843: 58, pl. 5, fig.
93; Küster, 1844: 54, pl. 8, figs. 1-5;
Reeve, 1877, pl. 7, fig. 61.

Auricula mysotis Draparnaud. Sowerby,
1842: 99 [misspelling of myosotis].
Auricula denticulata var. borealis (Conrad).

De Kay, 1843: 58, pl. 5. fig. 91.

?Auricula sayi Küster, 1844: 42, pl. 6, figs. 14,
15 [United States of America; location of
type unknown (nomen dubium)].

Auricula microstoma Küster, 1844: 52, pl. 1,
figs. 18, 19 [Budua, Dalmatia; location of
type unknown].

Auricula kutschigiana Küster, 1844: 54, pl. 8,
figs. 11-14 [Servola near Trieste; Lissa
Island; location of type unknown].

Auricula biasolettiana Kúster, 1844: 56, pl. 8,
figs. 18-20 [Niza; Trieste; coast of Dal-
пана; location of type unknown].

Auricula myosotis var. elongata Küster, 1844:
69, pl. 8, figs. 21, 22 [Zara; location of
type unknown].

Auricula myosotis var. adriatica Kúster, 1844:
69, pl. 8, figs. 23, 24 [Trieste; Istria; Dal-
matia; Zara; location of type unknown].

WESTERN ATLANTIC ELLOBIIDAE 195

Auricula ciliata Morelet, 1845: 77, pl. 7, fig. 4
[Alcácer do Sal, Alentejo, Portugal; lec-
totype herein selected BMNH
1893.2.4.831 (Fig. 47)].

Аипсша botteriana Philippi, 1846: 97 [Lesina
Island, Dalmatia; location of type un-
known].

Melampus denticulatus auct.
185152:

Alexia denticulata (Montagu). Leach, 1852:
97; Locard, 1882: 182; Adam, 1947: 39;
Sevo, 1974: 5, fig. 5.

Alexia obsoleta Pfeiffer, 1854a: 111 [Tergesti,
Adriatic Sea; location of type unknown];
Kobelt, 1898: 131, pl. 19, figs. 5, 6.

Alexia myosotis (Draparnaud). Pfeiffer,
1854b: 151; Pfeiffer, 1856a: 148; Binney,
1859; 172, pl..15, 119.88; plu 79, Tig: 16;
Binney, 1860: 4; Binney, 1865: 4, figs.
2-4; Tryon, 1866: 6, pl. 18, figs. 1, 2;
Pfeiffer, 1876: 365; Nevill, 1879: 227;
Verrill, 1880: 250; Locard, 1882: 183;
Apgar, 1891: 130; Schneider, 1892: 116;
Whiteaves, 1901: 208; C.W. Johnson,
1915: 178; Morse, 1921: 21, pl. 7, fig. 44;
Nobre, 1930: 165, pl. 7, fig. 70; Nobre,
1940: 36; Adam, 1947: 38; La Rocque,
1953: 262; Porter, 1974: 300; Sevo,
1974: 6, fig. 6.

Conovulus denticulatus (Montagu). Clark,
1855: 297.

Alexia bermudensis H. 8 A. Adams, 1855a:
33 [Bermuda; lectotype herein selected
BMNH 1969105 (Fig. 48)]: H. & A. Ad-
ams, 1855b: 241; Pfeiffer, 1856a: 152;
Pfeiffer, 1876: 367; Kobelt, 1901: 282, pl.
33, fig. 3; Fénaux, 1939: 43, pl. 1, fig. 6.

Conovulus (Alexia) denticulata (Montagu).
Woodward, 1856: 174.

Alexia payraudeaui “Shuttleworth” Pfeiffer,
1856a: 147 [Corsica; Nizza; Tergesti; lo-
cation of type unkown]; Pfeiffer, 1876:
365; Kobelt, 1898; 130, pl. 17, figs. 21,
22:

Melampus turritus (Say MS) Binney, 1859:

_ 174 [Rhode Island; type presumably de-
posited at ANSP, probably lost].

Auricula bicolor Morelet, 1860: 206, pl. 5, fig.
7 [Pico, Azores; lectotype herein se-
lected BMNH 1893.2.4.822 (Fig. 49)].

Auricula vespertina Morelet, 1860: 210, pl. 5,
fig. 9 [Pico, Azores; lectotype herein se-
lected ВММН 1893.2.4.825 (Fig. 50)].

Alexia micheli Bourguignat, 1864: 140, pl. 8,
figs. 34-36 [La Calle and Cherchell, Al-
geria; lectotype herein selected MHNG
(Fig. 51)]. Non Mittré, 1841.

Stimpson,

Alexia micheli var. triplicata Bourguignat,
1864; 141, pl. 8, figs. 37, 38 [La Calle,
Algeria; lectotype herein selected MHNG
(Fig. 52)].

Alexia algerica Bourguignat, 1864: 141, pl. 8,
figs. 23-26 [Algeria; lectotype herein se-
lected MHNG (Fig. 53)]; Kobelt, 1898:
128, pl. 17, figs. 18, 19.

Alexia algerica var. quadriplicata Bourguig-
nat, 1864: 142, pl. 8, figs. 27-30 [Algeria;
lectotype herein selected MHNG (Fig.
54)].

Alexia loweana Pfeiffer, 1866: 145 [Madeira
Island; location of type unknown].

Melampus myosotis (Draparnaud). Jeffreys,
1869: 106, pl. 4, fig. 2 [Voluta ringens
Turton illustrated (Fig. 43), probably type
material].

Alexia setifer Cooper, 1872: 153, pl. 3, figs.
A1-A3, А5-Аб [San Francisco Bay,
California; holotype ANSP 22513a (Fig.
55).

Alexia setifer var. tenuis Cooper, 1872: 154,
pl. 3, fig. A4 [San Francisco Bay, Califor-
nia; holotype ANSP 22513b (Fig. 56)].

Alexia (Auricula) myosotis var. hiriarti Follin &
Bérillon, 1874: 88 [Biarritz lighthouse;
lectotype herein selected MNHNP (Fig.
57).

Alexia setigera Cooper. Pfeiffer, 1876: 368;
Fénaux, 1939: 43 [error for setifer].
Auricula (Alexia) meridionalis Brazier, 1877:
26 [Port Adelaide, South Australia; holo-

type ANSP 22506a (Fig. 58)].

Auricula watsoni Wollaston, 1878: 269 [Ma-
deira; lectotype herein selected BMNH
1895.2.2.411 (Fig. 59)].

Auricula watsoni scrobiculata Wollaston,
1878: 269 [Salvages Islands (Madeira);
lectotype herein selected BMNH 1895.
2.2.417 (Fig. 60)].

Auricula bicolor var. subarmata Wollaston,
1878: 466 [Lanzarote (Canary Islands);
location of type unknown].

Auricula (Alexia) denticulata (Montagu). Fisch-
er, 1878: 309-312.

Alexia setifera Cooper. Nevill, 1879: 226 [un-
justified emendation of setifer].

Alexia borealis Say Cooper. Nevill, 1879: 227.

Alexia hiriarti Follin 8 Bérillon. Locard, 1882:
183.

Alexia biasoletina (Küster). Locard, 1882: 183
[misspelling of biasolettiana].

Alexia ciliata (Morelet). Locard, 1882: 184;
Kobelt, 1898: 129, pl. 17, fig. 20.

Tralia (Alexia) myosotis (Draparnaud). Dall,
1885: 277; Dall, 1889: 92, pl. 52, fig. 9.

196 MARTINS

Tralia (Alexia) myosotis var. ringens (Turton).
Dall, 1885: 278.

Tralia (Alexia) myosotis forma junior Dall,
1885: 278 [new name for Auricula ciliata
Morelet and Alexia setifer Cooper].

Alexia cossoni Letour neux 8 Bourguignat,
1887: 130 [Gabès and Cheiba, Cape
Bon, Tunisia; lectotype herein selected
MHNG (Fig. 61).

Alexia terrestris Letourneux & Bourguignat,
1887: 130 [El-Hamma, $ of Gabes, Tu-
nisia; holotype MHNG (Fig. 62)].

Alexia globulus Bourguignat, in Letourneux &
Bourguignat, 1887: 131 [Gabes, Tunisia;
holotype MHNG (Fig. 63); on museum la-
bel as Alexia ovum Bourguignat].

Alexia letourneuxi Bourguignat, т Letourneux
8 Bourguignat, 1887: 131 [Mandara,
near Alexandria, Egypt, and Djerba Is-
land, Tunisia; lectotype herein selected
MHNG (Fig. 64)].

Alexia pechaudi Bourguignat, ín Letourneux
8 Bourguignat, 1887: 132 [Macta near
Oran and Majerda, Tunisia; holotype
MHNG (Fig. 65)].

Alexia acuminata Morelet, 1889: 15, pl. 1, fig.
11 [Port Elizabeth, Cape Colony, South
Africa; specimen marked “type” broken,
lectotype herein selected BMNH
1893.2.4.838 (Fig. 66)].

Alexia pulchella Morelet, 1889: 15, pl. 1, fig.
10 [Port Elizabeth, Cape Colony, South
Africa; lectotype herein selected ВММН
1911.8.8.39 (Fig. 67)].

Alexia armoricana Locard, 1891: 132 [west
coast of France; lectotype herein se-
lected MNHNP (Fig. 68)].

Alexia exilis Locard, 1893: 62 [Le Croisic,
Loire-Inférieure; Porquerolles (France);
herein restricted to Porquerolles; lecto-
type herein selected MNHNP (Fig. 69)].

Alexia parva Locard, 1893: 62 [Le Croisic,
Loire-Inférieure (France); lectotype here-
in selected MNHNP (Fig. 70)].

Alexia ringicula Locard, 1893: 62 [Arrdudon,
Morbihan (France); lectotype herein se-
lected MNHNP (Fig. 71)].

Auricula (Alexia) myosotis
Pelseneer, 1894a: 73, figs.
[anatomy].

Alexia bicolor (Morelet). Kobelt, 1898: 134, pl.
24, fig. 3.

Alexia vespertina (Morelet). Kobelt, 1898:
135, pl. 24, fig. 4.

Alexia (Kochia) oranica Pallary, 1900: 240, pl.
6, figs, 2, 2a [Oran, Tunisia; lectotype
herein selected MNHNP (Fig. 72)].

Draparnaud.
195-208

Alexia myosotis marylandica Pilsbry, 1900a:
40 [Mouth of St. Leonards Creek, Patux-
ent River, Maryland; lectotype by Baker
(1964) ANSP 22483a (Fig. 73)]; C.W.
Johnson, 1934: 159.

Alexia myosotis bermudensis Pfeiffer. Pilsbry,
1900b: 504.

Alexia oranica Pallary. Kobelt, 1901: 280, pl.
31, figs. 8, 9.

Alexia bidentata Montagu forma americana
Kobelt, 1901: 312, pl. 33, figs. 1, 2 [Ber-
muda; type Senckenberg Museum,
Frankfurt-am-Main (not seen)].

Myosotella myosotis (Draparnaud). Montero-
sato, 1906: 126.

Phytia myosotis var. bermudensis (H. & A.
Adams). Peile, 1926: 88.

Phytia myosotis (Draparnaud). Ellis, 1926: 96,
pl. 2, fig. 3, pl. 5, fig. 49; Germain, 1931:
560, text figs. 295, 296, pl. 18, figs. 535,
536; McMillan, 1947: 264; McMillan,
1949: 67; M. Smith, 1951: 145, pl. 55, fig.
3, pl. 71, fig. 9; McMillan, 1968: 165;
Climo, 1982: 43-48, fig. 1, A-L.

Alexia (Myosotella) myosotis (Draparnaud).
Thiele, 1931: 466.

Phytia myosotis myosotis (Draparnaud).
Winckworth, 1932: 238.
Phytia myosotis denticulata (Montagu).

Winckworth, 1932: 238.

Alexia myosotis myosotis (Draparnaud). C.W.
Johnson, 1934: 159.

Alexia myosotis var. varicosa Fénaux, 1939:
44, pl. 1, fig. 3 [Provence, France; type
probably in Fenaux’s collection, Ecole
des Mines, Paris].

Alexia subflava Fénaux, 1939: 45, pl. 1, fig. 9
[Bermuda; type in Fénaux's collection,
Ecole des Mines, Paris (not seen)].

Phytia bermudensis (H. & A. Adams). Morri-
son, 1951b: 10.

Phytia myosotis marylandica (Pilsbry). Morri-
son, 1951b: 10; Burch, 1960a: 182
[chromosomes].

Phytia myosotis borealis (Conrad). Morrison,
1951b: 10.

Ovatella myosotis (Draparnaud). Meyer,
1955: 1-43, pls. 1, 2 [anatomy, taxon-
omy, life history]; Morton, 1955b: 119-
131, figs. [anatomy, life history]; Morton,
1955c: 127-168 [anatomy, taxonomy,
evolutionary relationships]; Bousfield,
1960: 14, pl. 1, fig. 10; Coomans, 1962:
90; Kensler, 1967: 391-406 [ecology];
Jacobson & Emerson, 1971: 65, text fig.;
Baranowski, 1971: 143; Abbott, 1974:
334, fig. 4103; Emerson 8 Jacobson,

WESTERN ATLANTIC ELLOBIIDAE 197

1976: 192, pl. 26, fig. 28; Hubendick,
1978: 1-45 [taxonomic relationships);
Morrell, 1980: 208-209; Rehder, 1981:
650, fig. 232; Jensen 4 Clark, 1986: 458,
figured.

Ovatella (Myosotella) myosotis (Draparnaud).
Zilch, 1959: 73, fig. 236; Cesari, 1973:
181-210 [taxonomy, distribution, ecol-
ogy]; Giusti, 1973: 124, figs. 4 A-N, pl. 2,
figs. 1-4, pl. 3, figs. 1-3; Giusti, 1976;
Cesari, 1976: 3-19, 5 pls. [taxonomy,
anatomy, polymorphism]; Martins, 1978:
24, pl. 3, figs. 4, 4a, 4b, pl. 4, figs. 4, 4a,
4b, pl. 5, figs. 5, 6, D; Martins, 1980:
1-24, pl. 2, figs. f-o [habitat].

Ovatella (Alexia) myosotis (Draparnaud). Rus-
sell-Hunter & Brown, 1964: 134.

Ovatella myosotis bermudensis (H. 8 A. Ad-
ams). Abbott, 1974: 334 [fig. 4105, erro-
neously stated to be Microtralia occiden-
talis, appears to be Myosotella myosotis
from Bermuda].

Description: Shell (Figs. 41-77) to 12 mm
long, oval-elongate, fragile to somewhat
solid, commonly pale yellow to purplish red,
rarely whitish. Spire high, with as many as
eight somewhat convex whorls; first three
whorls of teleoconch with spiral rows of pits,
becoming fewer as spire progresses (Figs.
76, 77); row of hairs, in juveniles, anterior to
spiral rows of pits (Fig. 77). Body whorl about
70% of shell length, smooth except for faint,
irregularly spaced growth lines. Aperture
about 80% body whorl length, oval-elongate,
anteriorly rounded; inner lip with small, very
oblique and somewhat twisted, white col-

umellar tooth; anterior parietal tooth stron-
gest, white, of variable thickness and perpen-
dicular to columellar axis; usually one, rarely
none, sometimes as many as four posterior
parietal teeth that gradually become smaller
posteriorly; outer lip sharp, often weakly re-
flected in gerontic specimens, commonly
with one, sometimes with as many as six
whitish tubercles. Partition of inner whorls
only in body whorl (Fig. 75). Protoconch as in
genus.

Animal grayish white to yellowish brown;
neck usually darkly pigmented; tentacles
subcylindric, darker than neck; rudimentary
anterior tentacles present; foot not trans-
versely divided, yellowish; mantle skirt gray-
ish with dark spots.

Radula (Figs. 78-80) having formula (20 +
11 +1+ 11 +20) x 80. Width of central tooth
base twice that of lateral teeth, with central
emargination, anterior portion of arms some-
what sinuous; crown small, posteriorly de-
pressed, unicuspid; mesocone triangular,
somewhat rounded. Lateral teeth eight to 13;
base quadrangular, elongate, oblique, with
rounded lateral prominence over anterior
third; crown cuneiform, about half length of
base, posteriorly rounded. Marginal teeth 17
to 25; base becoming reduced anteriorly,
projecting and square posteriorly; crown
pointing medially, bicuspid; endocone some-
what smaller than mesocone.

Stomach with anterior membranous cham-
ber, median muscular gizzard and posterior
membranous gastric caecum (Fig. 81).

Reproductive system (Fig. 82) with ovotes-

FIGS. 41-60. Myosotella myosotis (Draparnaud). (41) Voluta denticulata Montagu, lectotype (RAMM 4100),
Devon, England, sl 8.5 mm. (42) Voluta ringens Turton, lectotype (USNM 859011), British Isles, sl 8.4 mm.
(43) Voluta ringens Turton, figured in Jeffreys’ British Conchology, pl. 98, fig. 29 (USNM 67947), sl 8.5 mm.
(44) Voluta reflexa Turton, holotype (USNM 55370), British Isles, sl 9.2 mm. (45) Carychium personatum
Michaud, lectotype (MNHNP), Boulogne, France, sl 6.5 mm. (46) Auricula reflexilabris Orbigny, lectotype
(BMNH 1854.12.4.242), Lima, Peru, sl 9.0 mm. (47) Auricula ciliata Morelet, lectotype (BMNH 1893.2.4.831),
Portugal, sl 7.8 mm. (48) Alexia bermudensis H. & A. Adams, lectotype (ВММН 1969105), locality not given
[Bermuda], sl 7.6 mm. (49) Auricula bicolor Morelet, lectotype (BMNH 1893.2.4.822), Pico, Azores, sl 9.7
mm. (50) Auricula vespertina Morelet, lectotype (BMNH 1893.2.4.825), Area [Areia] Larga, Pico, Azores, sl
7.8 mm. (51) Alexia micheli Bourguignat, lectotype (MHNG), La Calle, Algeria, sl 9.2 mm. (52) Alexia micheli
var. triplicata Bourguignat, lectotype (MHNG), La Calle, Algeria, sl 8.0 mm. (53) Alexia algerica Bourguignat,
lectotype (MNHG), Mostaghanem, Algeria, sl 9.4 mm. (54) Alexia algerica var. quadriplicata Bourguignat,
lectotype (MHNG), Cape Caxine near Alger, Algeria, sl 6.8 mm. (55) Alexia setifer Cooper, holotype (ANSP
22513a), San Francisco, California, sl 7.1 mm. (56) Alexia setifer var. tenuis Cooper, holotype (ANSP
22513b), San Francisco, California, sl 6.4 mm; Baker (1964) gave the length as 7.7 mm, which does not
match the length of the shell marked as type. (57) Alexia (Auricula) myosotis var. hiriarti Follin & Bérillon,
lectotype (MNHNP), Biarritz lighthouse, France, sl 10.1 mm. (58) Auricula (Alexia) meridionalis Brazier,
holotype (ANSP 22506a), Port Adelaide, South Australia, sl 8.3 mm. (59) Auricula watsoni Wollaston,
lectotype (BMNH 1895.2.2.411), Madeira, sl 8.1 mm. (60) Auricula watsoni scrobiculata Wollaston, lecto-
type (BMNH 1895.2.2.417), Salvages Islands [Madeira], sl 7.5 mm.

MARTINS

FIGS. 41-60.

199

WESTERN ATLANTIC ELLOBIIDAE

FIGS. 61-77.

200 MARTINS

FIGS. 78, 79. Myosotella myosotis, radular teeth, Newport River, North Carolina, sl 5.1 mm. Scale 100 um.

tis light colored, between lobes of digestive
gland; hermaphroditic duct long, dilated,
convoluted; pallial gonoduct hermaphroditic
along its entire length; anterior mucous gland
and prostate gland cover entire length of
spermoviduct; bursa duct as long as sper-
moviduct, emptying near vaginal opening;
spermatheca spherical. Penis short, thick;

length; connectives of visceral nerve ring
long; right pleuroparietal connective twice as
long as left one; left parietovisceral connec-
tive longer than right one, sometimes with
ganglionic swelling on anterior third, from
which internal pallial nerve originates; rudi-
mentary osphradial ganglion arising from
pneumostomal nerve.

associated vas deferens adhering to penis.
Nervous system (Fig. 83) with cerebral
commissure one and one-half times width of
cerebral ganglion; left and right cerebropleu-
ral and cerebropedal connectives of same

Remarks: Myosotella myosotis is an ex-
tremely variable species especially known
from European coasts. Within one population
the shape of the shell can vary from slim and

FIGS. 61-77. Myosotella myosotis (Draparnaud). (61) Alexia cossoni Letourneux & Bourguignat, lectotype
(MHNG), Lagune de l'oued Cheiba, (Cap Bon), Tunisia, sl 7.3 mm. (62) Alexia terrestris Letourneux &
Bourguignat, holotype (MHNG), El Hamma, $ of Gabes, Tunisia, sl 5.4 mm. (63) Alexia globulus Bourguig-
nat, holotype (MHNG), Gabés, Tunisia, sl 5.7 mm. (64) Alexia letourneuxi Bourguignat, lectotype (MHNG),
Mandara, near Alexandria, Egypt, sl 5.7 mm. (65) Alexia pechaudi Bourguignat, holotype (MHNG), La
Mactra, near Oran, Tunisia, sl 5.2 mm. (66) Alexia acuminata Morelet, lectotype (BMNH 1893.2.4.838),
Natal, sl 5.0 mm. (67) Alexia pulchella Morelet, lectotype (BMNH 1911.8.8.39), Port Elizabeth, South Africa,
sl 5.0 mm. (68) Alexia armoricana Locard, lectotype (MNHNP) Brest, Finisterre, France, sl 5.1 mm. (69)
Alexia exilis Locard, lectotype (MNHNP), Porquerolles, France, sl 6.1 mm. (70) Alexia parva Locard, lecto-
type (MNHNP), Le Croisic, Loire-Inférieure, France, sl 5.0 mm. (71) Alexia ringicula Locard, lectotype
(MNHNP), Arrdudon, Morbihan, France, $1 5.0 mm. (72) Alexia (Kochia) oranica Pallary, lectotype (MNHNP),
Oran, Tunisia, $1 6.0 mm. (73) Alexia myosotis marylandica Pilsbry, lectotype (ANSP 22483a) Patuxent River,
Maryland, sl 8.0 mm. (74) Jamestown, Rhode Island, sl 6.7 mm. (75) Old Road, Shelly Bay, Bermuda, sl 6.3
mm. (76) Lateral view of spire and protoconch, Jamestown, Rhode Island. (77) Top view of spire and
protoconch, Sao Miguel, Azores. Scale 1 mm.

WESTERN ATLANTIC ELLOBIIDAE 201

il (E 10L 111 1M 2M 14M 15M

Ола

|

J

FIG. 80. Myosotella myosotis, radula, Beaufort,
North Carolina. Scale 10 um.

FIG. 81. Myosotella myosotis, stomach, Bermuda.
Scale 1 mm.

high spired to globose, and the color ranges
from pale yellow to purplish red (Martins, per-
sonal observations in Bermuda and Azores).
Similar variability occurs in the apertural mor-
phology, in which the number of parietal and
outer Пр teeth can vary considerably. It was
the variability of these characters that evoked
most of the many names given to this spe-
cies. According to Locard (1895), Drapar-
naud (1801) was aware of this variability
when he described Auricula myosotis, be-
cause the 113 syntypes included examples
of the dentate form later described by Mon-
tagu (1803) as Voluta denticulata. Michaud
(1831), who completed Draparnaud's work,
described Montagu's form as Carychium
personatum (Fig. 45).

Even quite recently the question of the con-
specificity of the European forms included in
the genus Myosotella has been extensively

FIG. 82. Myosotella туозой$, reproductive sys-
tem, Bermuda. A-C, transverse sections and their
locations. Scale 1 mm.

FIG. 83. Myosotella myosotis, central nervous sys-
tem, Bermuda. Scale 1 mm.

debated. Germain (1931) accepted two Eu-
ropean species, Phytia myosotis (Drapar-
naud), with only one posterior parietal tooth,
and Phytia denticulata (Montagu), with a
heavily dentate aperture. Winckworth (1932)
treated both as subspecies of Phytia. Watson
(1943) noted the differences between the two
forms but added that there are intermediates.
The slight differences he found in radular fea-
tures could be explained by the different sizes
of the specimens studied, and the differences
in shell morphology could be attributed to the

202 MARTINS

more saline habitat of Phytia denticulata.
Fénaux (1939), after examining hundreds of
specimens from a stretch of coast between
Toulon and Agde, southern France, found al-
most all the “species” described from Eu-
rope. Cesari (1973) was inclined to treat Ova-
tella denticulata as a synonym of Ovatella
myosotis but later (1976), as did Watson
(1943), he considered the case of Ovatella
denticulata unclear pending a definite ana-
tomical comparison. Considering the high de-
gree of shell variability of Myosotella myosotis
(sensu lato), a wide range of anatomical vari-
ability is to be expected. The same condition
is found in the Western Atlantic Melampus (M.)
bidentatus, which exhibits high variability in
shell morphology as well as in anatomical
characters (see the remarks under that spe-
cies). On the basis of the great range of vari-
ability in shell morphology, | think it justifiable
to consider Myosotella myosotis as the only
species living in Europe and North Africa. The
names Voluta ringens Turton, Voluta reflexa
Turton, Auricula tenella Menke, Carychium
personatum Michaud, Auricula botteriana
Philippi, Alexia letourneuxi Bourguignat, Al-
exia armoricana Locard, Alexia ringicula Lo-
card and Alexia oranica Pallary all pertain to
the dentate morph of Myosotella myosotis.

| have concluded previously (Martins,
1978, 1980) that Auricula vespertina Morelet
and Auricula bicolor Morelet from the Azores
are conspecific with Myosotella myosotis.
Upon inspection of the type material of Wol-
laston's Auricula watsoni and Auricula wat-
soni scrobiculata from Madeira (Figs. 59, 60)
| also include them in the synonymy of Муо-
sotella myosotis.

Shell morphology can be affected by envi-
ronmental factors. The Bermudian speci-
mens (Figs. 48, 75) are larger and thicker than
the specimens from New England (Fig. 74),
but similar to those | found in North Carolina
[Alexia myosotis marylandica Pilsbry (Fig. 73)]
and in the Azores. The thickening and en-
hanced color of the shell seen nearer the
warm regions is also observed in Melampus
(M.) bidentatus (see the remarks under the
species), and should be considered an envi-
ronmentally determined character of little
taxonomic value. The names Alexia myosotis
marylandica Pilsbry and Alexia bermudensis
H. 8 A. Adams, the latter considered a sub-
species by Abbott (1974), are obviously only
morphological variations of Myosotella myo-
$015. Alexia subflava Fénaux, also from Ber-
muda, was based upon a form with unusual

apertural features, but it is clearly within the
range of variation of Myosotella myosotis,
and it too must be considered synonymous.

Myosotella myosotis (Draparnaud) and Au-
riculinella (L.) bidentata (Montagu, 1801) are
often confused. The latter was erroneously
reported from America and Bermuda. Dall's
(1885) statement that Melampus (Leuconia)
bidentatus (Montagu) [= Auriculinella (Leuco-
phytia) bidentata (Montagu)] lived in America
was based on Binney's remarks about Myo-
sotella myosotis. Binney (1859: 174), after de-
scribing the animal, noted that it differed from
Н. 8 A. Adams’ illustration of the animal of
Alexia denticulata (1855b: pl. 82, fig. 5). He
mentioned that, “from the exterior of the an-
imal there appears no difference between it
and Melampus bidentatus.” Apparently Dall
(1885) wrongly concluded that the species in
question should also have the foot trans-
versely divided, a characteristic shared by
Melampus (M.) bidentatus and Auriculinella
(L.) bidentata, but not by Myosotella myoso-
tis. From Dall’s description of Melampus
(Leuconia) bidentatus (Montagu) it is clear
that he was confused about differences be-
tween the shell of Myosotella myosotis and
that of Auriculinella (L.) bidentata.

Kobelt (1901: 283) briefly described a sup-
posedly biplicate variation of Alexia bermu-
densis H. & A. Adams, to which he later (p.
312, caption of pl. 33, figs. 1, 2) gave the
name Alexia bidentata Montagu forma amer-
icana. The illustration hardly differs from that
of Alexia bermudensis (pl. 33, fig. 3), which H.
& A. Adams (1855a: 33) described as having
“columella biplicata” (Figs. 48, 75). As noted
above, Myosotella myosotis varies greatly in
apertural morphology, especially in the con-
spicuousness of the posterior parietal teeth.
Alexia bidentata Montagu forma americana
Kobelt is just a phenotypic variant of Myoso-
tella myosotis.

Three other names were applied to North
American specimens. Kuster (1844) de-
scribed and figured an Auricula sayi. п the
words of Binney (1859: 178), “Kúster's figure
represents no known American shell. There
exists, however, a strong resemblance be-
tween it and his figure of Alexia myosotis.”
Pfeiffer (1856a) tentatively assigned Kúster's
name to Marinula, and compared it with Au-
ricula infrequens C. B. Adams, 1852, from
Panama. After examining C. B. Adam’s type
material | disagree with Pfeiffer’s сотрап-
son. The dentition shown in Kúster's illustra-
tion resembles that of Creedonia succinea,

WESTERN ATLANTIC ELLOBIIDAE 203

although the shell is too globose and acumi-
nate to be referred with certainty to that spe-
cies. In view of the conflicting diagnostic
characters derived from the illustration and
from the description given by Kúster, | con-
sider Auricula sayi Kúster a nomen dubium.
The other two problematic names are
Melampus borealis Conrad and Melampus
turritus (Say MS) Binney, both from Rhode
Island, and both undoubtedly conspecific
with Myosotella myosotis. The former was
misidentified by Pfeiffer (1856a) who, based
upon misidentified specimens from Georgia
in the Cuming collection, wrongly assigned
them to Melampus bidentatus Say, var. y bo-
realis Conrad. Pfeiffer’s description of this
variety (1856a: 46) mentioned an “outer lip
with a white callus, regularly with 6-10 pli-
сае.” Меатриз (D.) floridanus, another
Georgian species, has as many as ten riblets
inside the outer lip, but it is doubtful that
Pfeiffer, who had introduced the latter spe-
cies 11 pages before, would have confused it
with Myosotella myosotis. Melampus (M.) bi-
dentatus normally exhibits the sort of dentic-
ulation on the outer lip mentioned by Pfeiffer,
but this feature never has been found in Му-
osotella myosotis. One must conclude that
Pfeiffer relied on misidentified specimens
when he identified his variety with Ме/атри$
borealis Conrad. The description of Меат-
pus turritus, found by Binney (1859) among
Say's unpublished manuscripts, was pub-
lished by that author only to provide addi-
tional information about Myosotella myosotis.
Myosotella myosotis can be differentiated
conchologically from Melampus (M.) bidenta-
tus and Melampus (D.) floridanus, with which
it associates, by its less globose shape, by its
shorter, anteriorly rounded and wider aper-
ture, and by its lack of riblets within the outer
lip. № differs from Creedonia succinea by its
pointed spire and by the dentition of its inner
lip, which in Creedonia has a very strong,
posteriorly located parietal tooth. Some
dwarf, thin-shelled forms of Tralia (T.) ovula
can be confused with the solid, deeply col-
ored forms of Myosotella myosotis. This fact
probably accounts for Dall's (1885) report of
the latter species from Jamaica. The nonmu-
cronate apex, the sinuous outer lip and the
white, equidistant teeth of the inner lip of Tra-
lia constitute sufficient diagnostic characters,
however. Gerontic individuals of Myosotella
myosotis have a weakly reflected outer lip, a
feature that led early authors to insist on in-
cluding this species in the genus Auricula.

Detailed studies of life history and anatomy
were published simultaneously by Morton
(1955b) and Meyer (1955).

Habitat: Myosotella myosotis lives in salt
marshes and adjacent areas, preferring piles
of rocks and detritus above the high-tide
mark. In Bermuda this species commonly
lives under piles of rocks, farther onto land
than any other halophilic ellobiid, a situation
also observed in the Azores (Martins, 1980).

Range: Myosotella myosotis is well known as
a Mediterranean and Eastern Atlantic spe-
cies. “Species” very similar to Myosotella
myosotis have been described from extra-
European shores, such as Orbigny’s (1837)
Auricula reflexilabris (Fig. 46) from the Pacific
coast of South America, Cooper's (1872) Al-
exia setifer (Figs. 55, 56) from California, Bra-
zier's (1877) Auricula (Alexia) meridionalis
(Fig. 58) from southern Australia and More-
let's (1889) Alexia acuminata (Fig. 66) and Al-
еха pulchella (Fig. 67) from South Africa.
Hanna (1939) included Cooper's species
among the “Exotic Mollusca in California”
and | concur with Paulson (1957) in conclud-
ing that the Californian Alexia setifer Cooper
does not differ from eastern American or Eu-
ropean specimens. Alexia setifer Cooper, as
well as Auricula ciliata Morelet, were named
on the basis of the presence of hairs on the
spire of juveniles. Clark (1855) first noted that
this condition occurs in Myosotella myosotis.
Taking into consideration the well-docu-
mented morphological plasticity shown by
Myosotella myosotis, | concur with Climo
(1982) in synonymizing Brazier's species, as
well as the others just mentioned.

The wide range of Myosotella myosotis is
attributed to its estuarine and supralittoral
habits; most probably the animals were car-
ried about in ballast or as egg masses laid on
deck equipment and cargo that came in di-
rect contact with marsh communities (Climo,
1982).

In the Western Atlantic this species occurs
from Halifax, Nova Scotia (Bousfield, 1960),
to Georgia, Bermuda and Cuba (Fig. 84). The
Cuban specimen at the USNM (383711)
should be classed as a spurious report until
further confirmation because it is far from the
range of the species in the Western Atlantic.
Dall's герой of this species from Jamaica 1$
doubtful and it has not been confirmed by
recent collections. Stimpson (1851), followed
by Binney (1859, 1865), Verrill (1880) and Dall
(1885), remarked that this species probably

204 MARTINS

45 30

FIG. 84. Myosotella myosotis, geographic distribu-
tion, Western Atlantic. Open circle, locality from
literature.

was introduced to the eastern coast of North
America.

Specimens Examined: MAINE: Castine
(MCZ 4180; USNM 492501); Portland (MCZ
uncatalogued; USNM 24865, 73394); New-
castle (MCZ 34005). NEW HAMPSHIRE:
Fabian Point, Great Bay, Newington (R.B.).
MASSACHUSETTS (ANSP 22508; USNM
27740, 27913): Manchester (USNM 39800);
Marblehead (MCZ 199478; USNM 492503);
Boston (MCZ uncatalogued; USNM 41240);
Woods Hole (ANSP 357609; MCZ 34004;
USNM 158953, 525155); New Bedford
(ANSP 22494; MCZ uncatalogued; USNM
139801). RHODE ISLAND (MCZ 34003;
USNM 539238): Newport (MCZ 68946,
163167; USNM 39799, 67730); Warren
(ANSP 60355); Maple Creek, Jamestown
(A.M.); Wickford (MCZ 294645). CONNECTI-
CUT: Branford (MCZ 34847; USNM 492502);
New Haven (USNM 83471). NEW YORK:
New York Harbor (USNM 492485); Cold
Spring Harbor (MCZ 294167); Staten Island
(MCZ 56738, 61847, 119477; USNM 59729,
407787, 492500). NEW JERSEY: Cape May
(MCZ uncatalogued). MARYLAND: Patuxent
River (ANSP 22483, 359154; USNM 492486);

St. Leonards Creek, Patuxent River (ANSP
60971; USNM 465806); Crisfield (USNM
618924). VIRGINIA: Mollusk (USNM 791448);
Watts Bay (USNM 701628); Chincoteague Is-
land (MCZ uncatalogued); Fisherman’s Island
(USNM 422292); Norfolk (USNM 637142).
NORTH CAROLINA: mouth of Newport River,
S of Beaufort (A.M.) SOUTH CAROLINA:
Charleston (MCZ uncatalogued); McClellan-
ville (USNM 663059). GEORGIA: Isle of Hope
(USNM 663053); Thunderbolt (USNM
663055). BERMUDA (ANSP 48594, 48595,
48596, 62743, 78217, 79937, 85588; MCZ
8972, 9971, 24407, 24408, 24409, 294163,
294166; USNM 6529a, 6537, 94436, 94437,
101401, 151271, 250298a, 492487, 492488,
482490): Hamilton (USNM 152134, 171941);
Hamilton Beach (MCZ uncatalogued); Fairy-
land (ANSP 99074; USNM 208070); Gibbet
Island (MCZ 294162); Flatts (ANSP 88572;
MCZ 294164; USNM 1719340); Shelly Bay
(MCZ 294165; USNM 492489); Old Road,
Shelly Bay (A.M.); N of Shelly Bay Beach
(A.M.); S of Coney Island (R.B.); Ferry Reach
Park (R.B.); N of Long Bird Bridge (A.M.);
Castle Harbour (ANSP 143320); Cooper’s Is-
land (ANSP 131644); Spitall Pond (A.M.);
Hungry Bay (ANSP 88580; USNM 171947;
A.M.); Paget (USNM 714209); W of Somerset
Bridge (A.M.); S of Ely’s Harbour (A.M.); Man-
grove Bay (A.M.). CUBA: Los Canos, Guan-
tanamo (USNM 383711).

Genus Laemodonta Philippi, 1846

Laemodonta Philippi, 1846: 98. Type species
by monotypy: Laemodonta [Auricula]
striata (Philippi, 1846) [= Pedipes octan-
fracta Jonas, 1845]. Non Martens, 1824,
nec Anton, 1839.

Laimodonta Bronn, 1847: 4 [nomen nudum,
fide Herrmannsen, 1852]. Non “Nuttall”
H. & A. Adams, 1855a.

Plecotrema H. & A. Adams, 1853: 120. Type
species by original designation: Ple-
cotrema typica H. & A. Adams, 1853.

Bullapex Haas, 1950: 199. Type species
by monotypy: Laemodonta (Bullapex)
cubensis (Pfeiffer, 1854).

Description: Shell to 9 mm long, oval-conic,
solid, sometimes hirsute. Umbilicus present.
Spire moderately high, sculptured with more
or less marked spiral cords. Body whorl 70-
75% shell length, sometimes truncate at
base, with same sculpture as spire. Aperture
about 70% length of body whorl, oval-elon-

WESTERN ATLANTIC ELLOBIIDAE 205

gate, narrow; inner lip with three subequal
teeth, one oblique columellar tooth and two
parietal teeth; outer lip thickened, with one to
three teeth about same size as parietal teeth.
Protoconch smooth, globose, prominent.

Remarks: Philippi (1846: 98), following the
description of his Auricula striata, noted,
“Laemodonta striata Adams (ubi?). Bronn
placed under this name this species from
Sandwich Islands [Hawaii]. The name Lae-
modonta appeared as a nomen nudum in a
sales catalogue of shells prepared by Bronn
(1847, fide Sykes, 1894), who had sent the
shells to Philippi. Although Laemodonta Phil-
ippi, 1846, was introduced in synonymy, this
name must be accepted as valid according
to the ICZN, Article 11, (а). Thiele (1931) and
Zilch (1959) used Laemodonta and since
1961 the name has been universally ac-
cepted (Clench, 1964; Franc, 1968; Abbott,
1974; Hubendick, 1978; Kay, 1979).

The names Laimodonta and Laemodonta
have been confused in many instances, the
second being taken wrongly for a misspelling
of the first. Laimodonta (Nuttall MS) H. 4 A.
Adams, 1853, was introduced for a group of
shells different from those assigned to Lae-
modonta Philippi. Often credit was given er-
roneously to Nuttall for the introduction of
Laimodonta. It appears, however, that Nuttall
never published the name (Sykes, 1894).
Nevill (1879) considered “Laimodonta Nut-
tall” [= emendation of Laemodonta Philippi]
and Laimodonta H. 8 A. Adams to be differ-
ent taxa and Ancey (1887) introduced A/-
lochroa to replace H. 8 A. Adams” suppos-
edly preoccupied name. Sykes (1894),
apparently unaware of Апсеу’$ introduction,
also stated that Laimodonta H. 8 A. Adams
was preoccupied, not by Philippi’s (1846) or
by Bronn's (1847) names, which he consid-
ered undescribed, but by Laimodon Gray,
1841, a genus of birds. Sykes proposed the
new name Enterodonta. Laimodon Gray,
1841, cannot be considered a homonym of
Laimodonta or of Laemodonta. According to
the ICZN, Art. 32 (a), in spite of the fact that
Philippi (1846) misspelled Bronn's name, his
spelling is to be considered the correct orig-
па! spelling. Laimodonta (Nuttall MS) H. & A.
Adams, although not a homonym of Philippi’s
name, has been abandoned in favor of Al-
lochroa in important malacological works
(Thiele, 1931; Zilch, 1959; Franc, 1968; Kay,
1979). A permanent solution to this problem
would be the placement of Laimodonta ‘‘Nut-

tall” H. & A. Adams on the Official List of
Rejected Names in Zoology.

Sykes (1894) and Hubendick (1956) in their
monographs on Laemodonta preferred the
name Plecotrema H. 8 A. Adams to Philippi’s
name. Sykes recorded in his synonymy Lira-
tor Beck, which he felt had not been properly
introduced. The name Lirator, indeed, was
used by Beck (1837) for an undescribed
Melampus (Lirator) multisulcatus from Opara
Island. Pfeiffer (1856a, 1876) tentatively iden-
tified Beck's species with Laemodonta striata
Philippi [= Pedipes octanfracta Jonas] and
Hubendick (1956) accepted Pfeiffer's opinion
without query. Because only circumstantial
evidence connects Beck's names to a recog-
nized species, however, one must conclude
that both Lirator and Melampus (Lirator) mul-
tisulcatus are nomina nuda.

Hubendick (1956: 111) stated that Aguayo
8 Jaume (1947) had given “strong reasons
for maintaining Plecotrema as the valid name
of the genus.” In fact, what Aguayo & Jaume
(1947: No. 132) had stated was, “in the im-
possibility of deciding about the priority of
Laemodonta [Philippi] 1847 [sic] over Laimo-
donta [Bronn] 1847, and about the validity of
Lirator [Beck], we have decided to use the
genus Plecotrema [H. 8 A. Adams] as many
modern authors do.” Later Hubendick (1978)
used Laemodonta Philippi and relegated Ple-
cotrema to synonymy. As stated above, Lae-
modonta Philippi is now universally ac-
cepted.

The genus Laemodonta has been assigned
erroneously to the subfamily Pedipedinae on
the basis of the shell. Although described as
“fairly uniform” (Hubendick, 1956: 110) or as
a “convenient group” (Sykes, 1894: 241), the
name Laemodonta is used now for a mixture
of genera. Examination of the radula of “Lae-
modonta” punctigera H. & A. Adams from
Malaysia has a very wide, rounded meso-
cone, typical of the Cassidula group, and that
species will be assigned to another genus
pending more research. H. & A. Adams
(1853: 120) had noted that Plecotrema [=
Laemodonta] was “а genus of small shells
allied to Cassidula.”

The present study of Laemodonta cuben-
sis leads me to include this West Indian spe-
cies within the Pythiinae on the basis of the
radula and the reproductive and the nervous
systems. Although preserved material of the
type species was not available for anatomical
comparisons, the remarkable resemblance of
shell morphology, especially apertural denti-

206 MARTINS

tion, of the West Indian species to Laemo-
donta octanfracta (Fig. 31) has led me to con-
clude that they are congeneric. № further
anatomical studies indicate the necessity of
taxonomic separation, the name Bullapex
Haas 15 available for Laemodonta cubensis.

| concur with Hubendick (1956) that the
subgenus Bullapex Haas cannot be justified
on the basis of shell characters alone. The
true umbilicus of the West Indian species is
often reduced to an umbilical depression,
similar to the pseudoumbilicus mentioned by
Haas (1950: 199) for Laemodonta clausa H. 8
A. Adams, 1853 [= Laemodonta octanfracta
(Jonas)]. According to Hubendick (1956: 114)
the inflated apex can be explained as an eco-
logically influenced character and as such 1$
unreliable. The apex (protoconch) of Laemo-
donta octanfracta, although not so prominent
as that of Laemodonta cubensis, appears to
be somewhat inflated (Fig. 92).

According to Sykes (1894) and Hubendick
(1956) the genus Laemodonta appeared in the
Eocene. These earlier species, unlike those
recorded from the Miocene, are smoother and
more similar to the West Indian Laemodonta
cubensis. The Miocene species have the
heavy sculpture of the Indo-Pacific group.
Hubendick concluded that the West Indian
species and the Indo-Pacific group probably
had common ancestors in the Tethys Sea.

Habitat: Because most of the data available
to me pertain to Laemodonta cubensis, de-
scription of the soft parts and comments on
the habitat are presented under that species.

Range: Hubendick (1956) noted the discon-
tinuous distribution of Laemodonta. Most of
the representatives are from the western
Indo-Pacific, with one species in the West
Indies and Bermuda. The genus is not repre-
sented in the Recent fauna of the Mediterra-
nean or Eastern Atlantic.

Laemodonta cubensis (Pfeiffer, 1854)
Figs. 85-87, 89, 90, 93-101

Plecotrema cubensis Pfeiffer, 1854b: 153
[Cárdenas, Cuba; location of type un-
known]; Pfeiffer, 1856a: 107; Pfeiffer,
1876: 348; Arango y Molina, 1880: 60;
Crosse, 1890: 259; Kobelt, 1900: 236;
Peile, 1926: 88; Aguayo & Jaume, 1947:
132; Hubendick, 1956: 111, text. fig. 1A,
pl. 23, fig. 7 [distribution].

Plecotrema cubense Pfeiffer. Sykes, 1895:
245; Pilsbry, 1900b: 504, pl. 62, fig. 11.

Laemodonta cubensis (Pfeiffer). Thiele, 1931:
464; Morrison, 1951b: 9; Morrison, 1958:
118-124 [habitat]; Abbott, 1974: 333, fig.
4101; Emerson & Jacobson, 1976: 190,
pl. 26, fig. 20; Rehder, 1981: 650, fig.
222; Jensen 8 Clark, 1986: 458, figured.

Laemodonta (Bullapex) cubensis (Pfeiffer).
Haas, 1950: 199, pl. 22, figs. 6-8; Zilch,
1959: 69, fig. 225; Clench, 1964: 123, pl.
79 [taxonomy, distribution]; Vokes 4
Vokes, 1983: 60, pl. 31, fig. 18.

Description: Shell (Figs. 85-87, 89, 90) to 3.5
mm long, oval, somewhat solid, pale yellow
to light brown, hirsute. Narrow umbilicus or
umbilical depression present. Spire moder-
ately high, whorls as many as six and one-
fourth, weakly convex, with two incised spiral
lines near suture; first whorls of teleoconch
with fine, compact spiral striae, crossed by
very fine, somewhat irregular growth lines.
Body whorl about 70% shell length, with in-
cised spiral lines. Aperture oval; inner lip with
three evenly spaced teeth; columellar tooth
oblique toward base, moderately strong; an-
terior parietal tooth smallest, oblique poste-
riorly; outer lip sharp, with two conspicuous
teeth, sometimes with one or two much
smaller tubercles posteriorly. Partition of in-
ner whorls occupying about three-quarters of
body whorl (Fig. 86). Protoconch whitish,
smooth, inflated, oblique or perpendicular to
columellar axis of teleoconch (Figs. 89, 90).

Animal whitish, translucent; tentacles long,
thin, subcylindric, translucent; foot entire,
rounded posteriorly. Pallial cavity long; kid-
ney long and thin; mantle gland curved, tu-
bular, empties near vaginal opening.

Radula (Figs. 93-97) having formula
[24 + (7 +7) + 1 +(7 + 7) + 24] x 100. Central
tooth at about same plane as lateral teeth;
base triangular, weakly emarginate anteriorly,
with lateral prominences at mid-length;
length of crown about half that of crown of
lateral teeth, unicuspid; mesocone some-
what sharp. Lateral teeth seven to ten; base
weakly bent medially at posterior third, with
lateral and medial prominences, the latter
anteriormost; crown about half the length of
the base, unicuspid, cuneiform. Transitional
teeth five to nine; base similar to that of lat-
eral teeth; crown bicuspid; endocone some-
what shorter and weaker than mesocone.
Marginal teeth 21 to 24; base becoming

WESTERN ATLANTIC ELLOBIIDAE 207

FIGS. 85-92. Laemodonta, Ovatella. (85) L. cubensis (Pfeiffer), West Summerland Key, Florida, sl 2.9 mm.
(86) L. cubensis, Grassy Key, Florida, sl 3.2 mm. (87) L. cubensis, Crawl Key, Florida, sl 0.84 mm. (88) O.
aequalis (Lowe), Sáo Miguel, Azores, sl 9.3 mm. (89) L. cubensis, lateral view of spire and protoconch,
Grassy Key, Florida. (90) L. cubensis, top view of spire and protoconch, Grassy Key, Florida. (91) O.
aequalis, top view of spire and protoconch, Säo Miguel, Azores. (92) L. octanfracta (Jonas), top view of spire
and protoconch, Hawaii. Scale 1 mm.

shorter and wider than that of lateral teeth,
developing lateral basal cusp covered by
next tooth; crown tricuspid; endocone, me-
socone and ectocone sharp, becoming
subequal, with mesocone somewhat longer
and stronger.

Digestive system with salivary glands small,

attaching posteriorly to esophagus through
large area. Stomach (Fig. 98) tripartite; ante-
rior portion corresponding to cardiac region,
thin, dilated; mid-portion very muscular, with
muscle also covering pyloric region; gastric
caecum somewhat thin, not muscular, receiv-
ing dilated, pouch-like posterior diverticulum

208 MARTINS

FIGS. 93-96. Laemodonta cubensis, radular teeth, Grassy Key, Florida, sl 3.5 mm. Scale 50 um.

C 1L2L 11L 1T 8T 1M 2M 3M 4M 11M 12M

N |

J KL)

FIG. 97. Laemodonta cubensis, radula, Grassy
Key, Florida. Scale 10 um.

anteriorly, at boundary with mid-region. Di-
gestive gland with two subequal lobes.

Reproductive system (Fig. 99) with ovotes-
tis between lobes of digestive gland; seminal
vesicle of hermaphroditic duct convoluted at
mid-length; pallial gonoduct hermaphroditic
to the vaginal aperture; anterior mucous
gland and prostate gland cover entire length
of spermoviduct; bursa duct about same
length as spermoviduct and empties just
posterior to vaginal opening; bursa spherical.
Penis short, thin; vas deferens adhering to
penis; penial retractor about as long as penis,
inserting on floor of pallial cavity.

Nervous system (Fig. 100) with cerebral
commissure short, about half width of cere-

FIG. 98. Laemodonta cubensis, stomach, Ber-
muda. Scale 1 mm.

bral ganglion; right cerebropedal and cere-
bropleural connectives two-thirds length of
left ones; left and right connectives of vis-

WESTERN ATLANTIC ELLOBIIDAE 209

FIG. 99. Laemodonta cubensis, reproductive sys-
tem, Hungry Bay, Bermuda. A-C, transverse sec-
tions and their locations. Scale 1 mm.

FIG. 100. Laemodonta cubensis, central nervous
system, Hungry Bay, Bermuda. Scale 1 mm.

ceral nerve ring equal; pleuroparietal connec-
tives very short; parietovisceral connectives
very long, the right one longer; visceral gan-
glion beneath tentacle retractor muscle.

Remarks: Laemodonta cubensis is the only
representative of the genus in the Atlantic. All
other species live in the Indo-Pacific region.
The West Indian species is somewhat iso-
lated conchologically, owing to its thinner,
much less sculptured shell. Kobelt (1900) and

Thiele (1931) were not sure whether this spe-
cies should even belong to this genus. As
stated in the remarks under the genus, this
species is included in Laemodonta because
of the great similarity of its apertural morphol-
ogy to that of the type species, Laemodonta
striata (Philippi, 1846) [= Laemodonta octan-
fracta (Jonas, 1845)]. The description of the
radula of Plecotrema clausa H. 8 A. Adams,
1853, a junior synonym of the type species,
given by Odhner (1925) is very similar to that
of Laemodonta cubensis except for the
lesser number of teeth in a row in the latter.

In the original description of Laemodonta
cubensis, Pfeiffer (1854b: 153) characterized
the shell as “hispidula” [slightly hairy], a fea-
ture also noticed by Haas (1950). A pilose
shell appears also in some Pacific species;
Garrett (1872) noted that his Plecotrema hir-
suta [= Laemodonta molinifera (H. £ A. Ad-
ams)] had short, curved hairs.

The presence of a pallial gland was some-
what unexpected in Laemodonta. This organ
of unknown function was first noticed by
Plate (1897) in Phytia scarabeus (Linnaeus)
and observed later in Carychium tridentatum
(Risso) (Morton, 1955b), in Cassidula labrella
(Deshayes) (Renault, 1966) and in Ovatella
aequalis (Lowe) (Martins, personal observa-
tion). Carychium lives inland, frequently in the
mountains and, although preferring humid
environments such as forest leaf litter, it is
obviously a terrestrial species. Pythia is also
considered a terrestrial ellobiid because it
lives in the upper fringe of mangroves. Ova-
tella aequalis lives just above the high-tide
limit and Laemodonta at or just below the
high-tide mark. Information is not available
concerning the precise habitat of Cassidula
labrella. Morton (1955b, c) advanced the hy-
pothesis that this was probably a case of par-
allel evolution in response to some environ-
mental parameter associated with terrestrial
life. According to that same author, the pos-
sible functions of the pallial gland range from
help in forming egg cases to aid in keeping
the body moist or secretion of bacteria-killing
substances as a protective device while the
animal is crawling. The presence of the pallial
gland in two supposedly marine ellobiids de-
папа$ a review of the hypotheses about the
evolution and function of this organ.

Laemodonta cubensis 1$ very distinct from
all other West Indian mollusks because of its
hirsute, oval shell and its apertural dentition.
Its protoconch and juveniles are very similar to
the protoconch and hirsute juveniles of the
Macaronesian and western European Ova-

210 MARTINS

90 75 16 OI 45 30

FIG. 101. Laemodonta cubensis, geographic dis-
tribution. Open circle, locality from literature.

tella aequalis (Lowe) (Figs. 89, 90). The
strength of the palatal tooth of the aperture
(Figs. 85, 88), the similarity of the radular teeth
and the presence о the pallial gland (personal
observations) also suggest a generic relation-
ship between Laemodonta cubensis (Pfeiffer)
and Ovatella aequalis (Lowe).

Habitat: Laemodonta cubensis lives at or just
below the high-tide mark, aggregating under
half-buried porous rocks, rotting wood and
leaves, and among the roots of propagules,
together with Pedipes, Blauneria, Microtralia
and Creedonia. lt is common, along with Pe-
dipes, in rocky areas, either under loose
stones near the sediment, or in crevices in
rock beds at about the high-tide mark. They
prefer the part of loose stones that touches
the sediment.

Range: Bermuda; Captiva Island, on the
western coast of Florida, south to the Florida
Keys; Bahamas and Cuba, Jamaica south to
Barbados; Mexico (Vokes & Vokes, 1983)
(Fig. 101).

Specimens Examined: FLORIDA: Third
Ragged Key above Sand Key (USNM
462737, 614608); Key Largo (MCZ 235475);
N of Tavernier Key, Key Largo (A.M.); S of
Ocean Drive, Plantation Key (A.M.); Indian

Key Fill, N of Indian Key Channel (A.M.); In-
dian Key (USNM 462893, 492557); Long Key
(A.M.); Grassy Key (A.M.); Crawl Key (MCZ
235469; A.M. Bonefish Key (ANSP 174978,
219861; МСУ 110178; USNM 599365);
Knight Key (A.M.); Bahia Honda Key (ANSP
104109; MCZ 235472; USNM 492464); West
Summerland Key (A.M.); Big Pine Key (ANSP
104110); Long Beach Drive and W end of Ko-
hen Avenue, both Big Pine Key (A.M.); Little
Torch Key (MCZ 235470); Big Torch Key
(A.M.); Ramrod Key (MCZ 235471); Sugarloaf
Key (ANSP 89559); Boca Chica Key (ANSP
104111; USNM 270348); Garden Key, Dry
Tortugas (USNM 492509, 492522); Seminole
Point (ANSP 105438); Captiva Island (ANSP
149410). BERMUDA (ANSP 62842, 293543;
MCZ 24229; USNM 250298): Fairyland
(ANSP 99078); N of Shelly Bay Beach (A.M.);
North of Long Bird Bridge (A.M.); near St.
Georges (ANSP 100821); Castle Harbour
(ANSP 143321); Hungry Bay (A.M.); W of
Somerset Bridge (A.M.); Ely's Harbour (A.M.);
Mangrove Bay (A.M.). BAHAMAS ISLANDS:
GRAND BAHAMA ISLAND: Hepburn Town,
Eight Mile Rock (ANSP 375427, 375455); Bell
Channel, Lucaya (ANSP 370709); Dead Mans
Reef [Sandy Bevan's Cay] (ANSP 371224);
Bahama Beach Canal (ANSP 371802); Silver
Cone Canal (ANSP 372886); North Riding
Point (ANSP 375563); West End (ANSP
368764); GREAT ABACO ISLAND: Mores Is-
land (MCZ 116720); Sand Bank, Crossing
Bay (MCZ 235474); S of Witch Point (MCZ
235478); Wilson City (MCZ 235479); North
Hawksbill Creek (ANSP 370566); ANDROS
ISLAND (ANSP 151873): Morgan's Bluff
(А.М.); South Mastic Point (A.M.); Mangrove
Cay (USNM 590609, 614605); Solomon
Pond, Mangrove Cay (USNM 614606); First
island off Mintie Bar, SE of South Bight
(USNM 590610, 614607); NEW PROVI-
DENCE ISLAND: Bar Point (A.M.); W of Rock
Point (A.M.); W of Clifton Point (A.M.); E of
Clifton Pier (A.M.); shore of Millars Road
(A.M.); Malcolm Creek (A.M.); ELEUTHERA
ISLAND: S of Rock Sound (MCZ 235473);
EXUMA CAYS: NE coast, Hog Cay (MCZ
235476), Western End, Hog Cay (MCZ
235477). CUBA (ANSP 22544): near Habana
(ANSP 130743); El Vedado, Habana (MCZ
uncatalogued). JAMAICA: Falmouth (ANSP
397272); Robin's Bay (USNM 442000). PU-
ERTO RICO: San Juan (R.B.). VIRGIN 1$-
LANDS: ST. THOMAS (USNM 6427).
LESSER ANTILLES: BARBADOS: off Laza-
reto (USNM 502107).

WESTERN ATLANTIC ELLOBIIDAE 211

Subfamily Pedipedinae
Fischer & Crosse, 1880

Pedipedinae Fischer 4 Crosse, 1880: 5.

Description: Shell to 11 mm long, globose
to elongate. Spire low to high, with as many
as six and one-half whorls. Body whorl 80-
90% of shell length. Aperture broad to nar-
row; columellar teeth one or two; parietal
teeth one or two; outer lip smooth, with one
strong tooth or with internal axial, ribbed cal-
losity (Pseudomelampus). Inner whorls re-
sorbed except in Pedipes and Creedonia.

Animal whitish; foot transversely divided
(except in Microtralia), posteriorly tapered, tip
rounded. Mantle skirt broad, fused posteri-
orly. Pallial cavity not occupying entire body
whorl; kidney white, long to broadly triangu-
lar; pneumostomal glands white and anterior
to kidney; anal gill well developed; mantle or-
gan lacking.

Radula having very variable formula. Cen-
tral tooth slightly posterior to lateral teeth,
unicuspid or tricuspid. Lateral teeth bicuspid.
Transition to marginal teeth gradual. Marginal
teeth with as many as five cusps.

Digestive system with mandible broadly
rectangular, composed of numerous longitu-
dinal fibers. Salivary glands white, small, fusi-
form. Digestive gland of two roughly equal
lobes; anterior lobe empties into crop
through wide anterior diverticulum, just be-
fore crop enters stomach; posterior lobe
empties into gastric caecum through poste-
rior diverticulum. Stomach tripartite, middle
section very muscular and with a caecum.

Reproductive system with hermaphroditic
duct not convoluted (except in Marinula s.s.),
posteriorly dilated; anterior mucous gland
and prostate gland extending over proximal
half of spermoviduct; bursa duct emptying
just posterior to female opening, at which vas
deferens separates from vagina (except in
Pseudomelampus and Leuconopsis). Penis
thick, usually with more or less developed
diverticulum, simple in Microtralia; vas defe-
rens free, enters penis apically; penial retrac-
tor short, attached to columellar muscle or to
floor of pallial cavity.

Nervous system with cerebral ganglia well
developed; cerebropedal connectives about
as long as cerebropleural connectives; pleu-
roparietal connectives and parietovisceral
connectives very short, somewhat longer in
Leuconopsis.

Remarks: Fischer & Crosse (1880) created
the subfamily Pedipedinae for Pedipes, the
only ellobiid genus then known to retain its
inner whorls. On the basis of radular charac-
ters Odhner (1925) added to the subfamily the
genera Marinula and Plecotrema |= Laemo-
donta], Thiele (1931) included Pseudomelam-
pus and Leuconopsis and Morton (1955c)
added Rangitotoa [= Microtralia]. In 1959 Zilch
transferred Rangitotoa to the Melampinae and
added Apodosis, which | consider a junior
synonym of Leuconopsis. Abbott (1974) listed
Microtralia within the Cassidulinae and erro-
neously considered Ovatella [sensu Myoso-
tella] to belong to the Pedipedinae. My anal-
ysis Of nervous and reproductive systems
leads to the inclusion of Microtralia in the Pe-
dipedinae and to the removal of Laemodonta,
Ovatella and Myosotella to the Pythiinae. A
new genus, Creedonia, is here created upon
the basis of the conchological and radular
characters of Creedonia succinea (Pfeiffer,
1854), formerly placed in the genus Marinula.
Creedonia does not resorb the inner whorls,
its central and lateral radular teeth are broad
with only a few in arow and the marginal teeth
have several endocones but lack ectocones.
The radulae in those species of Marinula stud-
ied have numerous, very long, narrow lateral
teeth and marginal teeth with one endocone
and several ectocones (Figs. 163-168).

The subfamily is best characterized ana-
tomically. The nervous and reproductive sys-
tems have very consistent patterns, whereas
the shell and radula vary somewhat. The
short connectives of the visceral ring cause
the concentration of those ganglia. This fea-
ture also exists in the Melampinae and sets
these two groups apart from the remaining
subfamilies, which have long visceral ring
connectives. The cerebral ganglia in the Pe-
dipedinae are proportionally much larger
than the other ganglia. The reproductive sys-
tem differs from that of the Melampinae by its
acinose ovotestis, its unconvoluted seminal
vesicle of the hermaphroditic duct, its longer
spermoviduct, its junction of the bursa duct
near the female opening, and its elaborate
penial complex, sometimes with a long diver-
ticulum. From the other three subfamilies it
differs by its elaborate penial complex with
free anterior vas deferens, and by its prostate
and anterior mucous glands that cover only
the posterior half of the spermoviduct.

The genus Leuconopsis deviates in some
anatomical features from the typical Pedipe-
dinae pattern in that its visceral nerve ring is

212 MARTINS

longer and its reproductive system is semi-
diaulic, the vas deferens and vagina sepa-
rating from the common spermoviduct half-
way along the pallial gonoduct. The visceral
nerve ring is not so long as that of any of the
species belonging to the Pythiinae and Ello-
biinae here studied and does not justify per
se the exclusion of Leuconopsis from the
Pedipedinae. The organization of the pallial
gonoducts of Leuconopsis resembles that of
Pseudomelampus (Martins, personal obser-
vation) in that the vas deferens separates
from the spermoviduct some distance before
reaching the female opening, giving rise to a
long, nonglandular vagina. The arrangement
of the reproductive organs in Leuconopsis is,
then, within the range of variation seen within
the Pedipedinae. Shell and radular charac-
ters also justify the inclusion of Leuconopsis
within the Pedipedinae.

The Pedipedinae are represented in the
West Indies by Pedipes, Creedonia, Leu-
conopsis and Microtralia, all of which are
readily distinguishable on conchological
characters. Pedipes 1$ globose, generally
heavily sculptured, with two strong columel-
lar teeth, a large oblique, posteriorly placed
parietal tooth and a readily visible callous
tooth on the outer lip, opposite the parietal
tooth. Creedonia has an apertural configura-
tion similar to that of Pedipes, but lacks the
tooth on the outer lip and has an elongate,
smooth and very thin shell. The minute Leu-
conopsis has an elongate, thick shell that
lacks the parietal and teeth on the outer lip.
The shell of Microtralia is very thin and trans-
parent and has a long, narrow aperture with
the outer lip smooth inside, one columellar
tooth and two anteriorly placed parietal teeth,
of which the anterior is the strongest.

Habitat: The Pedipedinae are a group of
small species that live at or below the high-
tide zone (Morton, 1955c; Martins, 1980),
sometimes reaching the low intertidal area
(Spencer, 1979). They live mainly in crevices
and under half-buried rocks. Some (Cree-
donia, Microtralia) live in the soft, black sed-
iment of the high-tide region of mangroves,
or in the spaces around the underground root
system of propagules and adult plants.

Range: Worldwide, warm and temperate ar-
eas. In the Western Atlantic the Pedipedinae
occur in Bermuda and from south Florida to
the West Indies, Central America south to

Brazil, Ascension Island and Tristan da
Cunha (Connolly, 1915).

Genus Pedipes Scopoli, 1777

Pedipes Scopoli, 1777: 392. Type species by
subsequent designation of Gray (1847a):
Pedipes afra (Gmelin, 1791) [= Pedipes
pedipes (Bruguière, 1789)].

Carassa Gistel, 1847 [1850]: 555 [substitute
name for Pedipes Scopoli].

Description: Shell to 6 mm long, globose,
solid to fragile, white to dark brown. Spire
low, with as many as five rapidly expanding
convex whorls, with incised spiral grooves.
Body whorl averaging 87% of shell length.
Aperture about 70% length of body whorl,
ovate, widely rounded at base; exposed por-
tion of columella flat and depressed, with two
strong columellar teeth, posterior one stron-
ger; parietal tooth strongest, oblique; outer
lip sharp, smooth or with thick, elevated
tooth opposite parietal tooth. Inner whorls
not resorbed. Protoconch with apex involute
in first teleoconch whorls, smooth, translu-
cent, yellowish to brown.

Animal grayish; tentacles long, pointed,
with transparent base, blackish toward tip.

Radula with 150 to 450 teeth in a row; first
30 to 40 rows without marginal teeth. Central
tooth at same level as lateral teeth; base
long, slightly indentate anteriorly, wide and
angular in first quarter, longitudinally de-
pressed in the middle, becoming narrower
posteriorly; crown as wide as base, falciform,
with long mesocone. Lateral teeth as narrow
as but longer than central tooth, laterally
compressed; base irregularly thickened lon-
gitudinally; crown falciform, laterally com-
pressed; mesocone long; endocone half
length of mesocone, pointed. Transitional
teeth four to eight, with base becoming
shorter, crown becoming wider; two or three
endocones. Marginal teeth cteniform, with
very thin, short base; wide rounded crown,
connected to base by long neck bent up-
wards; as many as six short endocones; me-
socone somewhat stronger than endocones.

Visceral mass coiled. Digestive gland yel-
lowish; posterior diverticulum becoming
pouch-like before entering anterior portion of
gastric caecum. Stomach tripartite; anterior
thin-walled, pouch-like section that receives
crop on left and empties into intestine on
right; middle thick-walled, very muscular giz-
zard; posterior membranous, extensible cae-

WESTERN ATLANTIC ELLOBIIDAE 213

cum that receives posterior diverticulum at
junction with gizzard. Ovotestis acinose, em-
bedded in posterior lobe of digestive gland;
hermaphroditic duct dilated along most of its
length; bursa spherical. Penis unevenly thick-
ened, convoluted in middle. Cerebral com-
missure long; connectives of visceral ring
very short; penial nerve originating from right
middle labial nerve.

Remarks: The name Pedipes was first used
by Adanson (1757), who gave a very detailed
and accurate description of a species from
Senegal later described as Pedipes pedipes
and accepted as the type species of the ge-
nus. Adanson, a contemporary of Linnaeus,
derived the generic group name from the
French word, pietin, meaning pedestrian,
which refers to the way in which the animal
progresses, first advancing the anterior half
of the transversely divided foot, then moving
the posterior half, seemingly advancing by
steps. Adanson’s work (1757) antedates the
starting point of zoological nomenclature and
therefore he is not credited with introducing
Pedipes. The first post-Linnaean use of Pe-
dipes was that of Scopoli (1777), who briefly
characterized Pedipes “Adanson” as having
the shell aperture diversely dentate. The only
known species, not mentioned by Scopoli,
was Adanson’s African “pietin,” introduced
by Bruguiere (1789) as Bulimus pedipes. He-
Их afra Gmelin, 1791 [= Pedipes pedipes
(Bruguiere, 1789)] was designated type spe-
cies of Pedipes by Gray (1847a).

Férussac (1821) used Pedipes in a re-
stricted sense but Blainville (1824), using
the sharp outer lip as the major diagnostic
character, included a heterogeneous as-
semblage, i.e., Conovulus [= Melampus] and
the opisthobranch genus Tornatella. Lowe
(1832), following Lamarck (1822) and Menke
(1828), treated Pedipes as a genus of the
family “Plicacea” and tried to prove that the
animal was a pectinibranch. After Crosse &
Fischer (1879) noticed that Pedipes was the
only Known ellobiid that did not resorb the
inner whorls of the shell, the genus could be
separated easily from otherwise conchologi-
cally similar groups, e.g., Marinula. In this
study the genus Creedonia too was found not
to resorb the inner whorls. The radula of Pe-
dipes too is unique because the posterior-
most 30 to 40 rows lack marginal teeth (Fig.
140).

The genus Pedipes is represented in the
Western Atlantic by two species. The larger

(6 mm) Pedipes mirabilis has a thick, globose,
generally heavily sculptured shell; the visible
part of the protoconch consists of hardly
more than one whorl and has a sinuous,
elongate apertural lip. The smaller (3.5 mm),
more elongate Pedipes ovalis has a thicker-
shelled, rock-dwelling form and a thinner-
shelled, mangrove-inhabiting form. The pro-
toconchs of both forms are identical, with
more than one and one-third whorls visible
and with round, not sinuous, apertural lips.

Habitat: The genus Pedipes lives in man-
groves near the sea, in which animals are
abundant under fallen leaves and branches
below the high-tide mark. They also live
along open rocky shores in crevices and un-
der stones frequently covered by waves at
high tide.

Range: Worldwide, warm temperate to trop-
ical regions. In the Western Atlantic they oc-
cur in Bermuda, southern Florida to Texas,
the West Indies and Central America south to
Brazil and Ascension Island.

Pedipes mirabilis (Mühlfeld, 1816)
Figs. 102-106, 108-110, 112-120

Turbo mirabilis Mühlfeld, 1816: 8, pl. 2, figs.
13a, b [Locality unknown, herein desig-
nated Cabo Rojo lighthouse, Puerto
Rico; type specimens presumed lost,
fide Clench (1964); neotype herein des-
ignated MCZ 188476a (Fig. 102)).

Pedipes mirabilis (Mühlfeld). Beck, 1837:
105; Pfeiffer, 1856a: 70; Pfeiffer 1876:
333; Mörch, 1878: 5; Nevill, 1879: 221;
Pilsbry, 1900b: 503; Dall & Simpson,
1901: 369, pl. 53, fig. 8; Aguayo &
Jaume, 1947: 218; Morrison, 1951b: 9;
Morrison, 1958: 121 [ecology]; Nowell-
Usticke, 1959; 88; Warmke & Abbott,
1961; 152, pl. 28, fig. j; Rios, 1970: 138;
Morris, 1973: 274, pl. 74, fig. 12; Rios,
1975: 159, pl. 48, fig. 767; Rosewater,
1975: 23; Emerson & Jacobson, 1976:
189, pl. 21, fig. 20; Rehder, 1981: 648,
fig. 234; Vokes & Vokes, 1983: 60, pl. 31,
fig. 16 [juvenile; not positively this spe-
cies]; Mahieu, 1984: 314 pp; Jensen &
Clark, 1986: 458, figured.

Pedipes quadridens Pfeiffer, 1840: 251
[Cuba; location of type unknown]; C. B.
Adams, 1849: 41, 42; C. B. Adams,
1851: 186; Pfeiffer, 1854b: 148; Shuttle-
worth, 1854b: 102; H. & A. Adams,

214 MARTINS

1855b: 149; Shuttleworth, 1858: 73;
Poey, 1866: 394.

Pedipes globulosus C. B. Adams, 1845: 12
[Jamaica; lectotype by Clench 4 Turner
(1950), MCZ 177347 (Fig. 103)]; Clench
& Turner, 1950: 288, pl. 49, fig. 9.

Pedipes globulsus “Petit” Pfeiffer, 1856a:
70 [Haiti; type from Cuming's collection,
not seen at ВММН]; Pfeiffer, 1876: 333.
Non “Férussac” H. & A. Adams, 1854
(nomen nudum).

Pedipes mirabilis (Mühlfeld) [in part] Arango у
Molina, 1880: 60; Dall, 1889: 92, pl. 47,
fig. 17; Crosse, 1890: 259; Kobelt, 1900:
255, pl. 24, figs. 19, 20; Maury, 1922: 54;
C. W. Johnson, 1934: 159; M. Smith,
1937: 145, pl. 55, fig. 8 [probably Pe-
dipes ovalis; pl. 67, fig. 17 is Pedipes
ovalis]; M. Smith, 1951: [same illustra-
tions as in first edition, 1937]; Clench,
1964: 119, pl. 76, figs. 1, 3, pl. 77 [fig. 2
is lectotype of Pedipes ovalis C. B. Ad-
ams; systematics, distribution]; Ап-
drews, 1971: 144, text fig. [figure prob-
ably is of Pedipes ovalis]; Abbott, 1974:
333, fig. 4096 [in part]; Andrews, 1977:
181, text fig. [figure probably is of Pe-
dipes ovalis].

Pedipes mirabilis Megerle. Peile, 1926: 88.

Description: Shell (Figs. 102-106, 108-110)
to 6 mm long, globose, very solid, white
to brown. Spire low, as many as five con-
vex whorls, sculptured with incised spiral
grooves and fine axial striae. Body whorl av-
eraging 88% of shell length, with average of
22 deeply incised spiral grooves. Sculpture
as on spire; spiral grooves sometimes subdi-
vided by fine spiral cords. Aperture about
70% of length of body whorl, widely ovate,
round to angular at base, sometimes with
weak angle at shoulder; columella flat and
weakly concave, with two strong, rounded,
subequal teeth perpendicular to columellar
axis; parietal tooth strongest, oblique and
slightly curved anteriorly; outer lip wide and
smooth in juveniles, thick and crenulated in
adults owing to grooves of body whorl; op-
posite parietal tooth one large tooth very
weakly extends inside aperture. Protoconch
with barely more than one whorl visible, ap-
ertural lip sinuous (Figs. 108-110).

Radula (Figs. 112-116) as in genus; formula
[120 + (6 + 70) + 1 + (70 + 6) + 120] x 120.

Stomach (Fig. 117) as in genus.

Reproductive system (Fig. 118) with her-
maphroditic duct with longitudinally dilated

seminal vesicle; bursa duct longer than sper-
moviduct and albumen gland conbined. Pos-
terior half of penis thicker than anterior por-
tion.

Nervous system (Fig. 119): left cerebrope-
dal and cerebropleural connectives longer
than right ones; left parietovisceral connec-
tive twice length of right one; visceral gan-
glion largest of five in visceral ring; left pleural
and left parietal ganglia smaller than right
counterparts.

Remarks: In spite of the great variability
shown by West Indian Pedipes, recent au-
thors consider all of the named forms con-
specific. According to Clench (1964) variabil-
ity in Pedipes is a result of colonization
strategy. Most colonies might have begun
from one individual or from one cluster of
eggs. Clench based this observation upon
the meager representation of Pedipes in mu-
seum collections, because he clearly stated
(р. 118), “there is nothing in the literature...
concerning their life history.” The colonies
are not so rare as Clench implied. Pedipes
species are among the most common West
Indian ellobiids just below the high-tide mark,
at least in mangroves (Martins, personal ob-
servation).

Pedipes mirabilis prefers piles of loose
rocks around the high-tide mark. The shell is
always thick, deeply grooved, with the aper-
ture constricted in adults by a thick outer lip
tooth. The body whorl of gerontic animals
shows asymmetric growth. The name
“quadridens” of Pfeiffer (1840) reflects the
change in apertural aspect with age, and the
names “globulosus” of С. В. Adams (1845)
and “globulus” of Pfeiffer (1856a) refer to the
allometric growth of this species.

In 1849 C. B. Adams cautiously introduced
Pedipes ovalis, calling attention to its strong
affinity with Pfeiffer's Pedipes quadridens. Al-
though the latter species is here considered a
junior synonym of Pedipes mirabilis, Pedipes
ovalis is recognized here as a separate spe-
cies on the basis of size, sculpture and pro-
toconch. The thick-shelled, heavily toothed
Pedipes ovalis that C. B. Adams described
from Jamaica rarely occurs in mangroves
(Martins, personal observation), where the
much thinner-shelled, smoother Pedipes tri-
dens Pfeiffer [= Pedipes ovalis C. B. Adams]
abounds. Their similar protoconchs and the
sizes suggest, however, that both rock-
dwelling and mangrove-dwelling forms are
expressions of the same species. The simi-

WESTERN ATLANTIC ELLOBIIDAE 215

FIGS. 102-111. Pedipes. (102) P. mirabilis (Mühlfeld), neotype (MCZ 188476a), Cabo Rojo lighthouse,
Puerto Rico, sl 4.7 mm. (103) P. globulosus C. B. Adams, lectotype (MCZ 177347), Jamaica, sl 4.6 mm.
(104) P. mirabilis, Puerto Cabello, Venezuela, sl 6.0 mm. (105) P. mirabilis, Rio Grande do Norte, Brazil
(ANSP 300179), sl 3.8 mm. (106) P. mirabilis, Morgan's Bluff, Andros Island, Bahamas, sl 5.0 mm. (107) P.
pedipes (Bruguiere), Senegal (AMNH 22590), sl 7.7 mm. (108) P. mirabilis, lateral view of spire and proto-
conch, Maravén, Venezuela. (110) P. mirabilis, top view of spire and protoconch, Shelly Bay, Hamilton,
Bermuda. (111) P. pedipes, top view of spire and protoconch, Sáo Miguel, Azores. Scale 1 mm.

larities of Pedipes ovalis with Pedipes mira-
bilis should, then, be interpreted as adapta-
tions for life in rocky environments. A more
detailed comparison between these two spe-
cies is presented under the remarks on Pe-
dipes ovalis.

Pedipes mirabilis is similar to the Eastern
Atlantic Pedipes pedipes (Bruguière), mostly
in the shape of the protoconch (Fig. 111).
The Eastern Atlantic species, however, has a
double outer lip tooth and a bifid, downward-
curved parietal tooth (Fig. 107).

216 MARTINS

FIGS. 112-115. Pedipes mirabilis, radular teeth. (112) Shelly Bay, Bermuda, sl 2.3 mm. (113-115) El Palito,

Venezuela, sl 3.2 mm. Scale 50 um.

CAL 9L 65L 75L80L1T 2T ST 1M

I

—~

2M 3M 4M 38M 40M 81M 124M

/

FIG. 116. Pedipes mirabilis, radula, El Palito, Ven-

ezuela. Scale 10 um.

H. & A. Adams (1854) listed a Pedipes
globulus Férussac, which might be confused
with the homonym introduced by Pfeiffer

AA

FIG. 117. Pedipes mirabilis, stomach, Bahamas.
Scale 1 mm.

WESTERN ATLANTIC ELLOBIIDAE ZY.

ot

FIG. 118. Pedipes mirabilis, reproductive system,
Clifton Pt., New Providence, Bahamas. A-C, trans-
verse sections and their locations. Scale 1 mm.

FIG. 119. Pedipes mirabilis, central nervous sys-
tem, Clifton Pt., New Providence, Bahamas. Scale
1 mm.

(1856a) for a West Indian specimen. Such a
name does not appear in Férussac (1821).
The Adams brothers might have intended to
refer to Pedipes ovulus, which Férussac
(1821: 109) described as “longer than afra [=

FIG. 120. Geographic distributions, Pedipes mira-
bilis (circles) and Leuconopsis manningi (star).
Open circle, locality from literature.

Pedipes pedipes], smooth and shiny, without
tooth on the outer lip.” Perhaps Férussac
was dealing with a specimen of Marinula,
which Connolly (1915) with doubt referred to
Marinula xanthostoma H. & A. Adams. Pe-
dipes globulus was described by Pfeiffer
(1856a) using Petit’s manuscript name in the
Cuming collection, and it is considered syn-
onymous with Pedipes mirabilis. The name
Pedipes globulus “Еегиззас” H. & A. Adams
should be considered a nomen nudum.

A Pedipes, tentatively assigned to Pedipes
mirabilis, was found in the Early Miocene
Cantaure Formation in Venezuela (Gibson-
Smith & Gibson-Smith, 1979). Recently
(1985) the Gibson-Smiths described those
fossils as Pedipes mirandus, which | consider
а junior synonym of Pedipes ovalis (see the
remarks for this species).

Habitat: Pedipes mirabilis usually lives on
rocky shores, often where wave action is
strong. The animals aggregate in fairly large
numbers under rocks at or just below the
high-tide mark.

Range: Bermuda; Florida, Texas; West In-
dies, Central America, northern South Amer-
ica to Sao Paulo, Brazil (Rios, 1975); Ascen-
sion Island (Fig. 120).

Specimens Examined: FLORIDA: Daytona
(USNM 162346, 253173); Indian River
(USNM 758222); Lake Worth (MCZ 205366;

218 MARTINS

USNM 599349); Palm Beach Inlet (MCZ
110215; USNM 543392); Boca Raton (ANSP
219865); S Bayshore Dr., Miami (USNM
701950); Biscayne Bay (MCZ 291105); Hau-
lover Beach Park, Biscayne Bay (USNM
809777). TEXAS: Port Aransas (MCZ 225522,
229626); Mustang Island, Port Aransas (MCZ
235614); South Padre Island (ANSP 319092;
USNM 758649); Port O'Connor (USNM
711183). BERMUDA (ANSP 48599, 62741;
MCZ 9952, 24251, 167937): Flatts (USNM
171948); Shelly Bay (MCZ 25523; A.M.); М of
Long Bird Bridge (A.M.); W of Somerset
Bridge (A.M.); Ireland Island (USNM 712378).
BAHAMA ISLANDS: GRAND ВАНАМА IS-
LAND: W of Eight Mile Rock (R.B.); Hepburn
Town, Eight Mile Rock (ANSP 370410); Car-
avel Beach, Freeport (ANSP 370228); Tama-
rind Shipway, Lucaya (ANSP 370708);
GREAT ABACO ISLAND: Wilson City (ANSP
299513; USNM uncatalogued); Sweeting's
Village (MCZ 24142); Sand Bank, Crossing
Bay (MCZ 116721); Mores Island (MCZ
116719); ANDROS ISLAND: Morgan's Bluff
(A.M.); South Mastic Point (A.M.); Mangrove
Cay (USNM 180462a); PARADISE ISLAND
(A.M.); NEW PROVIDENCE ISLAND: Bar
Point (A.M.); Delaport Point (A.M.); Rock
Point (A.M.); Clifton Point (A.M.); E of Clifton
Pier (A.M.); ROYAL ISLAND (MCZ 78360).
CUBA: El Vedado (MCZ 167983); Matanzas
Bay (ANSP 167481; MCZ 83308, 109334,
167984); Peñas Altas (MCZ 127866); Playa
de Bellamar (ANSP 222590, 345332); Ver-
salles (MCZ 92075); Muelle de la Aduana,
Matanzas (MCZ 188903); Chivera, Bahia de
Santiago (MCZ 167985); Cayo Francés (MCZ
167982); Guantánamo Bay (ANSP 313059).
JAMAICA (ANSP 22565, 22570, 22572; MCZ
117347, 117348, 185170; USNM 90459,
94747): Montego Bay (USNM 441609); Rob-
ins Bay (MCZ 167896; USNM 441978);
Jack's Bay (MCZ 167895; USNM 441836);
Manchioneal (USNM 492493); Port Morant
(USNM 423674); Rock Fort (MCZ 167894;
USNM 423792); Kingston (USNM 442594);
Kingston Harbor (MCZ 314005); Palisadoes
(USNM 442540); Mouth of Rio Cobre, Port
Royal (USNM 426870); Hunt's Bay (USNM
441675); Little River (USNM 492506). HAITI:
St. Louis (MCZ 167899; USNM 439397); Port
Salut (MCZ 167891; USNM 440000); Les
Cayes (USNM 439780); Ааит (USNM
367339, 440107); Baïe Anglaise, near Aquin
(USNM 439605); Saltrou (MCZ 167897,
167898, 223892; USNM 439341); W of Me-
tesignix (USNM 404730); Bizoton (USNM

439843). DOMINICAN REPUBLIC: Santo Do-
mingo (ANSP 60920; USNM 492507); Santa
Bárbara de Samaná (ANSP 173412; MCZ
57783); Cayo Chico, 4 km E of Santa Bárbara
de Samaná (MCZ 57784). PUERTO RICO: Pi-
ñones, W of Boca de Cangrejos (A.M.); Pu-
erta de Tierra, San Juan (A.M.); Punta Arenas,
N of Joyuda (A.M.); Cabo Rojo lighthouse
(MCZ 188476, 188476a); Humacao (MCZ
166297); Ensenada Honda, Culebra Island
(USNM 159675). VIRGIN ISLANDS: ST.
CROIX (USNM 621393, 706774): Christian-
sted (MCZ 188477); ST. THOMAS (ANSP
22569; USNM 119543); GUANA ISLAND:
North Beach (MCZ 89245); ST. JOHN (ANSP
22568). LESSER ANTILLES: ST. THOMAS
(MCZ 294220); ST. KITTS (MCZ 167935);
BARBUDA: Spanish Point (ANSP 353819);
GUADELOUPE (ANSP 22566; MCZ 181419);
MARTINIQUE (MCZ 167936, 294221; USNM
612694); Pointe Pie, 2.5 km S of Ste. Anne
(MCZ 248315); GRENADINES: Union, Admi-
ralty Bay, Bequia Island (MCZ 216484); BAR-
BADOS (MCZ 167900, 167939; USNM
502106); TOBAGO: Buccoo Bay (ANSP
188276); TRINIDAD (MCZ 90508): Toco
(MCZ 62326). CARIBBEAN ISLANDS: CAY-
MAN ISLANDS: Cayman Brac (MCZ 294222);
ARUBA (USNM 663655). CURAÇAO: Port
Marie & Daaibooi Baai (R.B.). COSTA RICA:
Portete (USNM 702836, 706405). PANAMA:
Toro Point, Fort Sherman (USNM 734066);
Limon Bay, inside Toro Point (USNM 732870;
R.B.); Fort Randolph (USNM 759237). CO-
LOMBIA: Sabanilla (MCZ 167890; USNM
103468, 193615). VENEZUELA: Cayo Punta
Brava (A.M.), Parque Nacional de Morrocoy,
Tucacas (A.M.); El Palito (A.M.); Puerto Ca-
bello (A.M.); Maravén, Borborata (A.M.).
BRAZIL: Praia do Forte, Natal, Rio Grande do
Norte (ANSP 300179). ATLANTIC ISLANDS:
ASCENSION ISLAND (USNM 735717).

Pedipes ovalis C. B. Adams, 1849
Figs. 121-148

Pedipes ovalis C. B. Adams, 1849: 41 [Ja-
паса; lectotype by Clench & Turner
(1950) MCZ 177349 (Fig. 121)]; C. B. Ad-
ams, 1851: 186; Pfeiffer, 1854b: 148; H.
8 A. Adams 1855b: 249; Pfeiffer, 1856a:
70; Pfeiffer, 1876: 333; Clench 4 Turner,
1950: 321, pl. 141, fig. 14 [lectotype fig-
ured]; Morrison, 1951b: 9; Morrison,
1958: 121 [ecology]; Morton, 1955:
127-168 [evolution].

Pedipes tridens Pfeiffer, 1854b: 148 [nomen
nudum].

WESTERN ATLANTIC ELLOBIIDAE 219

Pedipes tridens Pfeiffer, 1855: 122 [Bermuda
and Cärdenas, Cuba, herein restricted to
Bermuda; lectotype herein selected
BMNH 1967590 (Fig. 122)]; H. & A. Ad-
ams, 1855b: 249; Pfeiffer, 1856a: 72;
Pfeiffer, 1876: 333; Pilsbry, 1900b: 503,
pl. 62, fig. 10; Peile, 1926: 88; Haas,
1950: 198, pl. 22, fig. 4.

Pedipes naticoides Stearns, 1869: 108, text
fig. [Rocky Pt., Tampa Bay, Florida; ho-
lotype USNM 37598 (Fig. 123)]; Pfeiffer,
1876: 334; Dall, 1883: 323; Dall, 1885:
279, pl. 18, fig. 17; Simpson, 1889: 69.

Pedipes mirabilis (Mühlfeld) [in part]. Arango
y Molina, 1880: 60; Dall, 1889: 92, pl. 47,
fig. 17; Crosse, 1890: 259; Kobelt, 1900:
255, pl. 24, figs. 19, 20; Maury, 1922: 54;
C.W. Johnson, 1934: 159; M. Smith,
1937: 145, pl. 55, fig. 8 [probably Pe-
dipes ovalis; pl. 67, fig. 17 is Pedipes
ovalis]; М. Smith, 1951: [same illustra-
tions as in first edition, 1937]; Clench,
1964: 119, pl. 76, figs. 1, 3, pl. 77 [fig. 2
is lectotype of Pedipes ovalis C. B.
Adams; systematics, distribution]; An-
drews, 1971: 144, text fig. [figure prob-
ably is of Pedipes ovalis]; Abbott, 1974:
333, fig. 4096 [in part]; Andrews, 1977:
181, text fig. [figure probably is of Pe-
dipes ovalis]. Non Mühlfeld, 1816.

Pedipes insularis Haas, 1950: 197, pl. 22, fig.
3 [Lover’s Lake, St. George’s, Bermuda;
holotype FMNH 30171 (not seen); para-
type ANSP 212176 (Fig. 124)].

Pedipes mirabilis, forma ovalis C. B. Adams.
Robertson, 1960: 22.

Pedipes mirandus Gibson-Smith & Gibson-
Smith, 1985: 88, fig. 1 [Early Miocene
Cantaure Formation, Paraguana Penin-
sula, Venezuela; holotype NHMB No. H
17113 (not seen)].

Description: Shell (Figs. 121-139) to 3.5 mm
long, oval, solid to thin, yellow to dark brown;
spire low, whorls four and one-half, convex,
sculptured with incised spiral grooves; mi-
crosculpture of grooves composed of very
fine, irregular, compressed axial lamellae,
sometimes crossed by spiral lines; ribs
smoothish, sometimes with incised spiral
lines; slightly matte appearance caused by
fine growth lines crossing spiral ribs. Body
whorl averaging 85% of shell length, with 15
to 34 deeply incised spiral grooves. Aperture
about 70% of length of body whorl, widely
ovate to squarish, round to somewhat angu-
late at base; columella flat and weakly con-

cave, with two rounded teeth, the posterior
one stronger, anterior one sometimes very
weak; parietal tooth oblique, longest; outer
lip somewhat angular posteriorly, frequently
smooth in thin-shelled forms (Figs. 122-124,
126-128); thick-shelled forms usually with
strong, ridge-like tooth opposite parietal
tooth, extending inside aperture (Figs. 121,
125, 129, 132). Protoconch with more than
one and one-third whorls visible, apertural lip
round, not sinuous (Figs. 133-139).
Radula (Figs. 140-144) as in genus; for-
mula [75 + (5 + 50) + 1 + (50 + 5) + 75] x 120.
Stomach (Fig. 145) as in genus.
Reproductive system (Fig. 146) with her-
maphroditic duct anteriorly dilated to form
seminal vesicle; bursa duct shorter than sper-
moviduct and albumen gland combined. An-
terior half of penis thicker than posterior half.
Nervous system (Fig. 147) with left cere-
bropedal and cerebropleural connectives
about twice length of right ones; left parieto-
visceral connective about as long as right
one; visceral ganglion largest of five in vis-
ceral nerve ring; left pleural ganglion and left
parietal ganglion three times larger than right
counterparts.

Remarks: Pedipes ovalis is very variable
(Figs. 121-132). A stout, highly sculptured
form could be confused with Pedipes mirabi-
lis. In fact, C. B. Adams (1849: 41) introduced
his description of Pedipes ovalis with the
words, “Pedipes ovalis may be a variety of
Pedipes quadridens Pfeiffer [= Pedipes mira-
bilis (Múhltfeld)].” Аз С. В. Adams pointed
out, it differs from Pedipes mirabilis by the
smoothness of its body whorl and the less
conspicuous tooth on the outer lip. The outer
lip tooth in Pedipes ovalis is often ridge-
shaped and it gradually diminishes into the
aperture, whereas in Pedipes mirabilis this
tooth is more tubercle-shaped. In Pedipes
ovalis the anterior columellar tooth usually is
weaker than the posterior one. The most
consistent character differentiating these
species, however, is the protoconch, which
in Pedipes ovalis is larger and has a rounded,
not sinuous, lip.

Smoother, thin-shelled examples were
named Pedipes tridens by Pfeiffer (1855), Pe-
dipes naticoides by Stearns (1869) and Pe-
Чрез insularis by Haas (1950) (Figs.
122-124). This form differs from the typical
thick-shelled form in the greater number of
grooves on the body whorl and in the wider,
somewhat quadrangular aperture that some-

220 MARTINS

FIGS. 121-132. Pedipes ovalis C. B. Adams. (121) Lectotype (MCZ 177349), Jamaica, sl 3.1 mm. (122)
P. tridens Pfeiffer, lectotype (BMNH 1967590), Bermuda, sl 3.4 mm. (123) P. naticoides Stearns, holotype
(USNM 37598), Tampa Bay, Florida, sl 2.4 mm. (124) P. insularis Haas, paratype (ANSP 212176), Lover's
Lake, Bermuda, sl 2.4 mm. (125) Clifton Pt., New Providence, Bahamas, sl 2.3 mm. (126) Shore of Millars
Road, New Providence, Bahamas, sl 2.3 mm. (127) Crawl Key, Florida, sl 2.3 mm. (128) Plantation Key,
Florida, sl 3.5 mm. (129) Punta Arenas, Puerto Rico, sl 2.8 mm. (130) Isla Mujeres, Yucatán, Mexico (R.B.),
sl 2.6 mm. (131) Fort Sherman, Panama (USNM 620532), sl 3.3 mm. (132) Puerto Cabello, Venezuela, sl

3.0 mm.

times has a weak tooth inside the outer lip.
The smoothness of the body whorl is very
evident in the thin-shelled form although
there is much variability and overlap with the
thick-shelled form. Owing to unifying fea-
tures, such as the identical protoconch, and
the continuation and gradual disappearance
of the outer lip tooth into the aperture, how-
ever, the thin-shelled form should be consid-
ered conspecific with Pedipes ovalis.

The thick-shelled forms of Pedipes ovalis
live mostly in rocky habitats, whereas the
thin-shelled forms are predominantly man-
grove-dwellers. In Punta Arenas, Puerto
Rico, both species of Pedipes live in an area
in which mangrove trees cover the rocky
shore. At this site Pedipes ovalis showed a
wide range of thickness and corresponding
variability in the conspicuousness of the
tooth on the outer lip (Fig. 129). In the Florida

WESTERN ATLANTIC ELLOBIIDAE 221

FIGS. 133-139. Pedipes ovalis. (133) Juvenile, Crawl Key, Florida, sl 0.45 mm. (134) Juvenile, Crawl Key,
Florida, sl 0.55 mm. (135) Juvenile, Crawl Key, Florida, sl 0.55 mm. (136) Lateral view of spire and proto-
conch, Clifton Pt., New Providence, Bahamas. (137) Top view of spire and protoconch, Clifton Pt., New
Providence, Bahamas. (138) Top view of spire and protoconch, Punta Arenas, N of Joyuda, Puerto Rico.
(139) Top view of spire and protoconch, Isla Mujeres, Yucatán, Mexico. Scale 1 mm.

Keys, in which | failed to collect Pedipes mi-
rabilis and from which | could not confirm any
museum records referring to that species,
Pedipes ovalis in most mangroves appears
as Pfeiffer’s Pedipes tridens or Stearns’ Pe-

dipes naticoides. In rocky areas, however,
the sculpture and shape approach those of
Pedipes mirabilis.

Anatomical research yielded some small
differences in the reproductive and nervous

222 MARTINS

FIGS. 140-143. Pedipes ovalis, radular teeth. (140) Whole radula, Ely's Harbour, Bermuda, si 3.1 mm. (141)
Morgan's Bluff, Andros Island, Bahamas, sl 2.7 mm. (142, 143) Ely’s Harbour, Bermuda, sl 2.7 mm. Scale,
Fig. 140, 1 тт; all others, 50 um.

WESTERN ATLANTIC ELLOBIIDAE 223

С 1L2L3L 34L т 3T AT

MUA ASS

1M 2M

14M 60M 62M
DER ACTAS

FIG. 144. Pedipes ovalis, radula, Ely's Harbour,
Bermuda. Scale 10 um.

FIG. 145. Pedipes ovalis, stomach, Florida. Scale
1 mm.

systems, and counts of radular teeth are
lower in Pedipes ovalis. On that basis, but
mostly on the bases of the protoconch, the
generally more disparate sizes of the col-
umellar teeth, the shape of the outer lip tooth
and the maximal size, Pedipes ovalis is con-
sidered distinct from Pedipes mirabilis. The
resemblance of the two species can be inter-
preted as convergence due to adaptation to
the same environmental pressures of the
rocky shore. The gradation from the thick-
shelled, rock-dwelling forms to the thin-
shelled, mangrove-dwelling populations, to-
gether with the retention of the same pattern
of protoconch and shape of the tooth on the

ot...--

FIG. 146. Pedipes ovalis, reproductive system,
Florida. Scale 1 mm.

FIG. 147. Pedipes ovalis, central nervous system,
Florida. Scale 1 mm.

outer lip, amply justify the inclusion of Pe-
dipes tridens, Pedipes naticoides and Pe-
dipes insularis as junior synonyms of Pedipes
ovalis.

As stated under the remarks on the previ-
ous species, Gibson-Smith 8 Gibson-Smith
(1985) described a Pedipes mirandus from
the Early Miocene Cantaure Formation of
Venezuela. The authors did not mention the
shape of the protoconch, the decisive char-
acter for the separation of the Western Atlan-
tic species. Judging from the accentuated
difference in size of the columellar teeth,
however, | consider Pedipes mirandus a jun-

224 MARTINS

FIG. 148. Pedipes ovalis, geographic distribution.

ior synonym of Pedipes ovalis. The specimen
of the latter species that | collected in Vene-
zuela (Fig. 132) closely resembles the illustra-
tion of the holotype of Pedipes mirandus
(Gibson-Smith 8 Gibson-Smith, 1985: 88,

fig. 1).

Habitat: Pedipes ovalis often occurs with Pe-
dipes mirabilis under rocks and in crevices at
or just below the high-tide mark. The thinner-
shelled forms are very common in man-
groves under leaves, twigs and rocks at or
just below high-tide mark. The juveniles ven-
ture farther into the intertidal zone than do
any other West Indian ellobiid.

Range: Bermuda; Florida; West Indies; Mex-
ico south to Panama and Venezuela (Fig.
148).

Specimens Examined: FLORIDA: Waveland
(USNM 123531); Miami (ANSP 320358;
USNM 159439, 330934); Ocean Beach
(USNM 270714); Third Ragged Key above
Sand Key (USNM 462738); Key Largo (USNM
597459); Tavernier Key (USNM 492504);
Plantation Key (MCZ 188973, 291000,
291003); Ocean Dr., Plantation Key (A.M.);
Upper Matecumbe Key (USNM 492492); In-
dian Key (MCZ 167889; USNM 492520); In-
dian Key Fill, N of Indian Key Channel (A.M.);

Lignumvitae Key (ANSP 156683); Lower
Matecumbe Key (MCZ 167893; USNM
492495); Long Key (ANSP 219860; A.M.):
Grassy Key (ANSP 89560, 397279; MCZ
188970; A.M.); Crawl Key (MCZ 188972,
289998, 289999; A.M.); Bonefish Key (ANSP
227991); Knight Key (MCZ 188971); Bahia
Honda (ANSP 104115; MCZ 188969); West
Summerland Key (A.M.); Big Pine Key (ANSP
104114, 227999; MCZ 291104); W end of Ko-
hen Avenue and Long Beach Drive, both on
Big Pine Key (A.M.); Little Torch Key (MCZ
188974, 291108); Big Torch Key (ANSP
104112); Ramrod Key (MCZ 188975; USNM
599368); Sugarloaf Key (ANSP 89561,
104113; MCZ 188478); Boca Chica Key
(MCZ 167892; USNM 270349); Key West
(ANSP 22563; USNM 36017, 492494); SW
channel, Dry Tortugas (USNM 492505); Gar-
den Key, Dry Tortugas (USNM 590210); Fla-
mingo Key (ANSP 294313); Cape Sable (MCZ
291103); Seminole Point (ANSP 105432);
Sanibel Island (MCZ 84103); Tarpon Bay,
Sanibel Island (MCZ 84339); Captiva Island
(ANSP 149408); Starvation Key (ANSP
130059); Palmetto (A.M.); Mullet Key (USNM
652408, 653109; A.M.); Shell Key (USNM
466287); Tampa Bay (MCZ 239222; USNM
37598a); Anclote Key (ANSP 22564). MEX-
ICO: Isla Cancun, Quintana Roo (ANSP
285534). BERMUDA: (ANSP 48597, 48600,
48601, 48602; MCZ 9952a, 74809, 314027;
USNM 6523, 94438, 492496): Fairyland
(ANSP 99077, 111096; USNM 208071); Flatts
(USNM 171963); Shelly Bay (MCZ 225523);
Old Road, Shelly Bay (A.M.); Coney Island
(А.М.); N of Long Bird Bridge (A.M.); Nonsuch
Island (MCZ 248274); Lover's Lake (ANSP
212176); Cooper's Island (ANSP 131648);
Hungry Bay (A.M.); W of Somerset Bridge
(A.M.); Ely's Harbour (A.M.); Mangrove Bay
(A.M.). BAHAMA ISLANDS: BIMINI (ANSP
325624): East Well, East Bimini (ANSP
326449); N. end of Pigeon Cay, Bimini La-
goon (ANSP 326022; USNM 656173); S end
of Pigeon Cay (ANSP 326017); Cavelle Pond,
South Bimini (ANSP 325548); Tokas Cay
(ANSP 325831); GRAND BAHAMA ISLAND:
W of Eight Mile Rock (R.B.); Running Mon
Canal (ANSP 369780); North Hawksbill Creek
(ANSP 370569); Dead Mans Reef [Sandy Be-
van’s Cay] (ANSP 371226, 371285); Sweet-
ings Cay (ANSP 374312); Riding Point (ANSP
371521); West End (ANSP 368763, 371933);
GREAT ABACO ISLAND: West Point (ANSP
299478); Gorling Cay (ANSP 299549); AN-
DROS ISLAND: Morgan's Bluff (A.M.); South

WESTERN ATLANTIC ELLOBIIDAE 220

Mastic Point (A.M.); Danlin Bay (USNM
180671); Mangrove Cay (ANSP 325639;
USNM 180462); First island off Mintie Bar, SE
end of South Bight (USNM 271784); NEW
PROVIDENCE ISLAND: Delaporte Point
(A.M.); E of Clifton Pier (A.M.); Clifton Bluff
(MCZ 205367); Clifton Point (A.M.); Millars
Road (A.M.); Malcolm Creek (A.M.); ROYAL
ISLAND (MCZ 78360, 167901; USNM
468120); ELEUTHERA ISLAND: Governor's
Harbor (MCZ 167995); EXUMA CAYS: Hog
Cay (MCZ 225560, 225561); CAY SAL BANK:
Salt Lagoon, Cay Sal (USNM 513429). CUBA
(USNM 492498): Dimas (USNM 614603); Ha-
bana (ANSP 130744); Las Villas, Caibarién
(USNM 608763). JAMAICA (MCZ 177348a,
177349, 177350, 185170a; USNM 90460,
94748): Falmouth (ANSP 397266); Robin's
Bay (MCZ 167896a; USNM 441978a); Jack's
Bay (MCZ 167895а; USNM 441836a); Port
Morant (USNM 423674a); Palisadoes (USNM
442540a). HAITI: St. Louis (USNM 439397a);
Port Salut (USNM 440000a); Bizoton (USNM
439843a). PUERTO RICO: San Juan (R.B.);
Punta Arenas, N of Joyuda (A.M.); Cabo Rojo
lighthouse (МСУ 1884765). VIRGIN IS-
LANDS: ST. CROIX (USNM 706775); ST.
THOMAS (ANSP 22562). LESSER ANTILLES:
ST. KITTS (MCZ 167935a; USNM 492491);
GRENADA: Caliveny Harbor (ANSP 296716);
ST. MARTIN (MCZ 250474). MEXICO: Isla
Mujeres, Quintana Roo (R.B.). BELIZE: Twin
Cays (USNM 841329); Drowned Cays (ANSP
284811). PANAMA: Devil's Beach, Fort Sher-
man (USNM 620532). CARIBBEAN 15-
LANDS: ST. ANDREWS ISLAND (ANSP
155415). VENEZUELA: Puerto Cabello (A.M.).

Genus Creedonia new genus

Type species: Сгеедота succinea (Pfeif-
fer, 1854).

Description: Shell to 3.8 mm long, oval-elon-
gate, fragile. Spire moderately high, trun-
cated, with rounded apex; as many as four
smooth, weakly convex whorls. Body whorl
about 80% of shell length. Aperture oval-
elongate, about 70% of body whorl length,
posteriorly acuminate, rounded at base; col-
umella somewhat oblique and twisted; col-
umellar teeth two, posterior one stronger; pa-
rietal tooth a little stronger than posterior
columellar tooth; outer lip sharp, smooth. In-
ner whorls not resorbed. Protoconch large,
smooth, with nuclear whorls covered by first
whorls of teleoconch.

Radula with about 45 teeth in a row; central
tooth wide, with triangular base, small, uni-
cuspid crown; lateral teeth with strong en-
docone; transitional teeth with two еп-
docones; marginal teeth with as many as five
endocones.

Animal whitish; tentacles long, pointed.
Visceral mass coiled. Pallial cavity elongate;
kidney long, thin. Hermaphroditic duct some-
what dilated in the middle; penis with long
diverticulum. Nervous system with long cere-
bral commissure.

Remarks: The genus Creedonia is created
for Creedonia succinea (Pfeiffer) upon the ba-
sis of shell, radular and anatomical charac-
ters. This new genus is closely related to Pe-
dipes and Marinula, and the type species was
formerly included in one or the other genus.
Creedonia, like Pedipes, does not resorb its
inner whorls and, like Marinula, has a smooth
shell and a smooth outer lip. The three gen-
era characteristically have two columellar
teeth and one strong parietal tooth.

As stated above, Creedonia succinea for-
merly was considered to belong to the genus
Marinula. Only twice have some species of
Marinula been assigned tentatively to new
genera. Swainson (1855) introduced the ge-
nus Cremnobates in which he included his
three species Cremnobates cornea, Cremno-
bates parva and Cremnobates solida, all from
Tasmania. Hedley & Suter (1910) noted that
Cremnobates cornea is a junior synonym of
Ophicardelus australis (Quoy 8 Gaimard,
1832) and that Cremnobates solida is con-
specific with Marinula patula (Lowe, 1832).
They therefore selected Cremnobates parva
(Fig. 156) as type of the genus. Connolly
(1915) considered Cremnobates parva allied
to Marinula xanthostoma H. 8 A. Adams,
1855. Iredale (1936: 328) proposed Mar-
ipythia for Marinula xanthostoma H. & A. Ad-
ams on the basis of Connolly's opinion that
that species “could not be classed under
Marinula.” This is a misinterpretation of the
statements of Connolly (1915: 118) who, after
tracing the tortuous history of Marinula xan-
thostoma, concluded, “the typical form of
xanthostoma is on the extreme borderland of
Marinula,” but added that intermediate forms
occurred in different localities, a fact making
the connection with Marinula less doubtful.

Research on the anatomy of a Marinula cf.
xanthostoma H. 8 A. Adams, conchologically
related to Cremnobates parva, revealed a re-
productive system similar to that of Pedipes.

226 MARTINS

The reproductive system of Marinula pepita
King, 1832, the type species of the genus,
differs considerably from that of the Adams’
species, leading to the conclusion that they
are at least subgenerically separated. The
similarity of the radular teeth of Marinula
pepita to those of Marinula filholi (Hutton,
1878) (Figs. 163-168), conchologically allied
to Marinula xanthostoma, casts doubt upon
their generic separation. Because | lacked
an opportunity to examine the anatomy of
Cremnobates parva to assess its relationship
to Marinula xanthostoma, | think a decision
about the synonymy of the names proposed
by Swainson and Iredale is unwarranted.

The genus Marinula has been confused with
Ovatella [Pythiinae] on the basis of the appar-
ent similarity of the dentition of the inner lip. H.
& A. Adams (1855b) created the subgenus
Monica to include the Mediterranean Monica
firminii (Payraudeau, 1826) [= Ovatella firminii),
and the Madeiran Monica aequalis (Lowe,
1832) [= Ovatella aequalis] and Monica gracilis
(Lowe) [= Ovatella aequalis]. The shells of
Marinula are easily separated from those of
Ovatella on the basis of their apertural teeth.
Marinula, Pedipes and Creedonia all have two
conspicuous columellar teeth, whereas Ova-
tella has only one columellar tooth. The pari-
etal tooth of Marinula is the strongest of the
three inner lip teeth, whereas in Ovatella the
anterior parietal tooth is the strongest (Fig.
88). Connolly (1915) added as a diagnostic
character of the genus the absence of teeth
on the outer lip, but the Eastern Pacific
Marinula concinna (C. B. Adams, 1852) and
Marinula brevispira (Pilsbry, 1920) have a
thick, ridge-like tooth opposite the parietal
tooth. Anatomical research on these species
is needed to ascertain their phylogenetic re-
lationships, however.

Marinula is known from the Indo-Pacific
and it is well represented along the Pacific
coasts of Central and South America; it has
been reported from the South Atlantic Islands
and from South Africa as well (Connolly,
1915).

The new genus Creedonia differs from
Marinula by having a thinner, smaller shell
that is less than half the size of that of any
species included in Marinula, with the possi-
ble exception of Marinula mandroni Velain,
1877, which Connolly (1915) suspected to
have been named after a young specimen of
Marinula velaini Connolly, 1915. In Creedonia
the columella is twisted and oblique, instead
of flat and straight, and the anterior columel-
lar tooth is always conspicuous, whereas in

Marinula it is very small (Figs. 155-157). The
spire in Creedonia is more elevated, the apex
is truncate and perforated (Fig. 158) instead
of acuminate and obliterated as in Marinula
(Fig. 159). As stated above, Creedonia ani-
mals do not resorb the inner whorls of the
shell (Fig. 153), whereas those of Marinula
species do.

The radula of Creedonia succinea differs
from that of Marinula in its broad central and
lateral teeth and in the very small number of
teeth in a row (Table 3, Appendix). The mar-
ginal teeth have several endocones but no
ectocones, whereas in the Neozealandic
Marinula Вой (Hutton) and in Marinula pe-
pita King there are one or two endocones and
several ectocones (Figs. 163-168).

The genus Creedonia is named in honor of
the Rev. Joseph Dennis Creedon, Pastor of
Christ the King Church, Kingston, Rhode Is-
land, as an expression of my gratitude for his
support in this research and for his invaluable
friendship.

Creedonia succinea (Pfeiffer, 1854)
Figs. 149-154, 158, 160-162, 169-173

Leuconia succinea Pfeiffer, 1854b: 156 [Cár-
denas, Cuba; location of type unknown];
Pfeiffer, 1856a: 157; Pfeiffer, 1876: 370;
Arango y Molina, 1880: 61; Crosse,
1890: 260; H. & A. Adams, 1855b: 248.

Pedipes elongatus Dall, 1885: 279, pl. 18, fig.
4 [Marco, Florida; lectotype herein se-
lected USNM 859012 (Fig. 149); five
paralectotypes USNM 37599]; Dall,
1889: 92, pl. 47, fig. 4; Simpson, 1889:
60; Kobelt, 1900: 258, pl. 24, figs. 17, 18;
Maury, 1922: 54; C. W. Johnson, 1934:
159; M. Smith, 1937, pl. 67, fig. 4 [pl.
from Dall (1885)]; Emerson 4 Jacobson,
1976: 190; pl. 26, tig 2%

Marinula succinea (Pfeiffer). Morrison, 1951b:
9; Morrison, 1958: 118-124 [habitat];
Abbott, 1974: 333, fig. 4100 [not fig.
4108]; Vokes & Vokes, 1983: 60, pl. 31,
A

Description: Shell (Figs. 149-154, 158) to
3.8 mm long, oval-elongate, fragile, shiny,
translucent, pale yellow to golden brown.
Spire truncate, with as many as four and
one-half weakly convex, apparently smooth
whorls; very fine spiral lines visible under
high magnification, crossed by weak, irregu-
larly spaced growth lines; spiral depression
just below suture. Body whorl about 80% of
shell length, smooth. Aperture oval-elongate,

WESTERN ATLANTIC ELLOBIIDAE 227

FIGS. 149-159. Creedonia, Marinula. (149) Pedipes? elongatus Dall, lectotype (USNM 859012), Marco,
Florida, sl 3.9 mm. (150) C. succinea (Pfeiffer), Crawl Key, Florida, sl 2.3 mm. (151) C. succinea, Big Pine
Key, Florida, sl 3.3 mm. (152) C. succinea, Isla Mujeres, Yucatán, Mexico (R.B.), sl 4.3 mm. (153) С.
succinea, Isla Mujeres, Yucatán, Mexico (R.B.), sl 3.3 mm. (154) С. succinea, lateral view of spire and
protoconch, Crawl Key, Florida. (155) М. pepita King, syntype (BMNH 1968882), Chiloe Island, Chile, sl 10.1
mm. (156) М. parva (Swainson), New Zealand (USNM 98181), sl 6.4 mm. (157) M. filholi (Hutton), New
Zealand, (USNM 681303), $1 5.4 mm. (158) С. succinea, top view of spire and protoconch, Crawl Key,
Florida. (159) M. filholi, top view of spire and protoconch, New Zealand (USNM 681303). Scale 1 mm.

about 70% of length of body whorl, round at anterior columellar tooth conspicuous, pos-
base; columella somewhat oblique, twisted; terior columellar tooth twice the size of ante-
columellar teeth two, oblique toward base; rior; parietal tooth lamelliform, as large as or

228 MARTINS

somewhat larger than posterior columellar
tooth; outer lip sharp, smooth. Inner whorls
not resorbed (Fig. 153). Protoconch large,
smooth, whitish, translucent; nuclear whorls
enveloped by first whorl of teleoconch, leav-
ing pit in apex of shell (Figs. 154, 158).

Radula (Figs. 160-162, 169) with formula
[12 + (2 + 12) + 1 + (12 + 2) + 12] x 80. Base
of central tooth as wide as that of lateral
teeth, rhomboidal, with anterior end much
shorter than posterior, rounded; crown as
wide as posterior end of base; mesocone
small, triangular, with rounded tip; no ecto-
cones. Lateral teeth eight to 12; base qua-
drangular, medially bent at half-length; crown
as wide as posterior end of base, triangular,
with rounded tip; endocone about half the
length of mesocone, strong, weakly pointed.
Transitional teeth two, with base wider than
that of lateral teeth, with two subequal en-
docones. Marginal teeth 12 to 14; base be-
comes shorter and wider; mesocone be-
comes smaller as teeth approach lateral
edge of radula; first marginal tooth with three
subequal endocones; fourth endocone ap-
pears on fourth marginal tooth, fifth en-
docone on tenth marginal tooth.

Animal whitish, translucent; tentacles mod-
erately long, somewhat pointed, translucent,
with bulbous base. Foot transversely divided.
Pallial cavity elongate; kidney broad, triangu-
lar, white.

Digestive system with salivary glands
small, fusiform. Stomach globose, very mus-
cular; gastric caecum conspicuous, membra-
nous (Fig. 170). Digestive gland bilobed;
anterior lobe covers most of stomach and
empties into pouch-like posterior crop
through dilated anterior diverticulum; intes-
tine very dilated as it comes off the stomach.

Reproductive system (Fig. 171) with ovo-
testis acinose, embedded in posterior lobe
of digestive gland; hermaphroditic duct with
irregularly dilated seminal vesicle; fertilization
pouch bilobed, very conspicuous; albumen
gland large, triangular; posterior mucous
gland weakly convoluted; anterior mucous
gland and prostate gland cover posterior half
of spermoviduct. Bursa duct thick, shorter
than spermoviduct; bursa elongate. Penis
with several pouch-like dilations, with very
long diverticulum wrapped around esopha-
gus and salivary glands; short penial retractor
attaches to columellar muscle; vas deferens
short, free.

Nervous system (Fig. 172) with cerebral
commissure just shorter than width of cere-

bral ganglion; left cerebropedal and cere-
bropleural connectives shorter than right
ones; pedal commissure very conspicuous;
cerebral ganglia large, elongate laterally; left
pleural ganglion about one-fourth size of right
one; left parietal ganglion about one-tenth
size of right one; visceral ganglion largest of
visceral ring, somewhat smaller than pedal
ganglia.

Remarks: Creedonia succinea was originally
assigned by Pfeiffer (1854b) to the genus
Leuconia Gray [= Auriculinella Tausch, 1886].
The species appeared in the literature under
this name until placed by Morrison (1951b) in
the genus Marinula King, 1832, in which it
has remained until now.

Dall (1885) apparently was not aware of
Pfeiffer's species when he introduced Pe-
dipes elongatus for specimens from Marco,
Florida. Creedonia succinea is one of the few
species of ellobiids that shows little morpho-
logical variation. It cannot be confused with
any other West Indian species. The superfi-
cial similarity to the Mediterranean Ovatella
was already pointed out in the remarks under
the genus Creedonia. In Creedonia the col-
umellar tooth 1$ double and the parietal tooth
is the strongest or at least as strong as the
posterior columellar tooth. In Ovatella there is
only one columellar tooth and the first pari-
etal tooth 1$ the strongest. The same applies
to the introduced Myosotella myosotis, with
the difference that in this species the poste-
rior parietal tooth is either absent or weaker
than the anterior parietal tooth. The spire of
Creedonia succinea is truncate and the pro-
toconch gives it a mucronate appearance.
The elongate, smooth, translucent shell, with
flat whorls, separates Creedonia succinea
from the thin-shelled form of Pedipes ovalis
with which it occurs. Microtralia and Blaune-
ría also occur with Creedonia; the former
differs from Creedonia in having a narrow
aperture with much smaller inner lip teeth
and a very short spire. В/аипепа 1$ sinistral,
has a high spire and is white and transpar-
ent, whereas Creedonia is straw-colored to
brown.

Connolly (1915: 105), in his monograph on
the genus Marinula, apparently was not ac-
quainted with Pfeiffer's species. He men-
tioned “Pythia abbreviatus Beck,” criticizing
Pfeiffer's (1856a) questionable attribution of it
to Marinula in these terms: “whatever may be
its true genus, as the shell is said to come
from the Antilles it is quite unlikely to be a

WESTERN ATLANTIC ELLOBIIDAE 229

FIGS. 160-168. Creedonia, Marinula, radular teeth. (160-162) C. succinea, Long Key, Florida, sl 3.0 mm.
(163) M. filholi, New Zealand, sl 5.4 mm. (164) M. filholi, New Zealand, sl 5.5 mm. (165) M. filholi, New
Zealand, sl 5.4 mm. (166-168) M. tristanensis Connally [= M. pepita King], Gough Island (BMNH), $1 10.8
mm. Scale 50 um.

230 MARTINS

С iL 2L 11L 12L 1M 2M 3M 15M 16M

IAS ANA A

FIG. 169. Creedonia succinea, radula, Long Key,

Florida. Scale 10 um.

lee

an
‚I

]
dj

E

SA

el

FIG. 170. Creedonia succinea, stomach, Crawl
Key, Florida. Scale 1 mm.

FIG. 171. Creedonia succinea, reproductive sys-
tem, Crawl Key, Florida, sl 3.3 mm. A-C, trans-
verse sections and their locations. Scale 1 mm.

Marinula.” Beck (1837: 105) had listed, with-
out description, a “Pythia abbreviatus” from
the West Indies, placing the name after
Pythia aequalis (Lowe, 1832) [= Ovatella ae-

ppre. pipe cpe = IM a cc bg

/
` ИВ Аа и
a N \ = /
Pg. x N SS ==
` S ` >
` ` `

FIG. 172. Creedonia succinea, central nervous sys-
tem, Crawl Key, Florida, sl 3.0 mm. Scale 1 mm.

qualis] and Pythia patulus, which is question-
ably referred by Connolly (1915) to Marinula
xanthostoma H. & A. Adams. Pfeiffer (1856a)
did not see Beck's specimens but tentative-
ly assigned Pythia abbreviatus Beck to
Marinula, no doubt on the basis that Beck
listed it between two species that Pfeiffer
considered to be Marinula. The only other
species in the Western Atlantic that at first
glance could be confused with Creedonia
succinea is Myosotella myosotis, which
does not live in the West Indies. In spite of
the fact that some circumstancial evidence
seems to indicate that Beck's name refers to
Creedonia succinea, Рута abbreviatus Beck
must remain a nomen nudum.

Habitat: Individuals of Creedonia succinea
live about the high-tide mark, the juveniles
venturing a short distance into the intertidal
zone. They live within the sediment, some-
times 10 to 15 cm deep, and they occur fre-
quently under half-buried rotting wood or
rocks and on the roots of mangrove propa-
gules, together with Pedipes, Microtralia and
Blauneria.

Range: Georgia ?, Florida Keys and the Ba-
hama Islands south to Cuba and Jamaica;
Mexico (Fig. 173). The USNM record from
Isle of Hope, Georgia, collected by Hubricht,
is so distant from the normal range that it
could be explained better as the result of ac-
cidental transportation by currents.

Specimens Examined: GEORGIA: Isle of
Hope (USNM 663054). FLORIDA: S of Ocean
Drive, Plantation Key (A.M.); Lignumvitae Key

WESTERN ATLANTIC ELLOBIIDAE 231

(ANSP 156694); Long Key (A.M.); Grassy Key
(А.М.); Crawl Key (A.M.); Big Pine Key (ANSP
293553); Long Beach Drive and W of Kohen
Avenue, both Big Pine Key (A.M.); Newfound
Harbor (USNM 272639); Big Torch Key
(ANSP 104105); Sugarloaf Key (ANSP 89566,
104104); Ramrod Key (MCZ 235471a); Boca
Chica Key (USNM 590597); Key West (USNM
450693); Seminole Point (ANSP 105410);
Marco (ANSP 22578; USNM 37599, 859012);
Captiva Island (ANSP 149409); Mullet Key
(USNM 652409; A.M.); Mullet Key Bayway
(USNM 653110). BAHAMA ISLANDS:
GRAND BAHAMA ISLAND: South Hawksbill
Creek (ANSP 371809); ANDROS ISLAND:
South Mastic Point (А.М.). CUBA: Matanzas
(MCZ 131760). JAMAICA: Kingston (USNM
442584). MEXICO: N end of Ascension Bay,
Quintana Roo (USNM 736105); Isla Mujeres,
Quintana Roo (R.B.).

Genus Microtralia Dall, 1894

Microtralia Dall, 1894: 117. Type species by
monotypy: Auricula ? (Microtralia) mi-
nuscula (Dall, 1889) [= Leuconia occi-
dentalis Pfeiffer, 1854].

Rangitotoa Powell, 1933: 148. Type species
by monotypy: Rangitotoa insularis Pow-
ell, 1933.

Description: Shell to 3.8 mm long, subcylin-
dric, fragile, translucent white. Spire low to
moderately high, with as many as seven
weakly convex whorls. Body whorl 80% of
shell length. Aperture narrow, about 90% of
body whorl length; inner lip with small, ob-
lique columellar tooth; anterior parietal tooth
very near columellar tooth, strong; posterior
parietal tooth very small, about mid-length of
aperture; outer lip thin, sharp. Protoconch
smooth, globose; nuclear whorls deeply em-
bedded in first whorl of teleoconch.

Radula with 55 to 79 teeth in a row. Central
tooth at same level as lateral teeth; base
broad, triangular, anteriorly emarginate;
crown small, tricuspid. Base of lateral teeth
quadrangular, weakly bent medially; crown
less than half length of base, with large me-
socone, small ectocone. Transitional teeth
with one endocone. Marginal teeth wide,
pectinate, with as many as six ectocones.

Animal whitish to rusty brown, translucent.
Foot not divided transversely, posteriorly en-
tire, round. Eyes lacking. Tentacles short,
subcylindric. Hermaphroditic duct dilated an-
teriorly into a pouch-like seminal vesicle; an-

"90 75. 60 45 30

FIG. 173. Creedonia succinea, geographic distribu-
tion.

terior mucous gland covering posterior half of
spermoviduct; vas deferens free from penis.
Connectives of visceral ring very short.

Remarks: Since its introduction by Dall (1894)
the genus Microtralia has been considered to
belong to very different taxonomic groups. Its
uncertain taxonomic position is the result of
the different weights given by different au-
thors to the various taxonomic characters.
The etymology ofthe word implies similarity to
Tralia, a member of the Melampinae. Dall
(1894) tentatively placed Microtralia in the ge-
nus Auricula [= Ellobium], a member of the
Ellobiinae. Thiele (1931) considered Microtra-
lía a subgenus of Melampus. Powell (1933),
although recognizing the uniqueness of the
genus, followed Odhner's (1925) radula-
based classification and placed his Rangito-
toa, here considered а junior synonym of М/-
crotralia, in the Melampinae. Powell stressed
the radular affinities of his genus with the
Carychiinae. Morton (1955b), on the basis of
anatomy and habitat preferences, placed
Rangitotoa [= Microtralia] within the Pedipe-
dinae. Zilch (1959) treated Microtralia as a
subgenus of Melampus, and he considered
Rangitotoa as a separate genus of the
Melampinae. Abbott (1974) considered Mi-

232 MARTINS

crotralia a genus of the subfamily Cassiduli-
nae.

Although the shell is not typical of the
Pedipedinae, the dentition of the inner lip and
the protoconch of Microtralia are similar to
those of the more solid Pseudomelampus
and Sarnia (Figs. 180, 181). The central and
lateral teeth of the radula of this Eastern At-
lantic genus closely resemble those of Pe-
dipes, but the pectinate marginal teeth with
as many as six ectocones are very similar
to those of Pseudomelampus (Martins, per-
sonal observation). Analysis of the reproduc-
tive and nervous systems indicate the sys-
tematic position of Microtralia within the
Pedipedinae.

The Neozealandic Rangitotoa insularis
Powell, 1933, is quite similar to the West In-
dian Microtralia occidentalis (Pfeiffer, 1854),
especially in shell and radular characters,
and Climo (1982) considered them conspe-
cific (Fig. 179).

Habitat: These animals live near the high-
tide mark, under rocks partly buried in mud
(Powell, 1933). In West Indian mangroves Mi-
crotralia lives in the black sediment at the
high-tide mark, preferably under rotting, half-
buried branches (Martins, personal observa-
tion).

Range: Sporadic records from Easter Island
(Rehder, 1980), Hawaii (Pease, 1869), New
Zealand (Powell, 1933), Japan (Habe, 1961)
and South Africa (Turton, 1932) indicate an
Indo-Pacific distribution. In the West Indian
region the genus 1$ represented by Microtra-
lia occidentalis (Pfeiffer).

Microtralia occidentalis (Pfeiffer, 1854)
Figs. 174-178, 182-193

Leuconia occidentalis Pfeiffer, 1854b: 155
[Cárdenas, Cuba; location of type un-
known]; H. 8 A. Adams, 18556: 248;
Pfeiffer, 1856a: 157; Pfeiffer, 1876: 370;

Arango y Molina, 1880: 61; Crosse,
1890: 260.

Tralia (Alexia?) minuscula Dall т Simpson,
1889: 69 [Magill's Bay, Tampa, Florida,
and Exuma Island, Bahamas, herein re-
stricted to Magill’s Bay, Tampa, Florida;
lectotype herein selected USNM 61211
(Fig. 174); two paralectotypes USNM
859503].

Tralia minuscula Dall. Dall, 1889: 92.

Auricula ? (Microtralia) minuscula (Dall). Dall,
1894: 117, fig. 7 [Fig. 175].

Leucopepla occidentalis (Pfeiffer). Peile,
1926: 88.
Microtralia occidentalis (Pfeiffer). Pilsbry,

1927: 125; Morrison, 1951b: 10; Abbott,
1974: 334 [not figured; fig. 4105, errone-
ously referred to this species, represents
Myosotella myosotis]; Jensen & Clark,
1986: 456 ffig. on раде 456, wrongly
stated to represent this species, is of
Myosotella myosotis].

Auriculastrum (Microtralia) minusculum (Dall).
C.W. Johnson, 1934: 159.

Auriculastra nana Haas, 1950: 197, pl. 22,
figs. 1, 2 [Lover's Lake, St. George’s Is-
land, Bermuda; holotype FMNH 30169
(not seen); paratype ANSP 212177 (Fig.
176)].

Melampus (Microtralia) minusculus
Zilch, 1959: 65, fig. 208.

(Dall).

Description: Shell (Figs. 174-178, 182-184)
to 3.8 mm long, subcylindric, fragile, translu-
cent, white to yellowish. Spire low to moder-
ately high; whorls to five and three-fourths,
weakly convex, sculptured with very fine, un-
dulating spiral lines that extend over body
whorl. Body whorl about 80% of shell length,
crossed by faint, compact growth lines. Ap-
erture about 90% of body whorl length, nar-
row; inner lip with three teeth on anterior half;
columellar tooth small, oblique, twisted; an-
terior parietal tooth strong; posterior parietal
tooth very small, sometimes reduced to a

FIGS. 174-184. Microtralia, Rangitotoa, Pseudomelampus, Sarnia. (174) Tralia (Alexia?) minuscula Dall,
lectotype (USNM 61211), Magill's Bay, Tampa, Florida, sl 3.5 mm. (175) Auricula? (Microtralia) minuscula
Dall, Atkins Island, Bahamas (USNM 127487), si 2.3 mm; figured by Dall (1894, fig. 7). (176) Auriculastra
nana Haas, paratype (ANSP 212177), Lover's Lake, Bermuda, sl 3.2 mm. (177) M. occidentalis (Pfeiffer),
Hungry Bay, Bermuda, sl 3.5 mm. (178) М. occidentalis, Hungry Bay, Bermuda, $1 3.6 mm. (179) A. insularis
Powell, paratype (ANSP 242319), Rangitoto Island, Auckland, New Zealand, sl 3.2 mm. (180) P. exiguus
(Lowe), lectotype (BMNH 1875.12.31.109), Madeira, sl 5.8 mm. (181) S. frumentum (Petit), syntype? (BMNH
1843.11.24.58), Lima, Peru, sl 7.0 mm. (182) M. occidentalis, top view of spire and protoconch, Plantation
Key, Florida. (183) M. occidentalis, Hungry Bay, Bermuda, sl 3.1 mm. (184) M. occidentalis, lateral view of
spire and protoconch, Hungry Bay, Bermuda. Scale 1 mm.

233

WESTERN ATLANTIC ELLOBIIDAE

FIGS. 174-184.

234 MARTINS

FIGS. 185-188. Microtralia occidentalis, radular teeth. (185) Hungry Bay, Bermuda, sl 3.9 mm. (186) Grassy
Key, Florida. (187, 188) Hungry Bay, Bermuda, sl 3.9 mm. Scale 20 um.

barely visible callus at mid-length of aperture;
outer lip sharp, parallel to body whorl, sinu-
ous. Inner wall of whorls occupying less than
one-quarter of body whorl (Fig. 178). Proto-
conch globose; nuclear whorls deeply invo-
luted in first whorl of teleoconch; only small
portion of lip showing (Figs. 182, 184).

Animal whitish to rusty brown; tentacles
short, subcylindric, with tip weakly pointed or
somewhat flat and expanded. Eyes lacking.
Mantle skirt whitish with brownish tinge along
border. Pallial cavity somewhat elongate;
kidney broadly triangular, anteriorly rounded,
covering most of pallial cavity; pneumo-
stomal and anal openings prolonged by a
tube-like flap of mantle skirt; anal gill well de-
veloped.

Radula (Figs. 185-189) having formula [15
+ (3 + 16) + 1 + (16 + 3) = 15] x 95. Crown of
central tooth small, as wide as posterior end
of base, tricuspid; mesocone small, blunt to
weakly pointed; ectocones very small but
well defined. Lateral teeth 12 to 18; crown

ST

FIG. 189. Microtralia occidentalis, radula, Grassy
Key, Florida. Scale 10 um.

wider than base, bicuspid; mesocone
broadly rounded anteriorly, becoming more
pointed and longer toward marginal teeth.
Transitional teeth two to three, with small en-
docone, thinner and longer ectocone. Mar-

WESTERN ATLANTIC ELLOBIIDAE 235

FIG. 190. Microtralia occidentalis, stomach, Ber-
muda. Scale 1 mm.

ginal teeth 13 to 19; base short and wide,
with lateral flare, on which endocone of next
tooth articulates; Crown gradually widening
and mesocone gradually becoming shorter
and thinner toward margin; first marginal
tooth with two ectocones; additional ecto-
cones appearing on fourth, eighth and twelfth
marginal teeth; sometimes a sixth ectocone
appears on twelfth marginal tooth in some
rows.

Digestive system (Fig. 190) having diges-
tive gland with two subequal lobes. Posterior
crop dilated, receiving anterior diverticulum
just before joining stomach. Anterior portion
of stomach thin, with inner thickening be-
tween entrance of esophagus and exit of
intestine; mid-stomach gizzard-like, thickly
muscular; gastric caecum thin, dilated, re-
ceiving posterior diverticulum at junction with
gizzard.

Reproductive system (Fig. 191) with
ovotestis acinose, trilobed, conical, at poste-
rior tip of visceral mass, covering stomach;
hermaphroditic duct straight, with anterior,
pouch-like seminal vesicle connecting with
convoluted fertilization chamber; albumen
gland and posterior mucous gland large; an-
terior mucous gland and prostate gland cov-
ering posterior half of spermoviduct; bursa
duct as long as spermoviduct, thick, empty-

FIG. 191. Microtralia occidentalis, reproductive
system, Hungry Bay, Bermuda. A, B, transverse
sections and their locations. Scale 1 mm.

ing near opening of vagina; bursa oval-elon-
gate; vas deferens separates from oviduct
near opening of vagina. Penis short, thick;
associated vas deferens free, somewhat
longer than penis; penial retractor about as
long as penis, inserting on penis subapically,
attaching to anterior portion of floor of pallial
cavity.

Nervous system (Fig. 192) with cerebral
commissure somewhat shorter than width of
cerebral ganglion; left cerebropleural and
cerebropedal connectives longer than right
ones; connectives of visceral ring very short,
causing agglomeration of ganglia; pedal
commissure short but conspicuous. Cerebral
ganglia largest; pleural ganglia well devel-
oped; left parietal ganglion very small; right
parietal ganglion and visceral ganglion about
same size. Penial nerve branching from me-
dial lip nerve.

Remarks: Originally Pfeiffer (1854b) assigned
Microtralia occidentalis to the genus Leuco-
nia Gray, 1840, which, because it was preoc-
cupied, was renamed Leucopepla by Peile
(1926). Pilsbry (1927) showed that on the ba-
sis of shell characters Microtralia occidentalis
could not be placed in Leucopepla [= Auricu-
linella]. The latter genus belongs in the Ello-
biinae on the basis of its nervous and repro-

236 MARTINS

FIG. 192. Microtralia occidentalis, central nervous
system, Hungry Bay, Bermuda. Scale 1 mm.

ductive systems, and Microtralia (see re-
marks under the genus) rightly belongs in the
Pedipedinae.

Dall (1889), apparently unaware of Pfeif-
fer's name, described Tralia (Alexia?) minus-
cula (Fig. 174) for which he created, in 1894,
the subgenus Microtralia, tentatively remov-
ing it to the genus Auricula. The odd combi-
nations of names representing such different
groups indicate the extent to which Dall was
confused about the relationships of this small
species.

Microtralia occidentalis shows some mor-
phological variation within populations and
across its geographical range. Bermudian
specimens are brownish and have the tips of
the tentacles somewhat flat and broad. Flo-
ridian specimens are usually whitish, some-
times yellowish brown, and the tentacles are
subcylindrical with blunt or weakly pointed
tips. Bahamian examples are rusty brown and
the tips of the tentacles are intermediate in
shape between Bermudian and Floridian
specimens. The radulae of specimens from
Bermuda have rounder and somewhat shorter
cusps than do those from Florida, but other-
wise show no other morphological differ-
ences. Intrapopulational variations in shell
morphology, especially the height of the spire
and the strength of the apertural teeth, occur
throughout the range of the species.

Haas (1950) apparently was unaware of
Pfeiffer’s or Dall’s names when he introduced
Auriculastra nana from Bermuda, for he
did not refer to either author in the original
description. Haas’ species (Fig. 176) does
not differ from Microtralia occidentalis and it

must therefore be considered a junior syn-
onym of the latter.

Climo (1982), as noted under the remarks
for the genus, synonymized Rangitotoa insu-
laris Powell with Microtralia occidentalis, con-
sidering the former to have been introduced
in New Zealand and possibly also in Rapa Iti
Island and Easter Island. | concur with Cli-
mo’s taxonomic decision about the genera,
on the basis of the conchological and radular
similarities; however, the widespread distri-
bution of the genus (See the remarks for the
genus) and the anatomical differences ob-
served in Microtralia alba (Gassies, 1865)
from Hong Kong (Martins, 1992), preclude an
immediate synonymization of both species.

An anatomical peculiarity of Microtralia oc-
cidentalis is the absence of eyes, confirmed
by histological examination. Concealment of
the eyes under the skin has been reported for
several species of the genus Ellobium (Pelse-
neer, 1894a: 75, note 1). In the West Indian
species Ellobium (A.) dominicense the eyes,
although covered by thick skin, are readily
visible.

The shell of Microtralia occidentalis is not
confused easily with that of any other West
Indian ellobiid. It can resemble the very thin-
shelled juveniles of some populations of Tra-
lia ovula, however. Microtralia has a large,
rounded protoconch, faintly incised, undulat-
ing lines on the spire and a posterior parietal
tooth that is anterior to the mid-length of the
aperture. In Tralia ovula the apex is mu-
cronate, the spire has marked, pitted lines
and the posterior parietal tooth 15 in the pos-
terior half of the aperture.

Microtralia commonly occurs with Pedipes
ovalis, Laemodonta cubensis, Blauneria het-
eroclita and Creedonia succinea and 1$
readily distinguished from them. Pedipes is
globose and has a rounded, strongly dentate
aperture, Laemodonta is oval-elongate and
hirsute and has a heavily dentate aperture,
Blauneria has a sinistral shell and Creedonia
has a truncated spire and very different inner
lip dentition.

Habitat: Microtralia occidentalis lives at or
above the high-tide mark, buried in the
sediment sometimes 10 to 15 cm deep, in
the company of В/аипепа heteroclita and
Creedonia succinea. The animals are quite
common under partly buried, rotting wood or
porous rocks, and on the roots of mangrove
propagules, on which Laemodonta cubensis
and Pedipes ovalis also abound.

WESTERN ATLANTIC ELLOBIIDAE 237

90 75. 60 45 30

FIG. 193. Microtralia occidentalis, geographic dis-
tribution.

Range: Bermuda; Clearwater, Florida, south
to the Florida Keys and the Greater Antilles
(Fig. 193).

Specimens Examined: BERMUDA (USNM
250297); Fairyland (ANSP 99075; USNM
208069); Old Road, Shelly Bay (A.M.); Lover's
Lake, St. George's (ANSP 212172); near St.
George's (ANSP 1008220); Castle Harbour,
near Harrington House (ANSP 143322); Coo-
pers Island (ANSP 131645); Hungry Вау
(A.M.); S End of Ely's Harbour (A.M.); Man-
grove Bay (A.M.). FLORIDA: N of Tavernier
Creek, Key Largo (A.M.); S of Ocean Drive,
Plantation Key (A.M.); Lignumvitae Key
(ANSP 156682); Long Key (A.M.); Grassy
Key (A.M.); Crawl Key (A.M.); Bahia Honda
Key (ANSP 104108); Big Pine Key (ANSP
104102); W of Kohen Avenue, Big Pine Key
(A.M.); Big Torch Key (ANSP 104001; A.M.);
Sugarloaf Key (ANSP 89558); Boca Chica
Key (ANSP 152503; USNM 270350); Semi-
nole Point (ANSP 105409); Blue Hill, Hors Is-
land (ANSP 99199); Captiva Island (ANSP
131836); McGill's Bay, near Tampa (USNM
61211, 859503); Boca Ciega Bay (ANSP
9571); Pinellas Point (USNM 83255); Clear-
water Island (ANSP 9351). ВАНАМА IS-
LANDS: GREAT ABACO ISLAND: Mores Is-

land (MCZ 294207); ANDROS ISLAND: South
Mastic Point (A.M.); Stafford Lake (ANSP
294338); Mangrove Cay (USNM 270214b);
NEW PROVIDENCE ISLAND: W of Clifton
Point (A.M.); E of Clifton Pier (A.M.); shore
of Millars Road (A.M.); Bonefish Pond
(A.M.); AKLINS ISLAND: Pinnacle Point
(USNM 390857a); SAN SALVADOR (USNM
127487): Bob's Key, S. Ferdinand (USNM
360499). CUBA (ANSP 22482): near Habana
(ANSP 130794). JAMAICA: Falmouth (ANSP
397269); Robin's Bay (USNM 441980,
442113); Kingston (USNM 395452b); Rio Co-
bre, Port Royal (USNM 426889); Hunt's Bay
(USNM 441642); Rock Fort (USNM 467164).
HAITI: Gonave Island (USNM 380184). DO-
MINICAN REPUBLIC: Rio Guayabin by Sa-
baneta Road (ANSP 160398). PUERTO
RICO: Puerta de Tierra, San Juan (A.M.); Pu-
erto Real (A.M.).

Genus Leuconopsis Hutton, 1884

Leuconopsis Hutton, 1884: 213. Type spe-
cies by monotypy: Leuconopsis obsoleta
(Hutton, 1878).

Apodosis Pilsbry & McGinty, 1949: 9. Type
species by monotypy: Apodosis novi-
типа! Pilsbry & McGinty, 1949.

Description: Shell to 4 mm long, oval-conic to
oblong-conic, somewhat thin to solid. Spire
low to moderately high, with as many as six
and one-half flat, striated whorls. Body whorl
about 80% of shell length. Aperture about
75% length of body whorl, approximately
oval, posteriorly angulate; inner lip with strong
submedian columellar tooth, usually with
weak anterior secondary tooth; outer lip
sharp. Protoconch smooth; nuclear whorls
embedded in first whorl of teleoconch.
Radula with 87 to 111 teeth in a row. Base
of central tooth widened anteriorly, sharply
constricted posteriorly; crown thin, falciform;
mesocone long, sharp. Base of lateral teeth
abruptly bent medially at half length; crown as
wide as base, with strong endocone. Transi-
tional teeth lacking. Marginal teeth with strong
mesocone, weaker endocone and ectocone.

Remarks: The genus Apodosis was created
by Pilsbry & McGinty (1949) for the smallest
and rarest West Indian ellobiid. In the original
description the authors stated (p. 10) that
they were “strongly inclined to treat Apo-
dosis as a subgenus of the antipodal genus
Leuconopsis Hutton.” The shape of the shell,
oblong-conic in the Atlantic species vs oval-

238 MARTINS

conic in the type species (Fig. 217) the unim-
pressed suture and the inner thickening of
the outer lip led them to establish a new ge-
nus. Examination of additional specimens of
Leuconopsis novimundi revealed that the
thickness of the outer lip varies with the
thickness of the shell, and that the distinct
thickened outer lip actually does not appear
in some thin-shelled specimens. This varia-
tion was observed in Leuconopsis manningi
n. sp. from Ascension Island and in Leu-
conopsis rapanuiensis Rehder, 1980, from
Easter Island. The other characters men-
tioned by Pilsbry 4 McGinty are significant
only at the specific level. The most obvious
generic shell characters are the absence of a
parietal tooth and the presence of a weak
secondary columellar tooth just anterior to
the primary submedian columellar tooth. This
columellar structure is reminiscent of Pe-
dipes, Marinula and Сгеедота.

Powell (1933) illustrated the radula of Leu-
conopsis obsoleta (Hutton, 1878). The mor-
phology of the teeth is very similar to that of
the West Indian Apodosis novimundi. On the
basis of shell and radular characters, Apo-
dosis Pilsbry 8 McGinty must be considered
a junior synonym of Leuconopsis Hutton.

Habitat: The genus lives intertidally under
rocks (Hutton, 1884). Powell (1933: 150)
found Leuconopsis obsoleta “in sheltered
harbour bays towards high-tide, .. . and on
cliffs, just above high-tide mark, in situations
where fresh-water seepage occurs.” | col-
lected Leuconopsis novimundi on New Prov-
idence Island, Bahama Islands, in crevices of
cliffs, just above high tide, but | did not see
any indication of freshwater seepage.

Range: The genus Leuconopsis occurs in
the Pacific in Australia, New Zealand and
Easter Island. In the Atlantic it is represented
by Leuconopsis novimundi (Pilsbry & Мс-
Ginty) from the Florida Keys, Bahamas and
Jamaica, and by Leuconopsis manningi,
herein described, known only from Ascen-
sion Island. A possible third species from St.
Thomas is left unnamed owing to lack of suit-
able material.

Leuconopsis novimundi
(Pilsbry & McGinty, 1949)
Figs. 194-204

Apodosis novimundi Pilsbry & McGinty,
1949: 10, pl. 1, fig. 1 [Clifton Bluff, New
Providence, Bahamas; holotype ANSP

185474a (Fig. 194)]; Morrison, 1951b: 9;
Zilch, 1959: 70, fig. 227; Franc, 1968:
525; Abbott, 1974: 334, fig. 4102. Non
“Pilsbry & McGinty” Rosewater, 1975
[misindentification of Leuconopsis man-
ningi Martins n. sp.].

Description: Shell (Figs. 194-197) to 3.4 mm
long, oblong-conic, solid, uniformly light
yellow to pale brown. Umbilical area marked
by shallow excavation. Spire moderately
high, whorls аз many as six and one-half,
flat and heavily sculptured with numerous
spiral cords, intersected by compact axial,
somewhat regularly spaced, fine growth
lines, giving shell a matte appearance. Body
whorl convex, about 75% of shell length and
with same sculpture as spire. Aperture about
70% body whorl length, subaxial, narrowly
ovate; inner lip with partly hidden tooth at
point of juncture of columella and parietal
wall; occasionally secondary columellar
tooth present as faint callosity just anterior to
columellar tooth; outer lip sharp. Protoconch
oblong, smooth, transparent, with sinuous lip
(Fig. 197).

Animal whitish gray; foot dirty white, trans-
versely divided; tentacles short, transparent,
subcylindric, with rounded tip; mantle skirt
slightly lighter than rest of animal.

Radula (Figs. 198-201) having formula
(33 + 10 + 1 + 10 + 33) x 75+. Base of central
tooth with quadrangular anterior half, round
at tip; width of posterior half abruptly re-
duced to half; crown as wide as posterior half
of base, falciform; mesocone just over half
length of base, pointed. Lateral teeth bicus-
pid; mesocone sharp, as long as, but stron-
ger than, that of central tooth; endocone
sharp, almost as long and strong as meso-
cone. Marginal teeth tricuspid; ectocone be-
coming as long as endocone; mesocone be-
coming smaller, but remaining the strongest
cusp.

Digestive system with posterior crop wide,
with strong internal folds; stomach very mus-
cular, with gastric caecum where posterior
diverticulum empties.

Reproductive system (Fig. 202) semidiau-
lic, with vas deferens separating from oviduct
almost at half-length of pallial gonoduct; an-
terior mucous gland covers spermoviduct
nearly to separation of vas deferens; bursa
duct empties near female opening. Penial
complex dilated, pouch-like in mid-section;
posterior section thinner, coming out of
pouch as blunt diverticulum; penial retractor

ee

WESTERN ATLANTIC ELLOBIIDAE 239

FIGS. 194-200. Leuconopsis novimundi (Pilsbry & McGinty), Clifton Bluff, New Providence, Bahamas. (194)
Holotype (ANSP 185474a), sl 3.36 mm. (195) Shell length 2.73 mm. (196) Lateral view of spire and proto-
conch. (197) Top view of spire and protoconch. (198) Central, lateral and marginal teeth of radula. (199)
Marginal teeth of radula. (200) Marginal teeth of radula. Scale, Figs. 196, 197, 1 mm; Figs. 198-200, 20 um.

multifid, short, attaching to end of posterior
section of penis subapically; vas deferens
free, entering penis at base of thinner poste-
rior section.

Nervous system (Fig. 203) with cerebral
ganglia largest; left pleural ganglion larger
than right one; left parietal ganglion much FIG. 201. Leuconopsis novimundi, radula, Clifton
smaller than right one; visceral ganglion Bluff, New Providence, Bahamas. Scale 10 um.

240 MARTINS

FIG. 202. Leuconopsis novimundi, reproductive system, Clifton Bluff, New Providence, Bahamas. Penis
drawn under camera lucida, pallial gonoducts reconstructed from histological sections. A-L, transverse

sections and their locations. Scale 1 mm.

about as large as right parietal ganglion. Ce-
rebral commissure somewhat longer than
width of cerebral ganglion; right and left cere-
bropedal and cerebropleural connectives
roughly equal; left pleuroparietal connective
very small; all other visceral ring connectives
equal, about half length of cerebropleural
connectives.

Remarks: Leuconopsis novimundi shows
some anatomical deviations from typical
members of the Pedipedinae such as Pe-
dipes and Creedonia (see the remarks under
the subfamily). The origin of the vas deferens
from the mid-section of the spermoviduct-
vaginal tract was unexpected in this species
because Morton (1955b) stated that the sep-
aration of the vas deferens from the sper-
moviduct in Leuconopsis obsoleta occurs
at the vaginal opening, as in Ovatella [sensu
Myosotella]. Nevertheless, Leuconopsis novi-
mundi is placed in the Pedipedinae on the
basis of the presence of the double columel-
lar tooth in the aperture, the shape of the
crown of the central and lateral teeth, the
prostate and anterior mucous gland not

FIG. 203. Leuconopsis novimundi, central nervous
system, Clifton Bluff, New Providence, Bahamas.
Scale 1 mm.

reaching the female opening, and the rela-
tively short visceral nerve ring, as compared
to that of the Pythiinae or Ellobiinae.

WESTERN ATLANTIC ELLOBIIDAE 241

Rosewater (1975) erroneously identified a
small Leuconopsis from Ascension Island
as Apodosis novimundi. These specimens
(USNM 735149, 859015) differ from Leu-
conopsis novimundi in the differently sculp-
tured shell that lacks the matte appearance
and in the readily visible secondary columel-
lar tooth. They represent a previously unde-
scribed species that is introduced as Leu-
conopsis manningi п. sp. in this report (Figs.
205, 206). The protoconch of Leuconopsis
novimundi is very similar to that of Leuconop-
sis manningi in having a sinuous lip (Figs.
207, 208), a characteristic lacking in Leu-
conopsis rapanuiensis (Fig. 216).

Leuconopsis novimundi is readily distin-
guished from all other Western Atlantic ello-
biids by its oval-elongate shell that has a
matte appearance and by its lack of parietal
teeth in the aperture.

Habitat: The two specimens collected by R.
Robertson in 1964 on Pigeon Cay, Bimini, in
algae on mangrove roots were probably the
first ones to be collected alive. All other spec-
imens in museum collections, including those
of Pilsbry and McGinty, seem to have been
obtained from beach drift. Despite thorough
field work and patient rock-combing, all but
one of the specimens | found alive came from
a cave at the western tip of Clifton Bluff (Clif-
ton Pt.), New Providence Island, Bahamas,
kindly indicated to me by T. L. McGinty. The
cave formed from a double crack in the coral
bed, running from sea to shore. lt is open
above, so direct sunlight illuminates it a few
hours a day. At high tide water enters the
main opening from the ocean, as well as the
bottom of the double crack. One specimen
was found among stones that had collected
in one such crack, just above high-tide mark.
Eight others were found in the crevices as
deep as 20 cm in the wall and were obtained
by chipping away the wet layers of coral in a
band about 15 cm wide just above the black
zone. The animals apparently feed on the de-
tritus that collects in these crevices. The
eight specimens were found within a radius
of about 20 cm, together with Pedipes ovalis,
Laemodonta cubensis and young Melampus
(D.) monile. The other live specimen was
found at Morgan's Bluff, Andros Island, Ba-
hamas, under stones at the high-tide mark of
a tidal pool.

Jamaica

Range: Florida Keys, Bahamas,

(Fig. 204).

60 “45 30

FIG. 204. Leuconopsis novimundi, geographic dis-
tribution.

Specimens Examined: FLORIDA: Indian
Key (USNM 492557a). BAHAMA ISLANDS:
GRAND BAHAMA ISLAND: Gold Creek
(ANSP 369338); Hepburn Town, Eight Mile
Rock (ANSP 370409); Caravel Beach [John
Jack Point], Freeport (ANSP 370225); BIMINI
ISLANDS: N end of Pigeon Cay (ANSP
329623); ANDROS ISLAND: Morgan’s Bluff
(A.M.); Mangrove Cay (USNM 1804625); First
island off Mintie Bar, SE of South Bight
(USNM 271888); NEW PROVIDENCE IS-
LAND: Clifton Bluff (ANSP 185474; А.М.). JA-
MAICA: Jack’s Bay (USNM 441915).

Leuconopsis manningi n. sp.
Figs. 120, 205-212

Apodosis novimundi Pilsbry & McGinty.
[Type locality: English Bay, Ascension Is-
land; holotype USNM 859015 (Fig. 205);
11 paratypes USNM 859015 (Fig.
206)].Rosewater, 1975: 23. Non Pilsbry
& McGinty, 1949.

Description: Shell (Figs. 205-208) to 2.1 mm
long, ovate, solid, uniformly pale to dark
brown, smooth, shiny. Umbilical excavation
very weak. Spire short, whorls as many as
five and one-fourth, flat, sculptured with two

242 MARTINS

Mi?

pd à |

FIGS. 205-211. Leuconopsis manningi n. sp., English Bay, Ascension Island. (205) Holotype (USNM
859015), si 2.02 mm. (206) Paratype (USNM 859015), sl 1.58 mm. (207) Lateral view of spire and proto-
conch. (208) Top view of spire and protoconch. (209) Central and lateral teeth of radula. (210) Marginal teeth
of radula. (211) Marginal teeth of radula. Scale, Figs. 207, 208, 1 тт; Figs. 209-211, 20 um.

to seven incised spiral lines on shoulder in
adults, juveniles often spirally striated on en-
tire length; growth lines very faint. Aperture
75% of body whorl length, ovate; inner lip
with strong posterior columellar tooth about
mid-length of aperture, with conspicuous but
much smaller anterior secondary tooth; outer
lip sharp. Protoconch oblong, smooth, trans-
parent, dark brown, with sinuous lip.

Radula (Figs. 209-212) having formula
(32+11+1+11+32)x70+. Radular morphol-
ogy as in Leuconopsis novimundi.

Animal unknown.

Remarks: Leuconopsis manningi was first
mentioned in the literature by Rosewater
(1975), who misidentified it as Apodosis
novimundi. This new species differs from

WESTERN ATLANTIC ELLOBIIDAE 243

C iL 2L 11L 1M 2M 22M 23M 31M 32M

| | IM LA a Аа» y a à en
LU ALL MA MA à N y ) 7
\ ] PRA L a | у L
<a ES

FIG. 212. Leuconopsis manningi, radula, Ascen-
sion Island. Scale 10 um.

Leuconopsis novimundi in the ovate shape
of the shell, the proportionately longer,
nonmatte body whorl that usually has two
incised lines on the shoulder, the larger ap-
erture and the conspicuous secondary col-
umellar tooth. Leuconopsis manningi is
closely related to Leuconopsis rapanuiensis
Rehder, 1980, from Easter Island (Figs. 214-
216). The secondary columellar fold of the
latter is farther foreward than in Leuconopsis
manningi, the protoconch is stouter and has
a rounded instead of sinuous lip.

The single radula studied was obtained by
breaking the shell of a dried animal; the apex
of the shell was used for the SEM study of the
protoconch.

Following the suggestion of the late J.
Rosewater | name this species for R. B. Man-
ning, who collected the specimens in 1971.

Habitat: Intertidal pools, subtidal rocky
shore, with some coarse sand (from USNM
label).

Range: English Bay, Ascension Island (Fig.
120).

Leuconopsis sp.
Rig: 213

Description: Shell (Fig. 213) 4 mm long, oval-
conic, solid, white. Spire moderately high,
with about six weakly convex whorls. Body
whorl 77% of shell length. Aperture semilu-
nate, 67% of body whorl; inner lip with mod-
erately strong, horizontal columellar tooth
just anterior to mid-length of aperture; obvi-
ous secondary tooth somewhat anterior to
previous one; outer lip sharp.

Remarks: A single specimen from St. Thom-
as, Virgin Islands, originally from the Swift
collection (now ANSP 22599), differs from the
other Atlantic species of Leuconopsis in its
wider body whorl, in its anteriorly less ex-
panded outer Пр, which gives the арейиге a
semilunate aspect, and in its conspicuous
secondary columellar tooth at some distance
anterior to the primary tooth. The example is

undoubtedly a beach specimen, a fact that
might account for the absence of any visible
sculpture.

| am reluctant to erect a new species upon
a single worn specimen; the naming of this
probably new species must await collection
of more material.

Subfamily Melampinae Pfeiffer, 1853

Melampinae Pfeiffer, 1853a: 8 [corrected from
Melampea by H. & A. Adams, 1855b)].

Melampodinae Fischer & Crosse, 1880: 5
[unjustified emendation].

Description: Shell to 23 mm long. Spire low
to high. Body whorl usually more than 75% of
shell length. Aperture elongate, narrow; one
columellar tooth; one to five parietal teeth;
outer lip internally smooth or with as many as
18 riblets. Protoconch nipple-like, smooth,
with spiral axis perpendicular to columellar
axis of teleoconch; nuclear whorls only partly
covered by first whorl of teleoconch.

Animal white to black, generally brown;
foot transversely divided, bifid posteriorly.

Radula with central tooth posterior to lat-
eral teeth, triangular, with base deeply in-
dented; crown with narrow, sharp mesocone,
ectocones very small or absent. Base of lat-
eral teeth rectangular, medially bent in shape
of boomerang, with medial node on inner sur-
face; crown with strong mesocone pointing
laterally, sometimes with endocone or ecto-
cone. Transitional teeth with shorter base,
crown more elongate posteriorly and extra
denticle on ectocone. Marginal teeth with
very strong mesocone, one or two denticles
оп endocone and one to eight denticles on
ectocone; base and crown gradually become
shorter from innermost to outermost teeth
and denticles of ectocone gradually advance
to same level, becoming subequal in size.

Digestive system having mandible tripar-
tite, of numerous longitudinal fibers; stronger
central portion lining upper lip and folded, ta-
pering extremities lining lateral lips. Salivary
glands yellowish to white, long, fusiform.
Esophagus with internal, longitudinal folds,
ending posteriorly in a wide crop that receives
anterior diverticulum of digestive gland; a thin
muscular strand secures posterior crop to
posterior portion of muscular band of stom-
ach. Digestive gland brown to dark orange,
bilobed; anterior lobe very large, composed of
several lobules that empty into wide anterior
diverticulum; posterior lobe small, located

244 MARTINS

FIGS. 213-217. Leuconopsis. (213) Leuconopsis sp., St. Thomas, Virgin Islands (ANSP 22599), formerly in
Swift collection, sl 4.04 mm. (214) L. rapanuiensis (Rehder), holotype (USNM 756790), Easter Island, sl 2.87
mm. (215) L. rapanuiensis, paratype (USNM 756238) Easter Island, sl 2.34 mm. (216) L. rapanuiensis, top
view of spire and protoconch. (217) L. obsoleta (Hutton), Takapuna, Auckland, New Zealand (USNM
681309), sl 2.50 mm. Scale 1 mm.

partly beneath ovotestis, covering posterior
left portion of stomach and emptying into
small posterior diverticulum. Stomach glob-
ular, tripartite; anterior portion thin, weakly
muscular at cardiac aperture; middle portion
surrounded by thick band of muscle around
pyloric region; posterior portion of caecum
thin, receiving posterior diverticulum at ante-
rior border, just posterior to muscular band
and near attachment of muscular strand;

stomach attached to mantle by muscular fi-
bers extending from region opposite opening
of posterior diverticulum. Intestine dilates as
it leaves the stomach anteriorly just right of
esophagus, and has several convolutions
in midst of digestive gland; rectum parallels
right edge of pallial cavity; anal opening lateral
to pneumostome, on mantle lappet. Anal gill
bilobed, flanking rectum just posterior to
anus.

WESTERN ATLANTIC ELLOBIIDAE 245

FIGS. 218-229. Melampus (M.) coffeus (Linnaeus). (218) Lectotype (LSL), si 18.8 mm. (219) Paralectotype
(LSL), sl 11.0 mm. (220) Auricula biplicata Deshayes, holotype (ММНМР), $1 20.0 mm. (221) М. coffea var.
microspira Pilsbry, holotype (ANSP 61471), Progreso, Yucatán, Mexico, sl 12.8 mm. (222) Bermuda (USNM
11421), sl 15.2 mm. (223) Grand Bahama Island, Bahamas (MCZ 116679), sl 17.3 mm. (224) Anegada, Virgin
Islands (MCZ 229004), si 18.3 mm. (225) Isla Matica, Dominican Republic (R.B.), $1 10.4 mm. (226) Juvenile,
Puerto Real, Puerto Rico, sl 3.6 mm. (227) Tucacas, Venezuela, sl 10.5 mm. (228) Bahía [San Salvador],
Brazil (AMNH 22434), sl 12.0 mm. (229) Boa Viagem, Brazil (MCZ 219130), sl 19.5 mm.

246 MARTINS

Reproductive system of advanced semi-
diaulic type, the short spermoviduct separat-
ing into long, thin vagina and vas deferens
shortly after passing posterior mucous gland;
posterior mucous gland large, spiral, cover-
ing only small portion of spermoviduct; pros-
tate gland beneath posterior mucous gland,
not discrete; anterior mucous gland absent;
vagina running posteriorly, turning abruptly
and following columellar muscle anteriorly;
elbow of vagina attached by muscle fibers to
left corner of insertion of columellar muscle.
Bursa with short peduncle, at or very near
proximal end of vagina. Penis long, usually
simple; anterior vas deferens long, thin, en-
tering penis apically; penis and posterior vas
deferens run beneath upper tentacle retrac-
tors, over cerebral commissure and under
right tentacular nerve.

Nervous system having cerebral commis-
sure short; cerebropleural connectives about
same length as cerebropedal connectives;
pleuroparietal and parietovisceral connec-
tives very short.

Remarks: Baker (1963) showed that, al-
though the word Melampus has its origin in
the Greek, meaning “black foot,” it was La-
tinized and used by the Romans in the gen-
itive case, “melampi.” Hence Melampinae
must be used instead of Melampodinae.

During growth the radula changes consid-
erably in shape and number of teeth (Table 4,
Appendix). In very young animals the inner
edge of the arms of the base of the tricuspid
central tooth bears conspicuous promi-
nences. The first lateral tooth can be either
tricuspid or bicuspid. The marginal teeth
have an ectocone that becomes serrate, car-
rying as many as eight subequal cusps. As
the animal grows, certain characteristics are
retained, such as the splitting of the en-
docone and ectocone. Others are enhanced,
such as the serrations on the lateral edge of
the crown of the marginal teeth of young
Melampus s. s. before the ectocone be-
comes a distinct cusp. Some characteristics,
such as the ectocone and endocone of the
lateral teeth, become lost in most species.
Even in adults the same tooth in adjacent
rows might be inconsistent. The characteris-
tics given in the description must be consid-
ered as the general pattern in adult individu-
als.

In all animals examined the stomach
agrees with Koslowsky's (1933) and Marcus
8 Marcus’ (1965a) descriptions, rather than

with Morton 's (1955c) account, which did not
record the presence of the caecum and pos-
terior diverticulum. No evidence was found to
support Marcus 8 Marcus’ statement that the
stomach of Melampus gundlachi Pfeiffer,
1853 [= Melampus (M.) bidentatus Say, 1822]
is radically different from that of other West
Indian Melampinae.

Only two genera compose the subfamily
Melampinae: Melampus Montfort and Tralia
Gray. Melampus 15 easily distinguished from
Tralia on the basis of its shell. Melampus has
a much narrower aperture than does Tralia,
the dentition of the inner Пр 1$ restricted to
the anterior half, the anterior parietal tooth,
when present, is very small and the outer lip
is usually interiorly ribbed. Tralia has a strong
anterior parietal tooth and always has an-
other conspicuous parietal tooth on the pos-
terior half of the aperture. The ощег Пр 1$ sin-
uous and has only one ridge-like riblet
opposite the posterior parietal tooth. Zilch
(1959) listed Rangitotoa Powell in this sub-
family, but results of this study led me to con-
clude with Climo (1982) that Rangitotoa is a
junior synonym of Microtralia Dall, which
Zilch had listed as a subgenus of Melampus.
The genus Microtralia belongs to the Pedipe-
dinae by reason of the morphology of the re-
productive and nervous systems.

The Melampinae are separated from the
other ellobiid subfamilies on the basis of rad-
ula, reproductive system and nervous sys-
tem. The serration of the ectocones of the
marginal teeth of the radula has some parallel
only in Microtralia. The nonglandular pallial
gonoducts, the proximal position of the bursa
duct and the long, generally thin penis are
unique. The concentration of ganglia has
some parallel in the Pedipedinae, but in the
Melampinae the cerebral commissure 1$ rel-
atively shorter and the cerebropleural and the
cerebropedal connectives are much longer.

A planktonic veliger has been reported for
several Melampinae (Morrison, 1959; Rus-
sell-Hunter et al., 1972; Berry, 1977). This
condition is considered primitive, retained
from the estuarine habit of the ancestors of
the ellobiids.

Habitat: The Melampinae are the most com-
mon ellobiids in the Western Atlantic, living
mostly in salt marshes and in mangroves.
They can occur in zones of very low salinity
such as along the banks of rivers some miles
inland [Melampus (M.) bidentatus, Melampus
(D.) floridanus], or under rocks exposed to

WESTERN ATLANTIC ELLOBIIDAE 247

high-tide surf [Melampus (D.) monile, Tralia
(T.) ovula]. They are among the common gas-
tropods found in Stephenson 4 Stephen-
son’s (1950) upper intertidal gray zone.

Range: Worldwide distribution, except in the
Mediterranean region. п the Western Atlantic
the Melampinae extend from Newfoundland
to southern Brazil. Species are especially nu-
merous in the West Indian region.

Genus Melampus Montfort, 1810

Melampus Montfort, 1810: 319. Type species
by monotypy: Melampus coniformis
(Bruguière, 1789) [= Melampus coffeus
(Linnaeus, 1758)]. Non Gray, 1865
[Mammalia].

Conovulus Lamarck, 1816, pl. 459, fig. 2 a.
b., Liste, p. 12. Type species herein des-
ignated, Conovulus coniformis (Bru-
guière, 1789) [= Melampus coffeus (Lin-
naeus, 1758)].

Melampa “Draparnaud” Montfort. Schweig-
ger, 1820: 739 [unjustified emendation of
Melampus].

Conovula Lamarck. Schweigger, 1820: 739
[in synonymy; unjustified emendation of
Conovulus].

Conovulae Lamarck. Férussac, 1821: 104
[unjustified emendation of Conovulus].

Conovolus Lamarck. Sowerby, 1839b: 10 [er-
ror for Conovulus].

Conovulum Lamarck. Sowerby, 1842: 119
[unjustified emendation of Conovulus].

Maelampus Montfort. Reeve, 1877, pl. 1 [in
synonymy; error for Melampus)].

Description: Shell ovoid, white to dark
brown, uniform or with light spiral bands or
axial stripes. Aperture high, narrow, with wid-
est point above columellar tooth; first parietal
tooth reduced or absent; outer lip sharp, with
lirae.

Animal grayish blue to black; tentacles
subcylindric, pointed.

First lateral tooth of radula with or without
ectocone, always lacking endocone.

Salivary glands attaching to ventral portion
of esophagus, right one in front of left one.
Esophagus heavily pigmented.

Reproductive system having ovotestis of
dark yellow radiating tubules with or without
dark brown spots, shallow-conic to leaf-like,
rounded or lobed at base, deeply split on
right side; gonadial artery entering ovotestis
from left, bifurcates and radiates, covering
lower surface of gland. Hermaphroditic duct

brown, moderately long, forming very convo-
luted seminal vesicle, passing beneath pos-
terior diverticulum. Albumen gland spiral.
Posterior end of short, nonglandular sper-
moviduct with pouch-like prevaginal cae-
cum; bursa large, banana-shaped, located
partly against mucous gland, partly embed-
ded in digestive gland, beneath heart. Penis
simple, thin; penial retractor usually running
beneath albumen gland, entering between
two major bundles of columellar muscle and
inserting together with muscle attachment;
often penial retractor attaches to floor of
lung, bifurcates or runs on top of columellar
muscle.
Free-swimming veliger larva present.

Remarks: Ignorance of the specific habitat of
the supralittoral pulmonate genus Melampus
Montfort led to taxonomic misplacement of
the group. Some early workers tried to incor-
porate habitat information in their classifica-
tion schemes and therefore early 19th cen-
tury nomenclatorial history of this genus is
related not only to shell characters but also to
knowledge of the habitats of species.

Montfort (1810) separated Melampus con-
iformis (Bruguière) [= Melampus coffeus (Lin-
naeus)] from Bulimus Bruguiere on the basis
of apertural details, and from Auricula La-
marck [= Ellobium Rodding] and Scarabus
Montfort [= Руа Róding] on the basis of its
conical shape. Bulimus Bruguiere was а
large, heterogeneous assemblage of mostly
terrestrial and fluviatile mollusks. Montfort
(1810: 320) seems to have had reliable infor-
mation about the marine habitat of Melampus
(M.) coffeus, for he specified (1810: 20), “Ce
mollusque est marin, il vit sur les côtes de
Cayemne, et principalement contre le rocher
du Conétable, qui est en avant de la rade.” It
is also probable that Montfort had access to
live or preserved material, although in the
description he did not mention the external
appearance of the animal. Only such an ob-
servation would justify the choice of the ap-
propriate name Melampus, meaning black
foot, a conspicuous characteristic of the type
species.

Lamarck, apparently unaware of Mont-
fort's work, also used shell shape and habitat
in his classification. In 1812 he used the ver-
nacular Conovule, Latinized by him in 1816 to
Conovulus, for those fluviatile shells with the
outer lip simple and sharp, which he previ-
ously had included under Auricula. Upon be-
ing informed that the animals in question

248 MARTINS

were terrestrial, however, Lamarck (1822:
136) suppressed Conovulus, reuniting those
species with Auricula. Nevertheless, the name
Conovulus continued to be used occasionally,
either emended or as originally spelled (Beck,
1837; Anton, 1839; Gray, 1840; Clark, 1850,
1855).

Lowe (1832) was convinced that the genus
Melampus Montfort should be included
within the marine pectinibranchs, and he
stated, wrongly, that his Melampus aequalis
[= Ovatella aequalis] had branchial respira-
tion. Lowe knew of Montfort's comments on
the marine habitat of the type species,
Melampus (M.) coffeus, but he listed the latter
species, without justification, among Species
incertae: huc forsan referendae [Uncertain
species; perhaps to be referred to this place].
It was this misidentification that led Gray
(1847a) to consider Lowe's use of Melampus
Montfort distinct and thus erroneously to
designate a type species.

The genus Melampus 1$ the predominant
West Indian ellobiid group because of the
number of species and abundance of individ-
uals. Of the 18 recognized ellobiid species
belonging to ten genera, seven are in Melam-
pus. The very numerous individuals in salt
marshes (Morrison, 1951a; Martins, personal
observation) and in mangroves (Martins, per-
sonal observation) makes them very conspic-
uous occupants of those habitats.

Only two subgenera are recorded for the
Western Atlantic, Melampus s. s. and Detra-
cía Gray, which can be separated on the ba-
sis ОГ apertural morphology. In Melampus $.
s. the columellar tooth is small and the upper
parietal tooth 1$ the largest of the teeth on the
inner lip. In Detracia the columellar tooth 15
largest, usually strongly twisted, and the up-
per parietal tooth is small and hidden. Ana-
tomical differences reside mainly in the com-
paratively longer separation of the foot from
the visceral mass in Detracia, with conse-
quent elongation of the pallial and anterior
reproductive ducts. In Detracia the mantle or-
gan is conspicuously pouch-like, rather than
rounded and conforming to the general
shape of the mantle cavity.

The anatomical differences are related to
the degree of resorption of the inner whorls of
the shell. In Melampus s. s. resorption is so
extensive that less than half of the partition of
the body whorl remains (Figs. 225, 267),
whereas in Detracia at least 75% of that par-
tition remains (Figs. 302, 316, 340, 361). In
Detracia the parietal teeth appear on the in-

ner whorl as two conspicuous lamellae and
the region near the columellar tooth forms a
cavity that is occupied by the pouch-like
mantle organ.

Habitat: Ubiquitous in those habitats men-
tioned for the Melampinae.

Range: Worldwide. Most of the Western At-
lantic species live in the West Indies, but the
genus extends from Newfoundland to south-
ern Brazil.

Subgenus Melampus s.s.

Description: Shell oval-conic, spire low, eight
to 12 whorls; posterior parietal tooth stronger
than columellar tooth. Animal brownish to uni-
form black or with white markings. Medial
edge of arms of base of central tooth of radula
smooth or with very faint medial nodes. Vis-
ceral mass separated from foot by half a
whorl; mantle organ round, not pouch-like.

Remarks: Beck (1837) used Melampus B
[Beck (sic)] as a subgenus of Melampus Mont-
fort. In listing the species, however, he did not
include Melampus (M.) coffeus, the type of the
nominate genus, which he had listed under
the subgenus Conovulus B [Beck (sic)]. Gray
(1847a) considered Beck’s Melampus a valid
taxonomic name and he erroneously desig-
nated Melampus lineatus Say [= Melampus
(M.) bidentatus Say] as type species.
Melampus $. $. is represented in the West-
ern Atlantic by only two species: Melampus
(M.) coffeus (Linnaeus) and Melampus (М.) bi-
dentatus Say. In Florida, Bermuda and the
Greater Antilles, in which they overlap, the
two species show a wide range of variation in
shape and color, sometimes making their
separation difficult. Melampus (M.) bidenta-
tus can be recognized by the presence of at
least one, usually two or three marked spiral
grooves on the whorls of the spire and on the
shoulder of the body whorl. In Melampus (M.)
coffeus these grooves occur only on the first
four whorls (Figs. 230, 232, 233). In addition,
Melampus (M.) coffeus is conical, whereas
Melampus (М.) bidentatus is more ovoid. Al-
though variable in color, Melampus (M.) cof-
feus characteristically has as many as five
dark, olive-green bands on the body whorl
and usually it has a pinkish to purple patch
covering the columellar fold and the tip of the
columella. In Melampus (M.) bidentatus the

WESTERN ATLANTIC ELLOBIIDAE 249

bands are usually brownish and the col-
umella is white. Anatomical differences oc-
cur, as in the nervous system, in which the
right parietovisceral connective 1$ shorter
than that in Melampus (M.) coffeus; also in
this species the vagina is twice the length of
the posterior vas deferens, whereas т
Melampus (M.) bidentatus, although variable,
both ducts are much more alike in length.

Habitat: Melampus (M.) bidentatus is a com-
mon inhabitant of the North American salt
marshes and of the mangroves of the Florida
Keys, Bermuda and the Bahama Islands.
Melampus (M.) coffeus lives in mangroves
from Florida to Brazil. This species has not
been reported from salt marshes and it over-
laps with the former species only in man-
groves.

Range: The subgenus Melampus has a
worldwide distribution, not being restricted,
as are most ellobiid groups, to the tropics. In
the Western Atlantic the subgenus extends
from Newfoundland, Canada, to Brazil.

Melampus (Melampus) coffeus
(Linnaeus, 1758)
Figs. 218-256

Bulla coffea Linnaeus, 1758: 729 [type local-
ity unknown, herein designated to be
Barbados, West Indies; lectotype herein
selected, Linnaean collection, LSL (Fig.
218)].

Voluta coffea (Linnaeus). Linnaeus, 1767:
1187; Gmelin, 1791: 3438; Dillwyn, 1817:
505.

Bulimus coniformis Bruguiere, 1789: 339
[American coast, herein restricted to
Barbados, West Indies; location of type
unknown].

Melampus coniformis (Bruguière). Montfort,
1810: 319; Lowe, 1832: 292; C. B. Ad-
ams: 1849: 42; C. B. Adams, 1851: 186;
Shuttleworth, 1858: 73; Franc, 1968:
525.

Conovulus coniformis (Bruguière). Lamarck,
1816, pl. 459, figs. 2, a. b., Liste p. 12.

Auricula coniformis (Bruguiere). Férussac,
1821: 105; Lamarck, 1822: 141; Menke,
1830: 36; Gould, 1833: 67; Potiez &
Michaud, 1838: 202; Jay, 1839: 59;
Sowerby, 1839b: 10, fig. 298: Sowerby,
1842: 77, fig. 298; Küster, 1844: 31, pl. 4,
figs. 14-17; Reeve, 1877, pl. 7, fig. 57.

Pedipes coniformis (Bruguiere). Blainville,

1824: 245; Blainville, 1825: 325 [425], pl.
37 bis, fig. 4 [erroneously listed in plate
caption as Tornatelle coniforme].

Auricula biplicata Deshayes, 1830: 91 [type
locality unknown, herein designated to
be Barbados, West Indies; holotype
MNHNP (Fig. 220)]; Pfeiffer, 1854b: 148.

Melampus (Conovulus) biplicatus (Deshayes).
Beck, 1837: 106.

Melampus (Conovulus) coffeus (Linnaeus).
Beck, 1837: 106.

Auricula (Conovulus) coniformis (Bruguiere).
Anton, 1839: 48.

Auricula conoidalis (Bruguière). Sowerby,
1839b: 63, fig. 298; Sowerby, 1842: 187,
fig. 298 [referred to in text as coniformis].

Conovulus coffee (Linnaeus). Gray, 1840: 20
[error for coffea].

Auricula coniformis Férussac. Orbigny, 1841:
187, pl. 12, figs. 4-7 [plate caption incor-
rect; should be 4-7, not 1-3].

Auricula coniformis Lamarck. Reeve, 1842:
106, pl. 187, fig. 7.

Auricula olivula ‘“‘Moricand”’ Küster, 1844: 33,
pl. 3, figs. 11-33 [Bahía, Brazil; location
of type unknown].

Melampus coffea (Linnaeus). Mórch, 1852:
38; Pfeiffer, 1854b: 147; Pfeiffer, 1856a:
28; Binney, 1860: 4; Binney, 1865: 13,
fig. 15; Tryon, 1866: 8, pl. 18, figs. 7, 8;
Pfeiffer, 1876: 306; Mörch, 1878: 5;
Fischer 8 Crosse, 1880: 23, pl. 34, figs.
10, 10a; Crosse, 1890: 258; Hinkley,
1907: 71; Bequaert & Clench, 1933: 538.

Melampus coniformis (Lamarck). Shuttle-
worth, 1854b: 101.

Melampus coffeus (Linnaeus). H. & A. Adams,
1854: 9; H. & A. Adams, 1855b: 243, pl.
82, figs. 7, 7a; Binney, 1859: 162, pl. 75,
figs. 21, 25; Poey, 1866: 394; Nevill, 1879:
219; Arango y Molina, 1880: 59; Dall,
1885: 280, pl. 18, fig. 3; Dall, 1889: 92, pl.
47, fig. 3; Машу, 1922: 54; Peile, 1926:
88; M. Smith, 1937: 146, pl. 55, fig. 7, pl.
67, fig. 3 [pl. 67 copied from Вай (1885: pl.
18)]; Perry, 1940: 117, pl. 39, fig. 286;
Broek, 1950: 80; Morrison, 1951b: 8;
Dodge, 1955: 64-68 [history of nomen-
clature]; Perry & Schwengel, 1955: 197,
pl. 39, fig. 286; Morris, 1958: 228, pl. 40,
fig. 14; Coomans, 1958: 103; Morrison,
1958: 118-124 [habitat]; Nowell-Usticke,
1959: 88; Holle & Dineen, 1959: 28-35,
46-51 [shell morphometry]; Golley, 1960:
152-155 [ecology]; Warmke & Abbott,
1961: 153 [pl. 28, fig. n is of Melampus
(Detracia) monile (Bruguiere)]; Marcus 4

250 MARTINS

Marcus, 1963: 41-52 [early life history);
Marcus, 1965: 124-128 [systematics];
Marcus & Marcus, 1965a: 19-82, figs.
1-18 [distribution, ecology, anatomy);
Natarajan & Burch, 1966: 114 [chromo-
somes]; Scarabino & Maytia, 1968: 276-
278; Coomans, 1969: 82; Rios, 1970:
138; Vilas & Vilas, 1970: 91, pl. 10, fig. 21;
Princz, 1973: 183; Morris, 1973: 273, pl.
74, fig. 11; Abbott, 1974: 332, fig. 4088;
Humphrey, 1975: 196, pl. 22, fig. 27 [fig-
ured dorsal view looks very much like
Melampus (D.) monile (Bruguiere)]; Rios,
1975: 158, pl. 48, No. 764; Emerson 4
Jacobson, 1976: 192, pl. 26, fig. 26;
Berry, 1977: 181-226; Cosel, 1978: 215;
Rosewater, 1981; 161; Rehder, 1981:
646, fig. 362; Heard, 1982: 20, fig. 15;
Mahieu, 1984: 314; Jensen 8 Clark, 1986:
457, figured.

Melampus (Tralia) olivula (Kúster). H. & А. Ad-
ams, 1854: 11.

Melampus olivula (Küster). Pfeiffer, 18546:
147; Pfeiffer, 1856a: 23; Pfeiffer, 1876:
304; Lange de Morretes, 1949: 122;
Morrison, 1951b: 8.

Melampus biplicatus (Deshayes).
1856a: 21; Pfeiffer, 1876: 303.

Melampus caffeus (Lamarck) (Linnaeus). Dall,
1883: 322 [misspelling].

Melampus caffeus (Linnaeus).
1889: 68 [misspelling].

Melampus coffea, var. microspira Pilsbry,
1891: 320 [Progreso, Yucatán, Mexico;
holotype ANSP 61471 (Fig. 221)].

Melampus (Melampus) coffeus (Linnaeus).
Dall 8 Simpson, 1901: 368, pl. 53, fig. 13;
Thiele, 1931; 467; Zilch, 1959: 65, fig.
211; Vokes & Vokes, 1983: 6, pl. 22, fig.
18

Melampus coffeus coffeus (Linnaeus). C.W.
Johnson, 1934: 159.

Melampus (Melampus) coffea (Linnaeus). Al-
tena, 1975: 86, pl. 8, fig. 8; Gibson-Smith
8 Gibson-Smith, 1982: 116, fig. 1.

Pfeiffer,

Simpson,

Description: Shell (Figs. 218-233) to 23 mm
long, ovate-conic, solid, shiny to dull, whitish
to dark brown with olive tones, sometimes
monochrome, rarely with irregular axial mark-
ings, generally with as many as five olive
green to brown bands on body whorl, the one
just below shoulder markedly consistent;
pinkish to dark brown patch usually covering
tip of columella and columellar tooth. Umbil-
ical excavation visible in large specimens.

Spire low, whorls nine to 12, flat; the first
three and one-half to four whorls of teleo-
conch dark brown, spirally pitted; four rows
of pits in first whorl, gradually disappearing at
a rate of about one a whorl; remaining whorls
smooth or marked with very fine superficial
cords, not related to the preceding pits; body
whorl about 90% of total length, carinate at
shoulder, near its broadest point, smooth ex-
cept around columella, which 15 striated. Ap-
erture narrow, broadening anteriorly, subax-
ial, averaging 93% of body whorl length;
inner lip with small, oblique columellar tooth,
two parietal teeth, posterior one moderately
strong, perpendicular to columellar axis, an-
terior one minute, sometimes fused with pos-
terior one, sometimes absent; rarely with ad-
ditional parietal denticles above posterior
parietal tooth; outer lip sharp, with 13 to 18
even, white internal riblets not reaching edge.
Inner whorls greatly resorbed, partition ex-
tending into only half of body whorl (Fig. 225).
Protoconch smooth, translucent, brownish
(Figs. 230-233).

Radula (Figs. 234-251) with formula [32 +
(1 + 26) + 1 +(26 + 1) + 32] x 113. Base of
central tooth approximately same width as
that of lateral teeth, triangular, laterally con-
stricted on first third; crown small, posterior
edge with medial depression; mesocone
small, sharp; ectocones very small or absent.
Lateral teeth 20 to 36; crown strong, broadly
triangular, one-third of total length of tooth;
irregularities on crown give it tricuspid ap-
pearance, but distinct endocone or ectocone
not present. Marginal teeth 28 to 38; meso-
cone large, triangular, pointing medially; en-
docone small, sometimes divided into as
many as five small denticles that can give a
serrated appearance; ectocone appearing as
serrate edge in crown of first marginal tooth;
denticles becoming distinct around seventh
to tenth marginal tooth; from about 20th mar-
ginal tooth onward, ectocone becomes
cteniform with as many as eight denticles.

Animal (Fig. 252) brownish, mottled to uni-
form black. Mantle skirt broad, with numer-
ous mucous Cells; right and left margins pos-
teriorly fused to form pointed lappet of
posterior aperture canal. Pallial cavity deep,
not covering entire body whorl, opening to
outside through semicircular pneumostome
on right side of mantle skirt. Rectum delimits
right side of pallial cavity; bilobed anal gill
flanking rectum just posterior to anus. Long,
weakly developed hypobranchial gland just
to left of rectum. Kidney contiguous with hy-

WESTERN ATLANTIC ELLOBIIDAE 251

FIGS. 230-233. Melampus (M.) coffeus, lateral and top views of spire and protoconch. (230, 231) South
Mastic Pt., Andros Island, Bahamas. (232, 233) Tucacas, Venezuela. Scale, Fig. 231, 500 um; all others,
1 mm.

pobranchial gland, white, long, narrow, with
incomplete transverse foldings forming lon-
gitudinal medial atrium; kidney opening pap-
illose, emptying into mantle cavity just pos-
teriorto dorsal pneumostomal gland. Grayish,
white-spotted pneumostomal glands in front
of kidney, one on roof, other on floor of pallial
cavity. Mantle organ in anterior left corner,
well developed, dark brown to black. Heart
transparent, posterior to kidney and mantle
organ; posterior ventricle gives off short aorta
that branches anteriorly and posteriorly; an-
terior aorta large, passing beneath crop,
crossing to right over columellar muscle and
under right parietovisceral connective, and
emptying into large pedal sinus; posterior
aorta branches to digestive gland, intestine,
stomach and ovotestis; some blood collects
in wide circular vein of mantle skirt and passes
to pulmonary vein that opens into auricle; rest
of blood passes through kidney and mantle
organ, joining pulmonary vein at entrance of
auricle. Stomach (Fig. 253) as in subfamily.

Reproductive system (Fig. 254) basically
as described under Melampus s. I.; vagina
about one and one-half times length of body
whorl; posterior vas deferens about half the
length of vagina.

Nervous system (Fig. 255) having cerebral
ganglia joined by thick cerebral commissure,
usually heavily wrapped in connective tissue;
cerebral commissure shorter than width of a
single cerebral ganglion. Ten pairs of nerves
originate on cerebral ganglia; from anterior to
posterior, they are: large tentacular nerve in-
nervating tentacles; ocular nerve to eyes;
peritentacular nerve going to base of tenta-
cles; anterior labial nerve innervating sides of
mouth; thicker medial labial nerve with one
branch to sole of labial palps and two
branches to lips and right medial labial nerve
sends a branch to penis (penial nerve); pos-
terior labial nerve innervating ventral portions
of mouth; long cerebrobuccal connective;
thick cerebropedal connective; thin statocyst
nerve; and thick cerebropleural connective.

252 MARTINS

m ь

ЖА CK
To (7 te
A nt

FIGS. 234-242. Melampus (M.) coffeus, radula, Hungry Bay, Bermuda, sl 14.1 mm. (234) General view, left
half. (235, 236) Left lateral teeth. (237, 238) First left marginal teeth. (239) Central and first right lateral teeth.
(240, 241) Last left marginal teeth. (242) Last right marginal teeth. Scale, Fig. 241, 500 um; all others, 50 um.

WESTERN ATLANTIC ELLOBIIDAE 253

FIGS. 243-250.

254 MARTINS

CIE

23L T 1M

10M 11M 12M 18M 19M 25M 26M 27M

FIG. 251. Melampus (M.) coffeus, radula, Mullet
Key, Florida. Scale 10 um.

Buccal ganglia small, joined by short, thin
buccal commissure from which unpaired
pharyngeal nerve leaves, embedding into
buccal bulb just anterior to radular sac; a pair
of esophageal nerves leave medial anterior
portion of ganglia, splitting immediately into
anterior and posterior esophageal nerves, the
latter running posteriorly on sides of esoph-
agus; cerebrobuccal connectives securely
attached to sides of buccal bulb, at which
they emit a lateral buccal nerve; the salivary
gland nerve branches off cerebrobuccal con-
nectives and enters salivary ducts near junc-
tion with buccal bulb.

Pleural ganglia small, lacking nerves di-
rectly associated with them other than cere-
bropleural, cerebropedal and pleuroparietal
connectives; thin cutaneous lateropleural
nerve extends from lower posterior portion of
pleuropedal connective to lateral mid-foot re-
gion; left pleural ganglion about 30% of size
of right pleural ganglion; left pleuroparietal
connective twice length of right one.

Parietal ganglia unequal. Left ganglion half
size of right one, with fewer nerves; mantle
skirt artery nerve leaving ganglion anteriorly,
entering mantle skirt on left side, running
along artery, sending branch into dorsal por-
tion of mucous gland; thin external pallial
nerve going to posteroventral section of

mantle skirt; parietocutaneous nerve with
branch to origin of posterior left bundle of
columellar muscle; internal pallial nerve to
mantle skirt, bifurcating to left side and to
posteroventral portion. Nerves from right pa-
rietal ganglion are: external pallial nerve orig-
inating ventrally near pleural ganglion, cross-
ing beneath other right parietal nerves and
ramifying in floor of lung; thick pneumo-
stomal nerve following floor of pneumostome
and branching to innervate lips of pneumos-
tome; medial pallial nerve arising above
pneumostomal nerve and branching into
mantle skirt; internal pallial nerve arising
above pneumostomal nerve and branching
into mantle skirt; internal pallial nerve, closely
associated with pneumostomal nerve, going
to roof of mantle cavity; and aortic nerve in-
nervating wide aorta.

Visceral ganglion about same size as right
parietal ganglion, giving off these nerves:
thick pallial cutaneous visceral nerve crosses
from left to right to innervate mantle lappet;
thinner anal nerve goes to lower pneumos-
tomal gland and anal region; genital nerve
gives off thin branch to columellar muscle,
extends along posterior vas deferens, sends
branch to albumen and mucous glands and
continues to ovotestis; and thinner columel-
lar muscle nerve arises to right of genital
nerve and penetrates columellar muscle.

Pedal ganglia united in front by pedal com-
missure and posteriorly by thin subpedal
commissure. There are seven pairs of pedal
nerves: anteromedian pedal nerve goes for-
ward to mid-ventral foot; anterolateral pedal
nerve runs laterally and anteriorly; anterior
and posterior cutaneous pedal nerves go
midlaterally to wall of foot; posterolateral
pedal nerve runs posterolaterally; postero-
median pedal nerve goes to posterior mid-
ventral section; and posteropedal nerve goes
to posterior ventral portion of foot. Thick
cerebropedal and thinner pleuropedal con-
nectives insert close to each other on anterior
and lateral margins of ganglion respectively;
thin statocyst nerve inserts just above cere-
bropedal commissure; thin left pharyngeal
retractor muscle nerve originates posterior to

FIGS. 243-250. Melampus (M.) coffeus, radula. (243) Laguna Rincon, Bahía de Boquerón, Puerto Rico, sl
2.33 mm; central tooth hidden by tricuspid first lateral teeth. (244) Laguna Rincón, Bahía de Boquerón,
Puerto Rico, sl 2.33 mm. (245, 246) Laguna Rincón, Bahía de Boquerón, Puerto Rico, sl 3.48 mm. (247-249)
Shore of Millars Road, New Providence, Bahamas, sl 4.63 mm. (250) Left lateral teeth, with articulation of
medial node of base of one tooth with crown of next tooth, Punta Arenas, Puerto Rico, sl 19.9 mm. Scale

50 um.

WESTERN ATLANTIC ELLOBIIDAE 255

| рип
\ | gn
ру4 апп v

mL "pn ра!
FIG. 252. Melampus (M.) coffeus, anatomy. Right and anterior sides of mantle cavity cut, roof of lung

reflected to left; neck incised longitudinally, neck skin reflected laterally; insertion of penial retractor muscle,
columellar muscle and anterior aorta cut; floor of mantle removed, organs cleaned of connective tissue and

slightly separated. Scale 1 mm.

({ _avd

pe... u
br

FIG. 254. Melampus (M.) coffeus, reproductive sys-
tem, Grassy Key, Florida. Scale 1 mm.

FIG. 253. Melampus (M.) coffeus, stomach, Florida.
Scale 1 mm. sue wraps connectives associated with pedal
ganglia, statocyst nerve and proximal portion
of radular muscle nerve; right pharyngeal re-
tractor muscle nerve and associated connec-
tive tissue attach to posterior right side of

statocyst nerve, follows the latter forward
buccal bulb. Round statocyst with numerous

and, halfway, turns posteriorly and inserts in
radular muscle; wide sheet of connective tis-

256 MARTINS

pmpn pg st plpn
ppn |

pcpn

й
/

/ y + , NS: \
+ 4 ` ` `

an, ipan\ pnn“ ‘арт:
/ = `

N

plprc” plpc épan

|

// alpn phmn
-- ampn man

-

mpan 'prg plg срс pin pen “min ‘ptn

FIG. 255. Melampus (M.) coffeus, central nervous system, Grassy Key, Florida. Scale 1 mm.

statocones on anterodorsal surface of each
pedal ganglion.

Remarks: Dodge (1955) gave a detailed ac-
count of the nomenclatorial history of Melam-
pus (M.) coffeus (Linnaeus). The description
provided by Linnaeus (1758) was so brief and
general that it could be applied to almost any
species of Melampus. The specific charac-
ters mentioned were the aperture dentate on
both sides and conical shell, characters com-
mon among ellobiid species. A synonymy
was not given, references to any illustration
were not cited, and the locality was unknown.
In his copy of the twelfth edition of the Sys-
tema Linnaeus added a manuscript reference
to Lister (1770: fig. 59) (Dodge, 1955). Al-
though the abbreviation “Barb” [for Barba-
dos] is written between figures 59 and 60,
plate 834, of Lister, the sketchy representa-
tion of the outer lip in figure 59, chosen by
Linnaeus, suggests a species of the Indo-Pa-
cific genus Cassidula. Such was the interpre-
tation of Hanley (1855: 214), who regarded it
as being Bulimus auris-felis Bruguiére [= Cas-

sidula aurisfelis] “or some closely allied con-
gener.” Also according to Hanley, Cassidula
aurisfelis was not present in the Linnaean col-
lection although Linnaeus asserted that he
owned the specimen he used for the descrip-
tion of his Bulla coffea.

The history of the interpretation of the
name Bulla coffea exemplifies the confusion
of the various authors about the identity of
Linnaeus’ species. One of the illustrations
cited in early works for the Linnaean species
is a dorsal view of the Auricula Midae non-
fimbriata, bidens of Martini (1773, 2: 126, pl.
43, fig. 445). Although Martini's description
and figure are rather sketchy and inaccurate,
he cited in synonymy Petiver’s (1770) species
No. 493, Persicula barbadensis fasciatus. The
locality, Barbados, is within the range of the
species and agrees with Lister’s note. Chem-
nitz [1786, 9(2): 45, figs. 1043, 1044] also il-
lustrated and described under the name Vo-
luta coffea Linnaei his concept of Linnaeus’
species, referring to the tenth and twelfth edi-
tions of the Systema. Chemnitz criticized
Martini’s illustration as a “small, unimportant

WESTERN ATLANTIC ELLOBIIDAE 201

and unrecognizable figure of the present
much larger and impressive shell.” The
Chemnitz definition and illustration are
equally incorrect, however; the former men-
tions fine transverse striae, the latter closely
resembles Cassidula aurisfelis (Bruguière,
1789). An indication that Chemnitz was con-
fused about the true identity of Linnaeus'
species is the reference to the size. Cassidula
aurisfelis easily reaches 30 mm whereas
Melampus (M.) coffeus rarely surpasses 20
mm in length.

Probably with the intention of clarifying the
existing confusion Bruguière (1789) intro-
duced the name Bulimus coniformis, for
which he provided a fairly accurate descrip-
tion. He cited in the synonymy Lister's figure
59, Linnaeus (1758, 1767) and Martini’s fig-
ure 445. He rejected, however, Chemnitz'
figure, which shows an apertural view.

The thirteenth edition of the Systema nat-
urae (Gmelin, 1791: 3438) also failed to re-
solve adequately the question of the identity
of Melampus (М.) coffeus (Linnaeus). Gmelin
listed Voluta coffea, and the references to the
synonymy included Lister's figure 59, Fa-
vanne's (1780) figure H7 [miscited as fig. 47],
Martini’s figure 445 and Chemnitz’ figures
1043, 1044. Two pages before (p. 3436)
Gmelin introduced Voluta minuta, for which
he cited the same references to Lister and
Martini. He described two color patterns,
dark with two to six white bands, or white
with four alternating yellow and coffee-col-
ored bands. These features are not surpris-
ing, considering the variation in color that
characterizes the group. Reference to the
“three ribs on the outer lip” raises doubt
about the relationship of Voluta minuta to
Melampus (M.) coffeus, because the latter
species has many (13-18) riblets inside the
outer lip. Voluta minuta has been cited fre-
quently as junior synonym of Melampus (M.)
coffeus. Given both the impossibility of iden-
tifying Gmelin's species from the original de-
scription, and the ambiguity of the illustra-
tions cited in the synonymy, however, the
name Voluta minuta of Gmelin must be
treated as a nomen dubium.

Róding (1798: 106) introduced the names
Ellobium inflammatum and Ellobium barba-
dense and referred both to Voluta coffea
(Linnaeus). No locality was given for the
“Banded Midas ear,” Ellobium inflammatum,
and the additional references are the Lister
figure 59 and the Chemnitz figures 1043,
1044, already discussed and considered in-

conclusive. The only reference given for Ello-
bium barbadense was the unidentifiable Mar-
tini figure 445. Róding did not provide any
description and, as noted, the references
given are inconclusive. For this reason, Ello-
bium inflammatum and Ellobium barbadense
are to be considered nomina dubia.

According to Dodge (1955) Melampus (M.)
coffeus is not described in the Museum Ulri-
cae, and specimens of it are not in the
Queen's collection at Uppsala. Inspection of
the type material at the Linnaean Society of
London revealed a mixed lot of 12 specimens
representing two species. A label note by S.
P. Dance from 1963 attributes to Hanley the
selection of these 12 unmarked specimens.
Seven specimens are Melampus (D.) monile
(Bruguière), four represent a smaller form of
Melampus (M.) coffeus (Linnaeus) (Fig. 219)
and the twelfth specimen, nearly twice the
size of any of the others, conforms to Ме/ат-
pus (M.) coffeus (Linnaeus) of all authors (Fig.
218).

With all the inconsistencies in the descrip-
tions and figures applied by the early authors
to Bulla coffea Linnaeus, this name, as
Dodge (1955: 67) remarked, would be treated
as nomen dubium were it not for the exis-
tence in the Linnaean collection of a speci-
men that must be considered the ostensible
type. Equally important is the fact that the
name Melampus (M.) coffeus has been in
general use since 1854 and to remove this
well-established name would create unnec-
essary confusion in the literature. To prevent
further confusion, the large specimen in the
LSL (Fig. 218) is herein designated lectotype.

Another source of confusion was the
failure of some authors to distinguish be-
tween Melampus (M.) coffeus (Linnaeus) and
Melampus (M.) bidentatus Say. The geo-
graphical ranges of these two species over-
lap in Bermuda, most of Florida, the Gulf of
Mexico and the western Greater Antilles.
Melampus (M.) bidentatus is highly variable,
sometimes resembling the highest-spired
Melampus (M.) coffeus in shape, size and
color. The most distinctive characteristic, as
noted by Morrison (1958, 1964), is the ab-
sence of incised lines on the shoulder of the
body whorl of Melampus (M.) coffeus. This
characteristic can be used only in individuals
having more than four whorls. In fact, the first
three and one-half to four whorls of both spe-
cies have incised lines and are indistinguish-
able in this regard (Figs. 230-233). Binney
(1859) and Tryon (1866) perceived this differ-

258 MARTINS

ence, describing Melampus (M.) coffeus as
having microscopic revolving lines, as op-
posed to Melampus (M.) bidentatus, which
has revolving striae. Lines are very weak
cords whereas striae are finely impressed
spiral depressions. The lines in Melampus
(M.) coffeus are not incised and they do not
correspond to the continuation of the stria-
tions or rows of pits on the first whorls.

Failure to distinguish between Melampus
(M.) coffeus and Melampus (M.) bidentatus is
obvious in the work of Holle & Dineen (1959).
One of their conclusions is the hypothesis
that Melampus (М.) coffeus and Melampus
(M.) bidentatus are but subspecies. Natarajan
& Burch (1966), on the basis of chromosomal
counts, stated that these species hybridize. |
doubt, however, that they were dealing with
both species, but rather with two forms of the
variable Melampus (M.) bidentatus. The sup-
posed Melampus coffeus used in their re-
search was from Jekyll Island, Georgia. In-
spection of four major museum collections of
the east coast of the United States failed to
yield any record of Melampus (M.) coffeus
north of Florida. It is doubtful, therefore, that
this species was involved and the suggestion
that Melampus (M.) bidentatus and Melam-
pus (М.) coffeus hybridize is probably unten-
able.

Deshayes (1830) introduced Auricula bipli-
cata, of unknown locality. Beck (1837) re-
ported Deshayes' species as living in Amer-
ica and Binney (1859), probably on account
of Beck's reference to the name biplicata,
treated Auricula biplicata as a synonym of
Melampus (M.) bidentatus, without comment.
In the original description Deshayes (1830:
91) stressed that his species should not
be confused with those under the section of
the Conovules. In comparing Auricula bipli-
cata with Auricula coniformis (Bruguiere)
[= Melampus (M.) coffeus], Deshayes pointed
out that, although comparable in size [22 x 12
mm], his species differed from Bruguiere’s
species in color, shape and apertural denti-
tion. The holotype of Auricula biplicata
(Fig. 220) lacks the incised grooves on the
shoulder of the body whorl, characteristic
of Melampus (M.) bidentatus. The uniform
brownish color, the slender shape, the
strong, whitish posterior parietal tooth and
the greatly reduced anterior parietal tooth are
well within the range of variation that | have
observed in Melampus (M.) coffeus. | col-
lected in Florida specimens that conform
with Deshayes' species. | have designated

herein Barbados as the type locality for Au-
ricula biplicata Deshayes to avoid confusion
with a large form of Melampus (M.) bidenta-
tus, also found in Florida, which Pfeiffer
(1853b) described as Melampus gundlachi
(Fig. 271).

Melampus olivula (Kúster) is a Brazilian
morph of Melampus (M.) coffeus character-
ized by the absence of the first parietal tooth
(Fig. 228). Although Küster (1844: 33) referred
to an alleged publication by Moricand in
“Mémoir de la Société de Genève. VIII”, |
failed to see such a reference in the place
indicated. Accepting Pfeiffer's (1856a) au-
thority, who attributed Auricula olivula to
Moricand “ex citationibus auctorum,” one
can conclude that Moricand's publication of
the name is doubtful or obscure, and Kúster
(1844) is to be credited for its introduction.

As pointed out under the remarks for the
subgenus, the radular morphology of Melam-
pus changes considerably during the early
growth of the animal (Figs. 243-249). Among
the most noticeable differences are the wide
base of the central tooth bearing conspicu-
ous nodes on the inner edge of the arms and
the clearly tricuspid first lateral tooth with ec-
tocone and endocone that becomes serrated
in larger juveniles. In juveniles, as in adults,
the transitional tooth is characterized by a
shortening of the base and the presence of
an ectocone. The marginal teeth also have an
endocone. The number of cusps on the ec-
tocone can vary from row to row in the same
positional tooth.

A unique feature of the Melampinae is the
mantle organ. This squarish, black structure
was first mentioned by Koslowsky (1933:
178) for Melampus boholensis H. 8 A. Ad-
ams, and by Marcus & Marcus (1965а: 31) for
Melampus (М.) coffeus. Although of unknown
function, this highly vascularized organ prob-
ably is excretory or lymphatic, according to
the latter authors.

Another structure, here called the bilobed
anal gill, was described by Koslowsky (1933)
and by Marcus & Marcus (1965a) as a pair of
anal glands. It consists of two small, pro-
fusely ciliated tubular structures flanking the
anus and isolated from the pallial cavity. Ac-
cording to Marcus & Marcus (1965a: 38) this
structure, consisting of epithelial cells and
subepithelial secretory cells, probably se-
cretes some mucilaginous substance that
helps in holding the fecal pellets together and
in lubricating the anal groove. Renault (1966)
described the same structure in Cassidula la-

WESTERN ATLANTIC ELLOBIIDAE 259

brella Deshayes, but considered it а gill. п
the Pythiinae, to which Cassidula belongs,
and in the Pedipedinae | observed it to be a
single, well-developed, highly folded struc-
ture, closely resembling a gill. Renault noted
that this structure is at the bottom of a de-
pression in the mantle-skirt groove and con-
cluded that such a depression is comparable
to a reduced pallial cavity, seen in the ontog-
eny of some pulmonates and retained in
adult ellobiids. Although not denying its glan-
dular character, | am inclined to consider this
structure in Melampus (M.) coffeus as a gill
because of its homology with identical or-
gans in the Pedipedinae and Pythiinae.

The connectives of the visceral nerve ring
are short, but readily identifiable and of un-
equal length. In this respect Marcus 8 Mar-
cus’ illustration (1965a, pl. 2, fig. 7) of the
central nervous system of Melampus (М.)
coffeus is inaccurate in not showing evidence
of those connectives, a condition never
found in the ellobiids.

Habitat: Melampus (M.) coffeus, an inhabit-
ant of mangroves, lives at and above the
high-tide mark but can also occur intertidally,
commonly gathering on mangrove roots and
propagules above water level at high tide, but
descending to the muddy ground at low tide.
They prefer shady places (Marcus 8 Marcus,
1965a) and seem to be more active at night
(Golley, 1960).

Range: Bermuda; Florida to West Indies;
Gulf of Mexico, Central America south to Bra-
zil (Fig. 256). Scarabino 8 Maytia (1968) re-
ported this species from Uruguay, where four
beach specimens were collected; the authors
suggested that they might have been carried
by currents from nearby Brazil.

Specimens Examined: FLORIDA (ANSP
56829): Indian River (USNM 758220); S of
Sebastian Inlet (MCZ 143993); Coconut
Grove (MCZ 291328, 291330; USNM
603112); Miami (USNM 700802); Biscayne
Bay (USNM 603116); Brickell Hammock, Bis-
cayne Bay (MCZ 291325); Homestead (MCZ
291097); Card Sound (ANSP 84403); Turner
River, Card Sound (ANSP 93430); Barnes
Sound (MCZ 291095); Middle Key (USNM
338339); McGinty Key (ANSP 139532); Key
Largo (ANSP 194009; MCZ 246700, 291324;
USNM 529249, 603118); Tavernier (ANSP
325296; MCZ 201659); Tavernier Creek
(USNM 667400; A.M.); Tavernier Key (USNM

45

30

CE EEE E ee eel

— 90 15) 60. 45 30

FIG. 256. Melampus (M.) coffeus, geographic dis-
tribution. Open circle, locality from literature.

492563); Snake Creek (MCZ 291012); Snake
Key (ANSP 1054); Plantation Key (MCZ
199340, 291016; A.M.); Windley Key (USNM
603103); Indian Key Fill (A.M.); Lignumvitae
Key (ANSP 156326); MCZ 75605); Mate-
cumbe Key (USNM 822252); Lower Mate-
cumbe Key (USNM 700764); Rabbit Key
(ANSP 88135); Long Key (MCZ 176170;
A.M.); Grassy Key (МСУ 201673; A.M.);
Crawl Key (MCZ 291020; A.M.); Key Vaca
(MCZ 201672); Marathon (MCZ 153253);
Knight Key (A.M.); Bahia Honda Key (ANSP
89551;USNM 269777b); No Name Key (MCZ
142470); Little Pine Key (USNM 681643); Big
Pine Key (ANSP 89552; MCZ 201651; USNM
597455; A.M.); Ramrod Key (MCZ 291021);
Pavilion Key (ANSP 93431); Pelican Key
(MCZ 3967; USNM 39829); Key West (MCZ
201666; USNM 60754, 668244); Marquesas
Key (ANSP 73711); Mangrove Key (ANSP
365478); Flamingo Key (ANSP 294311; MCZ
235501, 275576, 291098; USNM 672441);
Cape Sable (ANSP 56825; USNM 603117);
East Cape, Cape Sable (MCZ 291017);
Sandy Key (USNM 603121); Lossman River
(ANSP 132368); Everglades City (MCZ
291096); Seminole Point (ANSP 105431);
Horr’s Island, Ten Thousand Islands (USNM
381325); Fakahatchee Key (ANSP 88139);

260 MARTINS

Royal Palm Hammock (MCZ 257250; USNM
492561); Marco Island (MCZ 201687,
291011; USNM 381332); Blue Hill Island,
near Goodland Point (ANSP 88134); Little
Marco (ANSP 93429); Gordon Pass (USNM
603098); Naples (MCZ 178104, 201661;
USNM 667406); Bonita Springs Beach (MCZ
201680, 291015); Bonita Beach (USNM
334075); Cayo Tuna, S. Carlos Bay (ANSP
106299); Carl E. Johnson Park, Little Carlos
Pass (A.M.); Mound Key, 2 km N of south end
of Estero Island (MCZ 201657); Dog Key, 1.5
km NW of middle of Estero Island (MCZ
201648); Starvation Key (ANSP 130057); Fort
Myers (ANSP 183181); Punta Rassa (ANSP
45075, 56826, 140849; MCZ 84097, 291018;
USNM 37601, 513900, 492573, 700853;
A.M.); Sanibel Island (ANSP 91378, 170651;
MCZ 13702, 55773, 201655, 291024; USNM
611799); Wulfert, Sanibel Island (МСА 13703,
13803); Pine Island (ANSP 93387); Pineaire,
Pine Island (MCZ 291023); Bokeelia (MCZ
291025; A.M.); Captiva Island (MCZ 291013;
USNM 513901); Blind Pass, Captiva Island
(MCZ 201658); Boca Grande, Gasparilla Is-
land (ANSP 142272); Little Gasparilla Island
(ANSP 131405); Punta Gorda (ANSP 45076);
Charlotte Harbor (USNM 592308); Nokomis
(ANSP 180747); Siesta Key (USNM 669348);
Sarasota (MCZ 201679; USNM 30625,
487314a); 45 km N of Sarasota (ANSP
294315); Long Boat Key (MCZ 201686); be-
tween Palma Sola and Cortez (MCZ 291014);
Palmetto (A.M.); Manatee River (ANSP
56832; MCZ 3968, 201664; USNM 492560);
Big Bend Road [Rt. 672] (A.M.); Tampa Bay
(MCZ 55757, 70569, 201645, 201654,
201664; USNM 37602, 73210, 193362,
196349a, 504481, 711484); Small Island
(MCZ 91358); Mullet Key (ANSP 76699;
USNM 652406; A.M.); Tierra Verde (MCZ un-
catalogued); Shell Key (USNM 466206,
466288a); key 2 km S of Pass-a-grille (MCZ
56177); Pass-a-grille (ANSP 148522; MCZ
138939, 162640); Bird Key (ANSP 134318;
MCZ 71009, 71595, 104946; USNM 36896,
37600); Terra Ceia (MCZ uncatalogued;
USNM 124285); Pinellas Point (MCZ uncata-
logued); St. Petersburg (USNM 341721a,
466195, 663066); SE of Gulfport (MCZ
201665); Gulfport (MCZ 88789); Bocca Key
(USNM 75409); Boca Ciega Bay (ANSP 9568;
MCZ 291329); Sand Key (ANSP 128525);
Clearwater Island (ANSP 9354, 176363; MCZ
105461; USNM 611786); island in Clearwater
Bay (ANSP 149851); Anclote River (A.M.);
Cedar Key (ANSP 362803; MCZ 201660;

USNM 27918). BERMUDA (ANSP 158807;
USNM 11421, 94432a, 173642, 228686): in-
let E of biological station, St. George's (MCZ
64705); Hungry Bay (ANSP 85712; MCZ
24247; А.М.); BAHAMA ISLANDS (ANSP
56830): GRAND BAHAMA ISLAND (ANSP
374525): Gold Rock Creek (ANSP 369339);
Running Mon Canal (ANSP 369778); Eight
Mile Rock (ANSP 173255); Hawksbill Creek
(ANSP 176351; MCZ 116679); GREAT
ABACO ISLAND (ANSP 362802, 362804;
MCZ 201683; USNM 492564, 492591): Witch
Point (ANSP 299482); Bootle Bay (ANSP
371879); Cherokee Sound (MCZ 133101);
BIMINI ISLANDS: Alicetown, North Bimini
(MCZ 144186); BERRY ISLANDS (MCZ
291332); ANDROS ISLAND (USNM 492572):
South Mastic Point (A.M.); Stafford Creek
(ANSP 189566); Mangrove Cay (ANSP
94525; MCZ 24137, 24138; USNM 269861,

269968b); Lisbon Point, Mangrove Cay
(USNM 269599); Linder Key (USNM
270483a); NEW PROVIDENCE ISLAND

(ANSP 184850; MCZ 85768): Nassau (USNM
160765, 467072, 568412); Old Fort (MCZ
107792); Dick’s Point (MCZ 107797, 113101,
291333); Adelaide (USNM 603873); Millars
Sound (A.M.); Millars Road (A.M.); Bonefish
Pond (USNM 618597; A.M.); Fox Hill, South
Beach (MCZ 107770); ELEUTHRA ISLAND:
Schooner Cays (ANSP 359332); Millars
Beach (ANSP 359351); ROYAL ISLAND
(MCZ uncatalogued; USNM 468114); CAT
ISLAND (ANSP 173256): Russell Creek
(ANSP 173260; MCZ 63384); Orange Creek
(ANSP 173254; MCZ 63385); Arthurstown
(ANSP 173661; MCZ 107832); Dumfries
(MCZ 107752); EXUMA CAYS: Hog Cay
(ANSP 285755); SAN SALVADOR ISLAND:
Riding Rock (USNM 360311); LONG ISLAND:
Salt Pond, Clarencetown (ANSP 189565;
MCZ 113099; USNM 589832, 590247);
Brett's Hill (MCZ 113336, 142299); Glenton's
(ANSP 173253; MCZ 113100; Cape Sta.
Maria (MCZ 113329). CUBA (ANSP 56791;
USNM 10967, 59724, 121518, 492569,
492581): Bahia de Santa Rosa (USNM
492556, 492568); Cape Cajón (USNM
492571a); Los Arroyos (USNM 492558);
Cayo Rapado (MCZ 201652); Dimas (USNM
492559a); Bahia Honda (USNM 492560); Ma-
riel (MCZ 131922; USNM 169938); Marianao
(ANSP 77006; MCZ 131953); Rio San Juan
(MCZ 127825); Cayo Cristo (MCZ 291321);
Cárdenas (MCZ 87886); Rio Yumurí (ANSP
87920, 167219; MCZ 83311, 131913,
131941, 201684); Cayo de las Cinco Léguas

WESTERN ATLANTIC ELLOBIIDAE 261

(ANSP 158050); Rancho Veloz, Sagua la
Grande (MCZ 201682); Caibarién (MCZ
131914, 131920, 131940); Cayo Francés
(MCZ uncatalogued); Muelles (MCZ 131915);
Cayo Salinas, Buena Vista Bay (MCZ
201681); Cayo Сописо (MCZ uncatalogued);
Isla de Cobos (MCZ uncatalogued); Punta
Alegre (ANSP 149212); Isla Turigano (MCZ
uncatalogued); Terraplen, Isla Turigano
(USNM 385661); S of Central Ramon (USNM
391797); Gibara (USNM 381469, 603096);
Banes (MCZ 59623); Penon el Fraile, Fraile,
Santa Cruz del Norte (USNM 807577); Playa
Cajio, Guira del Melena (USNM 803401);
Guantanamo Bay (MCZ 92675); Santiago
(USNM 373225, 603114); Tarallones de
Arena, near Santiago (ANSP 182935); Rio
Cauto (USNM 682787); Santa Cruz del Sur
(MCZ 131919, 201662); Finca, Sabanalmar
(MCZ uncatalogued); Cienfuegos Bay (ANSP
106093); Alto del Caracol (ANSP 222629);
Cienfuegos Bay, 1 km E of La Milpa (MCZ
uncatalogued); Cayo Blanco (ANSP 157942);
Batabanó (ANSP 93714; USNM 603115); La
Coloma (MCZ 84884, 131949); Punta de
Напа (MCZ 201668); Isla de Pinos (MCZ
48079, 48080). JAMAICA (ANSP 56822; MCZ
291233, 291318, 291322; USNM 94743,
492562, 492570): Green Island Harbor
(USNM 440805); Fort Clarence (USNM
433433); Montego Bay (ANSP 329154; MCZ
17452); Port Morant (USNM 375739); Cow
Bay (USNM 440974); Kingston (USNM
442610); Palisadoes (USNM 442466a); Port
Royal (USNM 395452c, 427004, 442268);
Hunt’s Bay (USNM 441673); Rock Fort
(USNM 374232); Phillipsfield (USNM
402222); Old Harbor (441009); Portland
(USNM 375680); near Portland Light, Port-
land Hills (MCZ uncatalogued); Little River
(USNM 128066); Great Goat Island (ANSP
344216); Little Goat Island (MCZ 291326);
Black River (USNM 441356); Savanna la Mar
(MCZ uncatalogued). HAITI: Fort Liberté
(USNM 426365); St. Louis (USNM 439390);
Моте Rouge (USNM 402680, 402715); Go-
nave Island (MCZ 82119; USNM 492531);
Port-au-Prince (MCZ 183920; USNM
403109, 403408, 440460, 440610, 442923);
Miragoane (MCZ 82072); Anse-à-Maissons,
Grand Cayamite (MCZ 82100); lle-a-Vache,
Soulette Bay (USNM 439191, 442850); Tor-
beck (USNM 383068, 403363, 439667,
439695b); Les Сауез (USNM 439742а);
Aquin (USNM 367358, 402839, 403255,
403561, 440163); Bizoton (USNM 439832).
DOMINICAN REPUBLIC: Monte Cristi (MCZ

57752, 291334); 19 km E of Monte Cristi
(USNM 471542); Sanchez (USNM 307261);
Rio Tapion, Puerto Libertador (USNM
618639); Puerto Plata (MCZ 90785, 291317,
291320); Santa Bárbara de Samaná (ANSP
173257; MCZ 57754); Sanchez (ANSP
173252; MCZ 57338); Isla La Matica, Playa
Boca Chica, E of Santo Domingo (R.B.). PU-
ERTO RICO: San Juan (USNM 161160,
169885); Cayo Maguey (MCZ uncatalogued);
Cabo Rojo lighthouse (MCZ 242179); Laguna
Rincón (A.M.); Boquerón Beach (A.M.); Pu-
erto Real (A.M.); Punta Arenas (A.M.); La Par-
guera (USNM 622804); Santurce (ANSP
175624); Piñones (A.M.); Culebra Island
(USNM 360536). VIRGIN ISLANDS: ST.
THOMAS (ANSP 56824, 56827; MCZ 89651,
291319; USNM 6363, 6385a): Benner Bay
(USNM 702725). ST. JOHN’S (MCZ uncata-
logued). TORTOLA (USNM 6484). ANEGADA
ISLAND (ANSP 249494; MCZ 229004). ST.
CROIX (ANSP 56831): Altons Lagoon (USNM
621394); Salt River (MCZ 110325). LESSER
ANTILLES: ST. MARTIN'S (ANSP 56821).
ANTIGUA (ANSP 109156; USNM 215049);
Fitches Creek (USNM 809739). GUADE-
LOUPE (MCZ uncatalogued). MARTINIQUE
(MCZ 56464): between Le Vauclin and Le
Francois (ANSP 253289; MCZ 229358). BAR-
BADOS (MCZ 148628). TOBAGO: Pigeon
Point (USNM 682273). TRINIDAD: S of Sho-
ran Site (USNM 608786); Caroni Swamp
(MCZ uncatalogued); Blue River (R.B.).
СУВАСАО: Schotteghat, near Willemstad
(ANSP 133971). CARIBBEAN ISLANDS:
GRAND CAYMAN ISLAND: 5 km N of George-
town (ANSP 209765). OLD PROVIDENCE IS-
LAND (USNM 687818): N of Ironwood Point
(ANSP 313209; USNM 678832, 678833; MCZ
270624). MEXICO: Tampico (USNM 219997);
Vera Cruz (USNM 769426); Rio Vinasco, Vera
Cruz (USNM 675266); Tuxpan, Vera Cruz
(MCZ uncatalogued); SE of Tuxpan, Vera Cruz
(USNM 675271); Mandinga Lagoon, Vera
Cruz (USNM 791711); Boca del Rio, Vera
Cruz (MCZ 155429); Isla de Carmen (USNM
809096); Ciudad del Carmen (USNM 702910);
Rio Champotón (MCZ 59747); Silam (ANSP
61470); Progreso (ANSP 61469, 61471);
Rio Lagartos (USNM 618635); Isla Cancún,
Quintana Roo (ANSP 285520); N end of

Ascension Bay, Quintana Roo (USNM
736142, 736691, 738632); Allen Point,
Ascension Bay, Quintana Roo (USNM

736695, 736892). BELIZE: Ambergris Cay
(ANSP 284797); Belize (ANSP 294323); Bo-
tanical Garden, Belize (USNM 426007);

262 MARTINS

Blackadore Cay (ANSP 282031); Robinson
Point (ANSP 281579); N of Punta Gorda
(ANSP 282494). GUATEMALA: Puerto Barrios
(MCZ 88877). NICARAGUA: Wounta (ANSP
97591; USNM 181854); Wounta River, near
Wounta (MCZ 14804); 16 km S of Wounta
(MCZ 137227) PANAMA (USNM 46182):
Galeta Island (USNM 703195, 732922,
732949); Toro Point, Limon Bay (USNM
732885); Colón (ANSP 107258; MCZ 45058);
Porto Bello (USNM 218173). COLOMBIA: Sa-
banilla (USNM 103175); Cartagena (MCZ
192431); Coveñas, Bolívar (USNM 364336).
VENEZUELA (MCZ 291327): S of Porlamar,
Isla Margarita (ANSP 240007; USNM 707796);
Punta Mangle, ESE of Punta Piedras, Isla Mar-
garita (MCZ 273661); La Orchila Island (USNM
656031); Carenero (784775); Tucacas (A.M.).
GUYANA: Demerara (MCZ 177296). SURI-
NAME: 16 km WNW of Paramaribo (MCZ un-
catalogued); Nickerie Strand, Zeedijk (MCZ
uncatalogued); Bigisanti (USNM 635225).
BRAZIL: Boa Viagem (MCZ 219130); Uru-
majó, Braganca, Pará (ANSP 244096); Praia
de Búzios, 20 km S of Natal, Rio Grande do
Norte (ANSP 300442); Praia Upanema, Areia
Branca, Rio Grande do Norte (ANSP 300320);
Rio Pirangi (ANSP 300343); Baía (AMNH
22434; USNM 119506,157674, 465525); Vic-
tória (MCZ uncatalogued); Rio de Janeiro
(ANSP 56828; MCZ 89650); Pinheiro Island,
Rio de Janeiro (USNM 598337). ATLANTIC
ISLANDS: FERNANDO NORONHA (MCZ un-
catalogued).

Melampus (Melampus) bidentatus
Say, 1822
Figs. 257-289

Melampus bidentatus Say, 1822: 245 [East
Florida, herein restricted to mouth of St.
John's River; type material presumed lost
(Baker, 1964); neotype herein designated
USNM 859014 (Fig. 257)]; Jay, 1839: 59;
H.& A. Adams, 1854: 10; Pfeiffer, 1854b:
147; Pfeiffer, 1856a: 45; Say in Binney,
1858: 84; Binney, 1859: 156, pl. 75, fig.
23; Binney, 1860: 4; Binney, 1865: 10,
figs. 11, 12; Тгуоп, 18668 ре 1884: 5;
Gould, 1870: 467, fig. 721; Binney 4
Bland, 1870: 286, fig. 7 [radula figured];
Pfeiffer, 1876: 316; Nevill, 1879: 219; Dall,
1883: 322; Whiteaves, 1901: 207; Morse,
1921: 21, pl. 7, fig. 46, pl. 9, figs. 46, 46a

[external anatomy and morphology];
Peile, 1926: 88; Pilsbry, 1927: 125-126;
Hausman, 1932; 541-545 [ecology];
Hausman, 1936: 127; M. Smith, 1937:
146, pl. 55, fig. 11; plo G7) igs 12 бя
copied from Dall (1885: pl. 18)]; Morton,
1955c: 127-168 [anatomy, evolution];
Holle & Dineen, 1957: 90-95 [life history);
Morrison, 1958: 118-124 [habitat]; Mor-
ris, 1958: 40, fig. 15; Holle & Dineen,
1959: 28-35, 46-51 [shell morphometry,
taxonomy]; Baker, 1964: 151; Russell-
Hunter & Brown, 1964: 143; Russell-
Hunter & Meadows, 1965: 409 [physiol-
ogy]; Russell-Hunter 4 Apley, 1966, 392-
393 [early life history]; Apley et al. 1967:
455-456 [annual reproductive turnover];
Coomans, 1969: 82; Apley, 1970: 381-
397 [life history]; Jacobson & Emerson,
1971: 64, text fig.; Russell-Hunter et al.,
1972: 623-656 [early life history]; Grandy,
1972: 106-109 [winter distribution]; Mor-
ris, 1973: 273, pl. 74, fig. 8; Abbott, 1974:
331, fig. 4087; Lesser et al., 1976: 69-77
[population density]; Emerson 8 Jacob-
son, 1976: 192, pl. 26, fig. 25; Orton,
1976: 1-57 [ecology]; Andrews, 1977:
181, figured [common name coffee
Melampus erroneously applied]; Price,
1977: 295-312 [central nervous system];
Fitzpatrick & Sutherland, 1978: 23-28
[population density]; Moffett, 1979: 306-
319 [locomotion]; Andrews, 1981: 77,
text fig.; Rehder, 1981: 645, fig. 361;
Heard, 1982: 19, fig. 15; Moffett, 1983:
950; Ridgway, 1983: 950; Thompson,
1984: 44-53 [diet]; Jensen & Clark, 1986:
457, figured.

Melampus bidentatus var. lineatus Say, 1822:
246 [Coasts of Maryland and New Jer-
sey, herein restricted to Bivalve, New
Jersey, type material presumed lost
(Baker, 1964); neotype herein desig-
nated USNM 859013 (Fig. 259)]; Say т
Binney, 1858: 85; Pfeiffer, 1854b: 147;
Pfeiffer, 1856a: 46.

Melampus obliquus Say, 1822: 377 [South
Carolina; type material lost (Binney,
1859)]; Pfeiffer, 18546: 147; Pfeiffer,
1856a: 30; Say т Binney, 1858: 27; Bin-
ney, 1860: 4; Pfeiffer, 1876: 306;
Mazyck, 1913: 2.

Auricula cornea Deshayes, 1830: 90 [New
York; location of type unknown]; Jay,
1839: 59.

Melampus (Melampus) lineatus Say. Beck,
18374107:

WESTERN ATLANTIC ELLOBIIDAE 263

Melampus (Melampus) obliquus Say. Beck,
1837: 107.

Melampus (Melampus) corneus (Deshayes).
Beck, 1837: 107.

Auricula bidentata (Say). Gould, 1841: 197,
fig. 130; De Kay, 1843: 57, pl. 5, figs. 92,
1-3; Kúster, 1844: 41, pl. 6, figs. 7-11.

Auricula jaumei Mittré, 1841: 67 [Hampton,
Virginia; location of type unknown].

Auricula obliqua (Say). De Kay, 1843: 58.

Melampus lineatus Say. Gray, 1847a: 179;
Dall, 1885: 282, pl. 18, figs. 9, 12; Dall,
1889: 92, pl. 47, figs. 9, 12; Apgar, 1891:
181, figs. 46-48; Mazyck, 1913: 2; C.W.
Johnson, 1915: 178; Maury, 1922: 55;
C.W. Johnson, 1934: 159; Webb, 1942,
pl. 11, fig. 20; La Rocque, 1953: 262;
Coomans, 1958: 103; Bousfield, 1960:
14, pl. 1, fig. 10; Coomans, 1962: 90;
Baker, 1964: 152; Baranowski, 1971:
143.

Melampus corneus Stimpson.
1851: 51; Porter, 1974: 301.

Melampus gundlachi Pfeiffer, 1853b: 126
[Cayo Blanco, Cuba: location of type un-
known]; Pfeiffer, 1854a: 147; Pfeiffer,
1856a: 20; Pfeiffer, 1876: 303: Arango y
Molina, 1880: 59; A.E. Smith, 1884: 277;
Crosse, 1890: 258; Kobelt, 1900: 229, pl.
29, figs. 1, 2; Pilsbry, 1900b: 504; Mor-
ton, 1955с: 9; Holle 8 Dineen, 1959: 28-
35, 46-51.

Melampus redfieldi Pfeiffer, 1854a: 112 [Ber-
muda; location of type unknown]; Pfeif-
fer, 1854b: 147; Pfeiffer, 1856a: 33; Bin-
ney, 1859: 170; Pfeiffer, 1876: 308;
Kobelt, 1900: 232, pl. 29, figs. 8, 9; Pils-
bry, 1900b: 504.

Melampus ? jaumei (Mittré). Pfeiffer, 1854b:
147; Pfeiffer, 1856a: 25.

Melampus bidentatus var. borealis ‘ Conrad”
Pfeiffer, 1856a: 46 [Georgia; type in
Cuming collection, fide Pfeiffer, not
found at ВММН].

Auricula gundlachi (Pfeiffer). Reeve, 1877, pl.
6, fig. 50. Non Gassies, 1869.

Auricula redfieldi (Pfeiffer). Reeve, 1877, pl. 7,
fig: 52;

Auricula bidenta (Say). Reeve, 1877, pl. 7, fig.
54 [error for bidentata, corrected in the
Index].

Melampus spiralis “Pfeiffer” Melvill, 1881:
155-173 [misidentification]. Non Pfeiffer,
1855.

Melampus coffeus var. gundlachi Pfeiffer.
Davis, 1904: 126, pl. 4, fig. 9; Maury,
1922: 55; Peile, 1926: 88.

Stimpson,

Melampus coffeus var. redfieldi Pfeiffer.
Davis, 1904: 126, pl. 4, fig. 10; Peile,
1926: 88.

Melampus coffeus var. bishopi Davis, 1904:
127, pl. 4, fig. 13 [Bermuda; lectotype
selected by Baker (1964) ANSP 86925
(Fig. 262)].

Melampus coffeus var. verticalis Davis, 1904:
127, pl. 4, fig. 12 [Bermuda; lectotype
selected by Baker (1964) ANSP 86927
(Fig. 263)].

Melampus coffeus var. alternatus Davis,
1904: 127, pl. 4, fig. 11 [Bermuda; lecto-
type selected by Baker (1964) ANSP
86926 (Fig. 264)].

Melampus coffeus gundlachi Pfeiffer. C.W.
Johnson, 1934: 159; Perry, 1940: 177.

Melampus bidentatus corneus (Deshayes).
Morrison, 1951b: 8.

Melampus bidentatus lineatus Say. Morrison,
1951b: 8; Burch, 1960a: 177-208, pl. 1,
fig. 1, pl. 4, fig. 62 [chromosomes]; Burch,
1960b: 454, fig. 5 [chromosomes]; Nat-
arajan & Burch, 1966: 111, figs. 8, 13
[chromosomes].

Melampus bidentatus bidentatus Say. Morri-
son, 1951b: 8;

Melampus bidentatus redfieldi Pfeiffer. Mor-
rison, 1951b: 8

Melampus (Micromelampus) bidentatus Say.
Morrison, 1959: 25.

Melampus bidentus Say. Porter, 1974: 300
[error for bidentatus].

Melampus (Melampus) bidentatus Say.
Vokes & Vokes, 1983: 60, pl. 22, fig. 12.

Description: Shell (Figs. 257-280) to 20 mm
long, ovate-conic to elongate-oval, solid to
thin, shiny to dull, often whitish, commonly a
uniform yellowish to dark brown, with irregu-
lar axial markings or with as many as five
transverse brown bands on body whorl. Um-
bilical excavation present. Spire low to mod-
erately high, whorls eight to 11, flat, spirally
pitted or grooved; apex frequently eroded.
Body whorl about 85% of total length, stri-
ated, with one, usually two or more incised
striae on round shoulder. Aperture subaxial,
about 90% body whorl length, narrow, with
base moderately broadening; inner lip with
small, oblique columellar tooth and two pari-
etal teeth, the posterior strongest, perpen-
dicular to columellar axis, anterior one
minute, often absent; additional parietal den-
ticles sometimes present; outer lip sharp, in-
ternal lamellae uneven, white, as many as 17,
not reaching edge and only three or four ex-

264 MARTINS

FIGS. 257-276.

WESTERN ATLANTIC ELLOBIIDAE 265

FIGS. 277-280. Melampus (M.) bidentatus, lateral and top views of spire and protoconch. (277) Hudson,
Florida. (278) Jekyll Island, Georgia. (279) Long Key, Florida. (280) South Mastic Pt., Andros Island, Baha-
mas. Scale, Fig. 279, 500 um; all others, 1 mm.

tend inward. Resorption of inner whorls ex- Stomach (Fig. 286) as in subfamily.

tensive, remaining internal partition less than
half a turn (Fig. 267). Protoconch smooth,
translucent, brownish (Figs. 277-280).
Radula (Figs. 281-285) with formula [26 +
(1 + 17) + 1 +(17 + 1) + 26] x 100. Radular
morphology the same as that of Melampus

Reproductive system basically as de-
scribed under Melampus s.l.; posterior vas
deferens about 70% of vagina length (Fig.
287).

Nervous system (Fig. 288) with left parietal
ganglion one-fourth size of right one; visceral

(M.) coffeus.
Animal as in Melampus (M.) coffeus.

ganglion slightly larger than right parietal
ganglion. Length of cerebral commissure

FIGS. 257-276. Melampus (M.) bidentatus Say. (257) Neotype (USNM 859014), from Smith collection, St.
John’s River, Florida, sl 9.2 mm. (258) Specimen figured by Binney (1865:10, fig. 11) (USNM 39818), sl 14.8
mm. (259) M. bidentatus var. lineatus Say, neotype (USNM 859013), Bivalve, New Jersey, sl 8.1 mm. (260)
Specimen figured by Binney (1865:10, fig. 12) (USNM 39818), sl 12.6 mm. (261) Cedar Island, North
Carolina, sl 18.2 mm. (262) M. coffeus var. bishopi Davis, lectotype (ANSP 86925), South Shore, Bermuda,
sl 12.1 mm. (263) M. coffeus var. verticalis Davis, lectotype (ANSP 86927), South Shore, Bermuda, sl 11.2
mm. (264) M. coffeus var. alternatus Davis, lectotype (ANSP 86926), South Shore, Bermuda, sl 11.6 mm.
(265) Specimen figured by Binney (1859, pl.75, fig. 30) as “Melampus floridanus Shuttleworth” (USNM
39835), sl 6.8 mm. (266) Crawl Key, Florida, sl 2.15 mm. (267) Knight Key, Florida, sl 14.8 mm. (268) Belize
(USNM 426007a), sl 12.0 mm. (269) Tampico, Mexico (USNM 219997а), sl 8.8 mm. (270) Myrtle Grove,
Louisiana (USNM 628753), sl 15.0 mm. (271) Long Key, Florida (USNM 193363), sl 18.8 mm. (272) Narrow
River, Wakefield, Rhode Island, sl 10.1 mm. (273) Narrow River, Wakefield, Rhode Island, sl 12.6 mm. (274)
Narrow River, Wakefield, Rhode Island, sl 9.3 mm. (275) Narrow River, Wakefield, Rhode Island, sl 10.7 mm.
(276) Narrow River, Wakefield, Rhode Island, sl 10.0 mm.

266 MARTINS

FIGS. 281-284. Melampus (M.) bidentatus, radular teeth. (281) Narrow River, Rhode Island, sl 11.1 mm.
(282, 283) Hungry Bay, Bermuda, sl 14.4 mm. (284) Skidaway Island, Georgia, sl 2.3 mm. Scale, Fig. 284,
50 um; all others, 100 um.

equals width of one cerebral ganglion; left
pleuroparietal connective twice length of
right pleuroparietal; left parietovisceral con-
nective about half the length of right parieto-
visceral. Internal pallial nerve crosses pneu-
mostomal nerve to right, anastomoses with
branch of medial pallial nerve, follows floor of
pneumostome toward aperture and turns up
to roof of mantle cavity, just to left of upper
pneumostomal gland; pneumostomal nerve
branches to mantle skirt, runs to pneumo-
stomal aperture, sending branches along the
way to mucous gland of mantle skirt, splitting
to innervate lips of pneumostome.

Remarks: Melampus (M.) bidentatus Say 1$
more variable than any other Western Atlan-
tic ellobiid. The shape of the shell (Figs. 257-
276) varies from almost globose to moder-
ately high spired and oval-elongate. The
thickness of the shell shows a geographic
gradient, increasing toward the warmer parts
of the range; Auricula cornea from New York,
described by Deshayes (1830), exemplifies

the much thinner-shelled northern variety of
Melampus (M.) bidentatus. The intergrading
color pattern ranges from monochromic to
axially striped to discretely banded, the latter
pattern being common among juveniles. п
some populations the size of some individu-
als reaches that of some gerontic Melampus
(M.) coffeus. The latter consistently has an
accentuated conic shape as opposed to the
slender profile of comparably-sized Melam-
pus (M.) bidentatus. The number of riblets on
the outer lip also shows geographical grada-
tion with a tendency toward higher counts in
the warmer parts of the range. These riblets
are of unequal size and only four or fewer
large ones penetrate deep into the aperture.

As noted in the synonymy above, Melam-
pus lineatus and Melampus obliquus, pro-
posed by Say (1822: 246, 377, respectively),
are here considered to be Melampus (M.) bi-
dentatus. The name Melampus bidentatus
was applied to the thin, monochromic east
Florida specimens, Melampus lineatus to the
smaller, banded variety of Melampus (M.) bi-
dentatus from Maryland and New Jersey and

WESTERN ATLANTIC ELLOBIIDAE 267

A 7M 8M 13M 14M 18M 19M 20M

CU AMAIA

\

13M 14M 20M 21M

8M 9M

ar

FIG. 285. Melampus (M.) bidentatus, radula. A,
Woodville, North Carolina; В, Hudson, Florida.
Scale 10 um.

Melampus obliquus to the rather thick form
with a very oblique columellar tooth from
South Carolina.

Binney (1859: 159), who had access to the
type material of Melampus bidentatus and
Melampus lineatus, noted the variability of
the species and wrote about the latter name,
“| have met with none sufficiently marked to
form a variety, much less a distinct species.”
Nevertheless, the name Melampus lineatus
has been used to designate a subspecies or,
erroneously, to substitute for Melampus (М.)
bidentatus Say (Dall, 1885). The reasons for
the use of Melampus lineatus resided in the
misplacement of Аиисште!а (Leucophytia)

FIG. 286. Melampus (M.) bidentatus, stomach,
North Carolina. Scale 1 mm.

bidentata (Montagu, 1801) in the genus
Melampus, which would have constituted a
case of secondary homonymy. Dall's confu-
sion arose from Binney's contention (1865)
that the animal of Myosotella myosotis from
America was apparently similar to Melampus
(M.) bidentatus Say, and that it did not agree
with the illustration of Alexia given by H. 8 A.
Adams (1855b, pl. 82, fig. 5). Binney did not
say in what way they differed, however. Be-
sides the color, the main difference between
the animal of Myosotella myosotis (Drapar-
naud) and that of Auriculinella (L.) bidentata
(Montagu, 1801) is that the foot of the latter
species is transversely divided (Morton,
1955b). Because Melampus has this same
characteristic, Dall probably assumed that
the Alexia myosotis referred to by Binney, like
Melampus, would have a transversely di-
vided foot. The shells of Myosotella myosotis
and of Auriculinella (L.) bidentata can be con-
fused easily and Dall's (1885: pl. 18, fig. 13)
illustration of Melampus (Leuconia) bidentata
(Montagu) is a facsimile of Binney’s (1865: 4,
fig. 4) representation of Alexia myosotis.
Say's last name, Melampus obliquus, was
applied to a form similar to Melampus (D.)
monile (Bruguiere, 1789). Say (1822: 377)
added that in the collection of the Academy
of Natural Sciences of Philadelphia there
were specimens from the West Indies which
conformed with his description of Melampus
obliquus. Binney (1859: 167), who had seen

268 MARTINS

FIG. 287. Melampus (M.) bidentatus, reproductive system. A, Narrow River, Rhode Island; B, Cedar Island,
North Carolina; C, New Smyrna, Florida; D, Hungry Bay, Bermuda. Scale 1 mm.

specimens belonging to the first two of Say's
names, noted, “It is not known what shell
Say had in view when the above description
was written. No authentic specimen was pre-
served, and no author has seen any shell
from that locality answering to the characters
laid down.” In 1865 Binney relegated Say's
Melampus obliquus to his ““Spurious species
of Melampus.” On Cedar Island, North Caro-
lina, however, | found specimens of Melam-
pus (M.) bidentatus that fit Say's description
of Melampus obliquus (Fig. 261). Melampus
obliquus Say must, then, be treated as a jun-
юг synonym of Melampus (М.) bidentatus
Say.

Pfeiffer introduced several species that
should be related to Melampus (M.) bidenta-
tus. In his comments on Melampus gundlachi
from Cayo Blanco, Cuba, Pfeiffer (1853b)
noted that their juveniles were much more
brightly colored than those of Melampus (M.)
coffeus and he did not mention incised
grooves. The elliptic-ovate shape of the shell,

however, indicates to me that Pfeiffer's name
can be treated as a synonym of Melampus
(M.) bidentatus Say.

Another of Pfeiffer's (1854a) introductions,
Melampus redfieldi from Bermuda, was dis-
tinguished from Melampus (М.) coffeus by
the presence of very compressed riblets in
the outer Пр. Because it occurs in the latter
species (personal observation), this condition
is not a useful character. The fact that Pfeiffer
referred to the striations on the shell sug-
gests that he was actually describing a spec-
imen of Melampus (M.) bidentatus.

Pfeiffer (1854b, 1856a) erroneously identi-
fied a variety of Melampus (M.) bidentatus
Say with Melampus borealis (Conrad) [= My-
osotella myosotis (Draparnaud)]. Neither
Pfeiffer's description nor the locality given
(Georgia) agrees with those of Conrad (1832).
Pfeiffer (1876: 316) later explained that he
was referring to a museum label in the Cum-
ing collection, rather than to Conrad (1832).

In 1841 Mittré described Auricula jaumei

WESTERN ATLANTIC ELLOBIIDAE 269

FIG. 287.

from Hampton, Virginia. It was said to differ
from Deshayes’ Auricula cornea in having the
inside of the outer lip consistently ribbed.
Melampus (M.) bidentatus is highly variable in
this respect (Holle & Dineen, 1959) and this
character is therefore unreliable for taxonomy.
Auricula jaumei is conspecific with Melampus
(M.) bidentatus.

Melvill (1881) mentioned Melampus spiralis
Pfeiffer from Cedar Keys, on the western
coast of Florida. Pfeiffer (1856a), however, in
his observations on Melampus spiralis, noted
that this species was made known to him by

Cuming as Melampus pallescentis Sowerby,
1839. Because Sowerby provided an incom-
plete description of the latter taxon, Pfeiffer,
based on Sowerby’s figure (1839a т Beechy:
pl. 38, fig. 28), thought that Melampus pall-
escentis Sowerby should be referred to
Melampus luteus Quoy 8 Gaimard, 1832,
from the Philippines. It appears, then, that the
Melampus spiralis of Pfeiffer was not based
on West Indian material. Melvill’s citation
(1881) of “Melampus spiralis Pfeiffer’ should
be treated as a misidentification of Melampus
(M.) bidentatus.

270 MARTINS

FIG. 288. Melampus (M.) bidentatus, central ner-
vous system. A, Narrow River, Rhode Island; B,
New Smyrna, Florida. Scale 1 mm.

Morrison (1951b) recognized several sub-
species of Melampus (M.) bidentatus: M. b.
corneus from Prince Edward Island, Canada,
to Staten Island, New York; М. b. lineatus
from New Jersey to North Carolina; M. b. bi-
dentatus from South Carolina to Florida,
Texas and the West Indies; and M. b. redfieldi
from Bermuda. The latter supposed subspe-
cies also lives in Florida (Pfeiffer, 1876, with a
question mark; personal observation) and
shows as much variation as the other forms
of Melampus (M.) bidentatus. Shell thickness
and color seem to be linked to temperature,
the thinner and paler forms being typical of
colder regions. This link would account for
the similarities between shells from Bermuda
and those from Florida (see the remarks un-
der Myosotella myosotis). Holle & Dineen
(1959) reported the continuous variation in
shell characters along the range of the spe-
cies, thus rendering Morrison's treating the
morphs as subspecies unjustifiable.

Morrison (1951b, 1964) placed Melampus
(M.) bidentatus in the subgenus Micromel-
ampus Möllendorff, 1898, which had been
erected, according to Zilch (1959), to include
the small Melampus. Only anatomical study

of the type species, Melampus nucleolus
Martens, 1865, from Amboina, will allow an
objective decision concerning the taxonomic
status of this subgenus. In any case, the ob-
served wide range of size in Melampus (М.)
bidentatus does not justify its inclusion in the
subgenus Micromelampus and my anatomi-
cal study strongly favors its inclusion in
Melampus s.s.

It appears that Say's type material, to
which Binney referred (1859), is not in the
Academy of Natural Sciences of Philadelphia
and was presumed lost (Holle 8 Dineen,
1959; Baker, 1964). Because of documented
confusion between Melampus (M.) bidenta-
tus and Melampus (M.) coffeus, neotypes
that conform to original descriptions and ac-
cepted usage are herein designated for
Melampus bidentatus and Melampus linea-
tus. Inspection of material from South Caro-
lina present in the collections of the various
museums did not provide any specimens
that could be identified with Say's Melampus
obliquus. As noted above, | found on Cedar
Island, North Carolina, a colony of Melampus
(M.) bidentatus that agree with Say's descrip-
tion of Melampus obliquus. Without ade-
quate material from the type locality, South
Carolina, and with the disuse into which
Say's name has fallen, it is inappropriate and
unnecessary to designate a neotype for
Melampus obliquus.

Habitat: The salt-marsh snail, Melampus (M.)
bidentatus, is very common in the high-tide
fringe of salt marshes and mangroves. Its
habits have been studied only in the New En-
gland area, in which this mollusk lacks con-
generic competitors. When Say described
this species (1822) he noted its abundance
and he considered it an important item in the
diet of marsh birds.

Although dependent upon the rhythm of
the tides, the salt-marsh snails are more ac-
tive during the low tides at twilight. Their diet
includes diatoms and detritus of vegetal and
animal origin (Hausman, 1936; Thompson,
1984). The animals gather to spend the win-
ter in partial hibernation (Grandy, 1972).

Apley (1970) and Russell-Hunter et al.
(1972) provided detailed studies of the early
life-history of this species. The snails become
sexually mature at a shell length of 5 to 6 mm
and have a life span of three to four years.
Copulation and hatching of veliger larvae are
closely synchronized with spring tides. Ge-
latinous masses of eggs are laid within one

WESTERN ATLANTIC ELLOBIIDAE 271

90 75 60 45 3

FIG. 289. Melampus (M.) bidentatus, geographic
distribution.

day after copulation. About 13 days later
free-swimming veligers hatch and spend
about six weeks in the plankton before set-
tling.

In the mangroves Melampus (M.) bidenta-
tus commonly lives under fallen leaves and
debris at the high-tide fringe, and is abundant
on the margins of sheltered inland lagoons. In
this habitat it coexists with Melampus (M.)
coffeus.

Range: New Brunswick, Canada, to the Flor-
ida Keys, the Gulf coast of the United States
south to Belize; Bermuda; Bahamas, Cuba,
Jamaica, Hispaniola, Tortola (Fig. 289). The
Tortola record is suspect and has not been
confirmed by recent collections.

Specimens Examined: NEW BRUNSWICK:
Buctouche Bay (MCZ 200471). PRINCE ED-
WARD ISLAND (MCZ 34048): Mount Stuart
(ANSP 106545); Bunbury (ANSP 106678,

106938). NOVA SCOTIA: Windsor (ANSP
326060); West Peswick, Halifax (MCZ
297898). MAINE: Newmeadows River (MCZ

104992); Middle Bay (MCZ 294188); Cape
Elizabeth (USNM 129331, 190466); Biddeford
(ANSP 22370, 77421, 242121; MCZ 34049,
34810); USNM 492528). NEW HAMPSHIRE:

Jackson's Landing, Durham (MCZ 193855);
Sagamore Creek (MCZ 275395). MASSA-
CHUSETTS (MCZ 201644): Ipswich (MCZ
147833); Gloucester (ANSP 102400; MCZ
201639; USNM 159104); Bass Rocks,
Gloucester (USNM 408730); Goodharbor
Beach, East Gloucester (MCZ 199906);
Manchester (USNM 398130); Danvers (MCZ
3963, 34052, 61508, 133195; USNM 504484);
Salem (MCZ 201633, 201636; USNM 73424);
Lynn (MCZ 34053; USNM 224905); Revere
(ANSP 89557); Revere Beach (MCZ 147714,
199909); Magazine, Cambridgeport (MCZ
141058; USNM 590055, 600303); Charles
River, Boston (ANSP 56837); Boston Harbor
(MCZ 141060); Neponset River, Milton (MCZ
142330); East Milton (MCZ 55146); Cohasset,
North Scituate (MCZ 139271); Duxbury (MCZ
14884, 163165, 201638; USNM 492523,
492525, 492535); Gurnet Lights, Plymouth
(MCZ 199905); Barnstable (ANSP 44599,
56840; USNM 34404, 159105, 487652);
Sandy Neck (MCZ 182392); Truro (USNM
603101); Playground Beach, Hyannis (ANSP
176264); Wellfleet (R.B.); Provincetown
(ANSP 134441; MCZ 147715, 199911; USNM
159108, 307158, 341102, 341104, 492532,
492533, 492536); Buzzard’s Bay (MCZ
199913); Eastham (MCZ 199907, 201632);
Pleasant Bay (MCZ 80561); Chatham (MCZ
34054); Eel Pond, South Chatham (MCZ
18812); South Dennis (MCZ 14547; USNM
492526); Grand Cove, South Dennis (MCZ
14548); Point Gammon, West Yarmouth (MCZ
112213); Lewis Bay (MCZ 200027); Nobska
Point (MCZ 140811); Briar Neck (MCZ
199904); Vineyard Sound (USNM 159107);
Nantucket (MCZ 34047); SE of Coskatee,
Nantucket (MCZ 167949); South Beach, Nan-
tucket Harbor (MCZ 167450); Popponesset
Beach (MCZ 182391); Waquoit (ANSP 46678);
Naushon Island (ANSP 163469; USNM
159110); Great Pond, Falmouth (USNM
660507); Woods Hole (MCZ 34051, 199908;
USNM 159106); Bathing Beach, Woods Hole
(USNM 340981, 340983, 340984; Little Sip-
pewisset Marsh, Falmouth (MCZ 294115;
A.M.); West Falmouth (ANSP 76171); Ware-
ham (MCZ 34050, 199910); Piney Point, Mar-
ion (MCZ 178105); New Bedford (ANSP
56846; MCZ 201635). RHODE ISLAND: New-
port (ANSP 56838; USNM 39814, 159109);
Nayatt (MCZ 71268, 199914; Jamestown, Co-
nanicut Island (A.M.); Seekonk River (ANSP
243912; MCZ 199912); Pawtuxet (MCZ
68940); Buttonwoods (MCZ 199915; USNM
492530); Apponaug (MCZ 151930; USNM

272 MARTINS

568388); Wakefield, Pettaquamscutt River
[Narrow River] (A.M.); Westerly (MCZ 14550;
USNM 492541, 590089). CONNECTICUT:
Stonington (ANSP 91947; MCZ 3965, 56902,
133196; USNM 858073; A.M.); Long Beach
(MCZ 231486); Oxecoset Brook (A.M.); Guil-
ford (USNM 478030, 568440); Pine Orchard
(MCZ 166061, 201630); Branford (MCZ
34055, 34925, 199916); New Haven (MCZ
291293; USNM 404295); Lighthouse, New
Haven (USNM 380818); Greenwich (ANSP
154660). NEW YORK: Harlem River (MCZ
156659, 167630; USNM 492538); Long Island
(MCZ 201637, 230929; USNM 307157,
492527, 492542); Greenport (MCZ 161271;
USNM 407784); Sand's Point (ANSP 145457);
Sea Cliff (ANSP 43785); Oyster Bay (USNM
307154); Cold Spring (USNM 504486); Towo
Point (USNM 694036); Orient (ANSP 133707,
218373; MCZ 124210); Shelter Island (ANSP
362811); East Patachogue (MCZ 288109);
Freeport (MCZ 248224); Coney Island (ANSP
362808); Far Rockaway (MCZ 54518,
139569); Sheepshead (MCZ 156658); Staten
Island (ANSP 132043; MCZ 78422, 201631;
USNM 59725, 307159, 492500b). NEW JER-
SEY: Raritan Bay (USNM 608377); Union
Beach, Raritan Bay (USNM 606609); E of Nan-
tuxant Point, Newport (ANSP 162143); Point
Pleasant (ANSP 65200); Harvey Cedars, Long
Beach Island (ANSP 106527); Beach Haven
(ANSP 155518); Shrewsbury River (ANSP
99006); Brigantine Island (ANSP 195015,
328333; USNM 611523); Atlantic City (ANSP
56843; MCZ 188902; USNM 492524); Vent-
nor (ANSP 99155); Margate City (ANSP
45192); Great Egg Harbor Bay (ANSP 326220;
А.М.); Avalon (ANSP 354825); Wildwood
Beach (ANSP 78982, 194600); Саре Мау
(ANSP 45098, 65201, 106229, 110131,
117532; USNM 119503, 124603, 406309,
504479); Cape May Harbor (ANSP 182641);
Carson’s Inlet (ANSP 133907); Bivalve (ANSP
113465; A.M.); Dividing Creek (A.M.); Fortes-
cue (ANSP 113466). DELAWARE: Woodland
Beach (USNM 522267); Rockhall (ANSP
112612); S of Fort Miles (USNM 621420); In-
dian River Inlet (MCZ 198052); М end of Cedar
Neck, S side of Indian River Bay (USNM
621418, 621419); Mouth of Cedar Creek
(USNM 406310); Assawoman Wildlife Area
(ANSP 302221). MARYLAND: Mayo Beach
(USNM 522288); Galesville (USNM 595602);
Chesapeake Beach (USNM 191595, 473893);
Parkers Creek (USNM 521920, 536709); near
Triton Beach (758314); Piney Island (USNM
667267); Patuxent River (ANSP 396602; MCZ

139263); Carrolls Bank, mouth of Patuxent
River (USNM 363927); Broome Island (USNM
473465); N of Solomons Island (USNM
600765); Solomons Island (USNM 424040);
Millstone Landing (USNM 512876); Benedict
(USNM 473456); St. Marys River (USNM
379539); Smiths Creek (USNM 518853);
Cobb Island (USNM 499519, 522921); Island
Creek (USNM 252064); Hope House, N of
Easton (USNM 602855); Huggins Point, S of
Leonard Town (MCZ 216712); Town Point
(ANSP 132462; MCZ 46420, 52344); Church
Creek, Cambridge (MCZ 200468); Parsons
Creek (MCZ 211150); Bishops Head (MCZ
139264); Ocean City, Sinepuxent Bay (USNM
601744, 601747); E of Dames Quarter (USNM
618922); SE of Chance Island (USNM
473331); Deal Island (USNM 468283); Cris-
field (USNM 595598); near Snow Hill, Chin-
coteague Bay (USNM 605563); West Bay,
Chincoteague Bay (USNM 591783). VIR-
GINIA: Chincoteague Island (MCZ 199443;
USNM 530826, 533596); Oyster (USNM
407459); Magotha (ANSP 275980; USNM
422297); Smiths Island (USNM 153352,
225965, 422294, 422295); Dameron Marsh,
Balls Creek and Ingram Bay (ANSP 305137);
Fisherman Island (USNM 422293); Brighton
(USNM 171118); Assateague Island (ANSP
352439; USNM 809536); Toms Cove, As-
sateague Island (USNM 209268); Watts Bay
(USNM 701629); Fairport (USNM 103258);
Potomac Beach (USNM 473900); Wicomico
Church (USNM 269147); Bretton Bay (USNM
628900); Greenvale Creek near Rappahan-
nock (MCZ 214466, 291314); Sweetheart Is-
land (USNM 269077); Fleeton (USNM
308939); Severn River (USNM 679373); Chis-
man Creek (USNM 603690); Poropotank,
York River (USNM 679375); Goodwin Island
(USNM 679374); Yorktown (USNM 474077);
Norfolk (MCZ 46542; USNM 407588, 504483,
539235, 667497); Granby and Lakewood
bridge, Norfolk (ANSP 219145; MCZ 221231;
USNM 653497); Boissevain Avenue, Norfolk
(ANSP 263386; MCZ 186692); Lamberts Point
(ANSP 263388), Elizabeth River (MCZ
186696); Lafayette Park, on bank of Lafayette
River (MCZ 186691). NORTH CAROLINA:
Bodie Island (МСУ 105478); Pee Island
(USNM 488826); Hatteras Island (USNM
637152); Cedar Island (A.M.); North River,
Woodville (A.M.); Williston (A.M.); Oyster
Creek (A.M.); Beaufort (ANSP 56839, 145683;
USNM 380566, 382982, 433203, 523610,
523620); Mullet Pond, Beaufort (USNM
380565); Brackish Pond (MCZ 141059); North

WESTERN ATLANTIC ELLOBIIDAE 273

River (ANSP 179830; MCZ 81930); Newport
River (A.M.); Wrightsville (USNM 492540);
Southport (ANSP 113647, 227028; MCZ
176143); Smith Island, Cape Fear (ANSP
95602); Cape Fear River (MCZ 175210).
SOUTH CAROLINA: Pawleys Island (MCZ
201642); Bull Island (USNM 543341); Sulli-
vans Island (USNM 492537); Charleston
(ANSP 56884; USNM 26571, 26574, 39823,
307156); Folly Beach (ANSP 300432); Edisto
Island (USNM 474096); Edisto Beach (ANSP
180786); Yemassee (USNM 492539; A.M.);
Lobeco (A.M.); Pocataligo (USNM 603106); N
of McClellanville (USNM 758249). GEORGIA
(ANSP 56835): Skidaway Island (A.M.); Cox-
boro Island (USNM 622491); Savannah (MCZ
200467; USNM 610311, 665761, 665762);
Jones Island, Savannah River (ANSP 164988);
Crescent (A.M.); Valona (A.M.); Рой King
George, Darien (USNM 707263); Sea Island
(ANSP 56847); Saint Simons Island (ANSP
123529); Brunswick (ANSP 183414; MCZ
186697; USNM 425961; A.M.); Jekyll Island
(A.M.). FLORIDA: Fernandina (MCZ 201208);
St. John's River (USNM 307155); Mayport
(ANSP 56844, 76095); Jacksonville (ANSP
56841; USNM 39815); St. Augustine (ANSP
45077, 66965, 140731; MCZ 186695; USNM
36014, 39825a, 492529); Fort Marion, St. Au-
gustine (ANSP 9523); Matanzas (USNM,
672437); Indian River (MCZ 186699, 291315;
USNM 758221); Ormond Beach (MCZ 82493);
New Smyrna Beach (MCZ 291026; A.M.); Oak
Hill (ANSP 22362; MCZ 201640; USNM
87624); Titusville (MCZ 143992); Banana
River (MCZ 201643); Grant (MCZ 105475);
Lake Worth (ANSP 69851; MCZ 291287;
USNM 253538); Fort Dallas (USNM 39817,
119502); Haulover Canal, head of Indian River
(ANSP 62720); Miami (ANSP 77506, 82844,
145885); Biscayne Bay (MCZ 291305); Sands
Key (MCZ 291309); Virginia Key, Biscayne
Bay (ANSP 189570); Coconut Grove (MCZ
291299, 291300; USNM 603111); Waveland
(USNM 103495, 123532); Madero Bay (MCZ
291289); Soldier Key (MCZ 174457); Middle
Key, Barnes Sound (USNM 405985); McGinty
Key (ANSP 189572; MCZ 103303); Pumpkin
Key (USNM 355115); Tavernier (MCZ
294246); Key Largo (ANSP 294318; MCZ
291281, 291302, 291313, 294228; USNM
603119); Tavernier Creek (USNM 667401;
А.М.); Tavernier Key (USNM 492549,
492552); Snake Creek (MCZ 294234); Plan-
tation Key (MCZ, 294230, 294231; A.M.);
Windley Key (USNM 603104); Indian Key
(USNM 26423, 27917, 462894a); Indian Key

Fill (A.M.); Lignumvitae Key (ANSP 156327,
189571); Lower Matecumbe Key (MCZ
291294; USNM 492553); Long Key (ANSP
76700; MCZ 291296, 291031; USNM 193363,
492578; A.M.); Grassy Key (MCZ 291288;
A.M.); Crawl Key (MCZ 174458, 291032,
291033, 294235; A.M.); Marathon, Key Vaca
(ANSP 189584); Knight Key (A.M.); Bahia
Honda Key (ANSP 104095; USNM 269777a,
270310); West Summerland Key (A.M.); New-
found Harbor (USNM 272688a); Little Pine
Key (USNM 681642); Big Pine Key (ANSP
104092, 104093; MCZ 291029, 291034,
294237, 294243; USNM 597453; A.M.); Big
Torch Key (ANSP 104096; A.M.); Howe Key
(USNM 681641); Ramrod Key (MCZ 291028,
291045, 294232, 294233); Sugarloaf Key
(ANSP 104094); Big Coppit Key (USNM
603102); Boca Chica Key (ANSP 104091,
152501; USNM 270326, 511180); Stock Is-
land (ANSP 149993; MCZ 59993; USNM
270269); Key West (ANSP 294309; MCZ
9947; USNM 27730, 529557, 596787,
667405, 670450); Shark Key (USNM 696979,
711532); Boca Grande Key (USNM 270262);
Flamingo Key (ANSP 294312; MCZ 291038,
294236, 294244); Cape Sable (MCZ 291035);
Lossman River (ANSP 132369); Everglades
City (USNM 683928); Marco Island (MCZ
294249); Little Marco (USNM 511210); Naples
(ANSP 189568; MCZ 291316); Sanibel (ANSP
170640; MCZ 13278, 13279, 294239,
294248); Pineaire, Pine Island (MCZ 291030);
Bokeelia (MCZ 294245; A.M.); Matlacha
Key (MCZ 291311); Starvation Key (ANSP
294320); Captiva Island (MCZ 294250); Punta
Gorda (USNM 492577); between Fort Myers
and Punta Gorda (USNM 531133); Charlotte
Harbor Bay (USNM 407410); Boca Grande,
Gasparilla Island (ANSP 142272); Little Gas-
parilla Island (ANSP 189592); Sarasota
(USNM 159254, 487213, 487314); 5 km N of
Sarasota (ANSP 195406); Long Boat Key
(MCZ 294252); between Palma Sola and Cor-
tez (MCZ 291037); Anna Maria Key (MCZ
291027); Palmetto (A.M.); Manatee River
(MCZ 3969; USNM 61052); Small Island,
mouth of Manatee River (MCZ 291282); Big
Bend Road [Rt. 672] (A.M.); Tampa (ANSP
56836, 294317, 362807; MCZ 201649,
291284, 294238; USNM 36061, 37609,
193361, 196349, 492574, 504487); Young
Lagoon, Tampa (USNM 37610); Hillsborough
River (USNM 100693); Palm River (MCZ
201647); Mullet Key (A.M.); Shell Key (USNM
466206a, 466288); S of Pass-a-grille (MCZ
201685, 291285, 294241); Pass-a-grille

274 MARTINS

(ANSP 294321); Pinellas Point (MCZ 294190;
USNM 194730); St. Petersburg (ANSP
132371, 132372, 132373, 132374; USNM
341721, 466194); Bird Key (ANSP 43568);
Gulfport (MCZ 294242); Boca Ciega Bay
(ANSP 189574; USNM 341722); Clearwater
(ANSP 189569, 294319, 294322; MCZ
294247; USNM 611786a); Anclote River
(А.М.); Hudson (A.M.); Aripeka (ANSP 73902,
151138, 151257); Little Blind Creek (ANSP
149525); Cedar Key (ANSP 56842, 194040,
194227, 362810; MCZ 199316, 201653,
201656, 291279; USNM 36012, 36013,
61700); Suwannee (ANSP 88138; MCZ
199315); Jena (USNM 484844); Adams Beach
(MCZ 186701); St. Marks (ANSP 56814,
56816); Рапасеа (MCZ 91696; USNM
706611); St. Andrews Bay (ANSP 83641;
USNM 667402, 667403, 667404); Anderson's
Bayou, St. Andrews Bay (ANSP 83653);
Ochlockonee (MCZ 199317); Port St. Joe Bay
(MCZ 29130, 291297); Panama City (MCZ
235949). ALABAMA: Cedar Point (MCZ
186702); Heron Bay (ANSP 315710); Dauphin
Island (USNM 701860); Coden Beach (USNM
422364); St. Anthony Bay (ANSP 365474).
MISSISSIPPI: Boat Harbor at Ocean Springs
(MCZ uncatalogued). LOUISIANA: Chan-
deleur Islands (USNM 189168); Grand Isle
(MCZ 186700; USNM 603100); Cherniere-au-
Tigre (ANSP 145106); Fort Pike, Orleans Par-
ish (MCZ 251101); Houma (USNM 653314);
Grand Lake, Calcasieu (USNM 160814,
467015); Myrtle Grove (USNM 628753); Cam-
eron (USNM 177952). TEXAS: Galveston
(ANSP 69414, 71522, 73692); East Lagoon,
Galveston (MCZ 217872); Seabrook (ANSP
105895); Clearlake (MCZ 217874); Quitano
Beach (MCZ 227841; USNM 600177); Free-
port (ANSP 254824); Matagorda (USNM
125534); La Vaca, La Vaca Bay (MCZ 223041;
USNM 126755); Sand Point, Keller Bay
(USNM 464802, 465269, 465278); Indianola
(USNM 26425, 26572, 39832); Rockport
(MCZ 217875); Carancahua Bay (USNM
134435); Aransas Pass (ANSP 322151; USNM
603108); Ransom Island (MCZ 200470); Ran-
som Island, Redfish Bay (MCZ 198174); Port
Aransas (ANSP 284785); Mustang Island
(MCZ 217873); Corpus Christi (МСА 294191;
USNM 603107); Port Isabel (ANSP 80044;
MCZ 193910; USNM 603109, 603110);
Brownsville (ANSP 351204); Callo del Oro
(USNM 473355). MEXICO: Tampico (USNM
219997a); N end of Ascension Bay, Quintana
Roo (USNM 735992, 735993, 736143); near
Allen Point, Ascension Bay, Quintana Roo

(USNM 736693). BELIZE: Belize (ANSP
96583); Botanical Garden (USNM 426007a).
BERMUDA (ANSP 56815, 56817, 56819,
56823, 77905; MCZ 3972, 9949, 201106,
201663, 201667, 201674, 201676, 291290,
291303, 291307, 291312; USNM 6540,
11420, 37605, 39805, 94432, 94433, 98152,
173643, 173644, 173645, 173646, 173647,
492582, 492584): Fairyland (ANSP 99053);
Hamilton Island (MCZ 201105); Hungry Bay
(ANSP 85712; MCZ 10381, 24248, 48123,
291306; USNM 492578, 492583; A.M.); South
Shore (ANSP 86925, 86926, 86927; MCZ
53060, 53061, 53062; USNM 109557,
109558, 109559); Ely's Harbour (A.M.). BA-
HAMA ISLANDS (MCZ 54619): GRAND BA-
HAMA ISLAND: Bootle Bay (ANSP 371880);
North Hawksbill Creek (ANSP 370567);
GREAT ABACO ISLAND: Witch Point (ANSP
359151; USNM 492565, 492580b); Mastic
Point Creek (USNM 618607); Angelfish Point
(MCZ 116711); BIMINI: Mangrove Creek
(ANSP 325614); Mosquito Point (ANSP
326148); BERRY ISLANDS (MCZ 294251);
ANDROS ISLAND: Morgan’s Bluff (A.M.);
South Mastic Point (A.M.); Mangrove Cay
(ANSP 189587; USNM 269477, 269855,
269968a, 270214d, 270216); Linder Key
(USNM 269360, 270483); NEW PROVIDENCE
ISLAND: Nassau (USNM 160766); Dick's
Point (MCZ 291280); South Beach, Fox Hill
(MCZ 294240); Bonefish Pond (USNM
618645; A.M.); Millars Sound (A.M.); Millars
Road (A.M.); ROYAL ISLAND (USNM
468115); ELEUTHERA ISLAND: Rock Sound
(MCZ 135932); CAT ISLAND: Russell Creek
(ANSP 294330; MCZ 291278, 294229);
Arthurstown (MCZ 291295); EXUMA CAYS:
Hog Cay (ANSP 285755); LONG ISLAND:
Brett's Hill (MCZ 113332, 291283, 291286);
Clarencetown (ANSP 173261); Glenton’s
(ANSP 189573; MCZ 291291); Pinder's (MCZ
113330); Salt Pond, Clarencetown (USNM
595950); AKLINS ISLAND: Pinnacle Point
(USNM 390856, 392237); Rooker Cay (USNM
390674); CAY SAL BANK: CAY SAL (MCZ
291331). TURK’S & CAICOS: TURK’S IS-
LAND: (USNM 492474a) CUBA (ANSP
56794, 56795, 56820, 189584): Bahia de
Santa Rosa (USNM 492556a); Dimas (USNM
492559); Cayo de las Cinco Léguas (ANSP
294325); Cayo Perro, Cárdenas (ANSP
189589); Marianao (ANSP 294324); Cayo
Maya, near Cayo Santa Maria (MCZ 294187);
Cayo Juan Garcia (MCZ 291308); Isla de Co-
bos (MCZ 294186); Cayo Romano (MCZ
291309); Gibara (USNM 603097); Finca, Sa-

WESTERN ATLANTIC ELLOBIIDAE 218

banalmar (MCZ 294189); Punta de los Colo-
rados, Bahia de Cienfuegos (MCZ 291292);
Alto del Caracol, Caracoles, S of Pinar del Rio
(MCZ 201675); Isla de Pinos (MCZ 48079;
USNM 130028). JAMAICA (ANSP 56793):
Green Island Harbor (USNM 440805); Pali-
sadoes (USNM 713079). HAITI: Gonave Is-
land (USNM 492531a). DOMINICAN REPUB-
LIC: Isla La Matica, Playa Boca Chica, E of
Santo Domingo (R.B.). VIRGIN ISLANDS: ?
TORTOLA (USNM 6485).

Subgenus Detracia Gray, 1840

Detracia Gray, 1840: 20. Type species by
monotypy: Detracia bullaeoides (Mon-
tagu, 1808) [unjustified emendation of
bullaoides].

Tifata H. & A. Adams, 1855b: 245. Type spe-
cies herein designated: Tralia (Tifata)
globulus (Orbigny, 1837).

Ensiphorus Conrad, 1862: 571. Type species
by monotypy: Melampus (Ensiphorus)
longidens Conrad, 1862 [Miocene].

Eusiphorus Conrad. Zilch, 1959: 65 [error for
Ensiphorus; in synonymy].

Description: Shell globose to oval-oblong to
fusiform; spire low to high, whorls ten to 13;
aperture very narrow; columellar tooth ob-
lique and stronger than parietal teeth; parti-
tion of inner whorls occupying almost all of
body whorl, parietal teeth continuing on wall
of inner whorl as conspicuous lamellae. Ani-
mal grayish blue to dirty white, sometimes
mottled with dark brown spots. Conspicuous
medial nodes on base of central radular
tooth. Visceral mass separated from foot by a
length equivalent to about one and one-half
whorls; mantle organ forming conspicuous
pouch; pallial gonoducts and cephalopedal
reproductive organs elongate.

Remarks: Detracia was created by Gray
(1840) for the West Indian species Melampus
bullaoides (Montagu) because it was said to
have only the columellar tooth on the inner
lip. Careful inspection of Melampus (D.) bul-
laoides reveals a hidden parietal tooth mid-
way on the inner lip and a long, vertical, cal-
lous anterior parietal tooth. The most obvious
diagnostic feature is the very strong, oblique
columellar tooth.

The genus Tifata was introduced by H. & A.
Adams (1855b) as a subgenus of Tralia and
was characterized by having two spiral, ele-
vated, lamellar plates at the forepart of the
inner lip. The genus Tralia had been intro-

duced by Gray (1840) on account of the
shape of the ощег lip, but H. & A. Adams
(1855b), followed by Binney (1865), used the
incorrect anatomical feature, “foot posteri-
orly acute,” to separate this taxon from
Melampus (Fischer & Crosse, 1880; Dall,
1885). This led to the erroneous inclusion of
Tifata within the genus Tralia. The pattern of
dentition of the inner lip used by the Adams
brothers to characterize Tifata is not con-
spicuous in the first two originally included
species, Tralia cingulata (Pfeiffer, 1840) [=
Melampus (D.) bullaoides (Montagu, 1808)]
and Tralia floridana (Pfeiffer, 1856) [= Melam-
pus (D.) floridanus]. In those species the first
parietal tooth is usually a broad, longitudinal
callosity, and the lamellar second parietal
tooth is too far posterior in the aperture to be
considered in the “forepart” of the inner lip.
In the next species listed, Tralia globulus (Or-
bigny, 1837) [= Melampus (D.) globulus], the
longitudinal callosity is less pronounced, ге-
sembling more an oblique lamella (Keen,
1971: fig. 2402). This agreement with the
original description justifies my choice of Tra-
lia globulus as type species of the subgenus
Tifata; to clarify further the taxonomy, |
hereby select a lectotype for this species
(Fig. 320). Nevertheless, all species originally
included in Tifata H. & A. Adams, including
Tralia pulchella (Petit, 1843), should be
placed in Detracia on the basis of the strong,
oblique columellar fold.

Morrison (19516, 1964) considered
Melampus (D.) monile (Bruguiere, 1789) to
belong to Pira H. & A. Adams, 1855, which he
elevated to generic rank. Pira was character-
ized as having three teeth on the inner lip.
The type species, Auricula kuesteri (Krauss
MS) Kúster, 1844 [= Melampus fasciatus (De-
shayes, 1830)], was illustrated by Küster
(1844: 34-35, pl. 4, figs. 10-13), who noted
the conspicuous twisted columellar tooth. Its
relative prominence 1$ evident in his figure 13,
which represents Kúster's variety A of that
species. Although a decision on the system-
atic status of Pira is at present unwarranted
owing to lack of anatomical data for compar-
isons, Melampus топйе (Bruguière) should
be included in Detracia on the basis of shell
and anatomical characters.

Conrad (1862) introduced Ensiphorus for a
Miocene species from Yorktown, Virginia,
that was characterized by its elongate,
slightly curved columellar tooth directed ob-
liquely upwards, a characteristic of Melam-
pus (D.) morrisoni. The type species, Ensi-

276 MARTINS

phorus longidens, illustrated by Conrad
(1866, pl. 4, fig. 12), resembles in its size and
shape a small Melampus (D.) bullaoides or a
Melampus (D.) floridanus, hence justifying
putting Conrad’s name in synonymy with De-
tracia.

Anatomical comparison of Detracia with
Melampus does not justify their generic sep-
aration. Thus Detracia is treated as a subge-
nus of Melampus [see the remarks under
Melampus $. 1.].

The subgenus Detracia is represented in
the West Indian region by five species, which
can be distinguished on the basis of shell
morphology. The type species, Melampus
(D.) bullaoides, is usually high spired, oval-
elongate and without any indication of a ca-
rina on the shoulder of the body whorl. The
spire has axial ribs and incised spiral
grooves, and in juveniles it is hirsute. The
spire of juvenile Melampus (D.) monile 1$ also
hirsute (Figs. 341, 345), but the shell is ovate-
conic and the midale riblets of the outer lip
are the strongest, whereas in Melampus (D.)
bullaoides the anteriormost riblet of the outer
lip is strongest. Some high-spired specimens
of North American Melampus (D.) floridanus
resemble Melampus (D.) bullaoides, but can
be distinguished from them by the absence
of axial ribs and hairs on the spire, as well as
by the conspicuous parietal callus that occu-
pies the site of the anterior parietal tooth.
This latter character also distinguishes
Melampus (D.) floridanus from its South
American counterpart, Melampus (D.) para-
nus, which lacks the parietal callus. In addi-
tion, the South American species never has
more than three riblets in the outer lip,
whereas the North American species has four
to eight riblets. Also included in this subge-
nus is Melampus (D.) morrisoni, a species
that has certain characteristics of Melampus
5. s., such as the readily visible upper parietal
tooth and a discrete anterior parietal tooth.
Placement in Detracia is based upon the
strong columellar tooth, narrow aperture,
long pallial gonoducts and the pouch-like
mantle organ.

Habitat: The West Indian species of Detracia
characteristically live farther from the water
than do the species of Melampus s.s. with
which they can occur. Melampus (D.) monile
is an exception in preferring the area of the
high-tide mark, in which it is very common
among piles of rocks frequently covered by
waves at high tide. Melampus (D.) morrisoni

prefers inland lagoons and, along with
Melampus (D.) bullaoides, lives in areas
reached only by spring tides. The North
American Melampus (D.) floridanus and the
South American Melampus (D.) paranus are
very closely related. Morrison (1951a) inter-
preted this similarity as a case of parallel ev-
olution. The North American species lives in
inland salt marshes of low salinity and curi-
ously it is absent from mangroves, whereas
its South American counterpart is a su-
pratidal estuarine species, living in man-
groves (Marcus & Marcus, 1965a).

Range: The subgenus Detracia is mostly
tropical and occurs in the Indo-Pacific and
the West Indies. In the Western Atlantic it
ranges from New Jersey to southern Brazil.

Melampus (Detracia) bullaoides
(Montagu, 1808)
Figs. 290-314

Voluta bullaoides Montagu, 1808: 102, pl. 30,
fig. 4 [Lincoln, England (error), herein
corrected to St. Vincent's, West Indies;
location of type unknown].

Auricula multivolvis Jeffreys, 1833: 518 [Scar-
borough, England (error), herein cor-
rected to St. Vincent’s, West Indies; ho-
lotype USNM 55308 (Fig. 290)].

Tornatella bullaoides (Montagu). Ferussac,
1821: 108.

Melampus bulla Lowe, 1832: 280 [unneces-
sary new name for Voluta bullaoides
Montagu].

Melampus (Melampus) bulla Lowe. Beck,
1837: 108.

Detracia bullaeoides (Montagu). Gray, 1840:
20 [unjustified emendation for bulla-
oides].

Auricula cingulata Pfeiffer, 1840: 251 [Cuba;
location of type unknown]; Küster, 1844:
40, pl. 6, figs. 4-6; Reeve, 1877, pl. 6, fig.
46.

Auricula oliva Orbigny, 1841: 189, pl. 12, figs.
8-10 [Outskirts of Havana, Cuba; lecto-
type herein selected BMNH 1854.10.4.
109 (Fig. 291)].

Melampus cingulatus (Pfeiffer). C. B. Adams,
1849: 42; С. В. Adams, 1851: 186; Shut-
tleworth, 1854b: 102; Pfeiffer, 1854b:
147; Pfeiffer, 1856a: 17; Binney, 1859:
161, pl. 75, figs. 12, 13; Binney, 1860: 4;
Poey, 1866: 394; Pfeiffer, 1876: 301;
Nevill, 1879: 219; Arango y Molina, 1880:
58; Crosse, 1890; 258.

WESTERN ATLANTIC ELLOBIIDAE PTT

Conovulus bullaoides (Montagu). Forbes &

Hanley, 1852: 197.

Melampus роеу! Pfeiffer, 1853b: 126 [Cuba;
location of type unknown); Pfeiffer,
1854b: 147; Pfeiffer, 1856a: 17; Pfeiffer,
1876: 301.

Melampus (Tralia) cingulatus (Pfeiffer). H. & A.
Adams, 1854: 11.

Melampus (Tifata) cingulata (Pfeiffer). H. 8 A.
Adams, 1855b: 245.

Melampus bullaoides (Montagu). Pfeiffer,
1856a: 18; Kobelt, 1901: 277, pl. 33, figs.
6-8.

Melampus oblongus Pfeiffer, 1856b: 393
[Bermuda; lectotype herein selected
BMNH 1968848 (Fig. 292)].

Tralia cingulata (Pfeiffer). Binney, 1865: 18,
fig. 19; Tryon, 1866: 9, pl. 18, fig. 10;
Fischer & Crosse, 1880: 22; Dall, 1883:
323.

Melampus ? bullaoides (Montagu). Pfeiffer,
1876: 301.

Melampus (Detracia) bulloides (Montagu).
Dall, 1885: 285, pl. 18, fig. 7; Dall, 1889:
92, pl. 47, fig. 7; Simpson, 1889: 68;
Maury, 1922: 56 [misspelling of bulla-
oides].

Melampus bulimoides (Montagu).
1901: 35 [error for bullaoides].

Melampus bulloides (Montagu). Davis, 1904:
126, pl. 4, fig. 4 [error for bullaoides].

Melampus (Detracia) bullaoides (Montagu).
Peile, 1926: 88.

Melampus (Detracia) bullaeoides (Montagu).
Thiele, 1931: 467; Zilch, 1959: 65, fig.
207 [error for bullaoides].

Detracia bulloides (Montagu). M. Smith,
1937: 147, pl. 55, fig: 1; pl. 67, fig: 7 pl.
67 copied from Dall (1885, pl. 18)]; Perry,
1940: 178: Coomans, 1958: 103; Porter,
1974: 300 [error for bullaoides].

Detracia bullaoides (Montagu). Morrison,
1951a: 18, figs. 1, 5 [systematics]; Mor-
rison, 1951b: 9; Perry 8 Schwengel,
1955: 198, pl. 53, fig. 359; Morrison,
1958: 118-124 [ecology]; Warmke 8 Ab-
bott, 1961: 153, pl. 28, fig. o; Coomans,
1969: 82; Morris, 1973: 274, pl. 74, fig.
13; Abbott, 1974: 332, fig. 4092; Hum-
phrey, 1975: 196, pl. 22, fig. 26; Emerson
8 Jacobson, 1976: 190, pl. 26, fig. 22;
Rehder, 1981: 648, fig. 349; Gibson-
Smith & Gibson-Smith, 1982: 117; Vokes
8 Vokes, 1983: 60, pl. 22, fig. 15; Jensen
8 Clark, 1986: 457, figured.

Detracia roquesana Gibson-Smith & Gibson-
Smith, 1982: 117, fig. 6 [Isla de los

Verrill,

Roques, Venezuela;
784718 (Fig. 300)].

holotype USNM

Description: Shell (Figs. 290-305) to 15 mm
long, globose to fusiform, solid, shiny, uni-
form whitish, yellow to brown or with as many
as three, rarely more, white, unequally wide
bands on body whorl; body whorl frequently
with axial zigzag markings or with combina-
tion of bands and markings. Deep umbilical
groove sometimes present in gerontic spec-
imens. Spire high, mucronate, whorls as
many as 13, flat and sculptured with well-
marked spiral grooves, axial ribs and a spiral
row of laterally compressed, short periostra-
cal hairs. Body whorl 70% of total length,
oval, smooth or with very faint spiral lines,
without carina on shoulder. Aperture length
about 80% of body whorl, very narrow, an-
gulated anteriorly, with base sometimes
broad and round in gerontic specimens; inner
Ир with a strong, oblique, twisted columellar
tooth and a small, oblique, hidden parietal
tooth just posterior to conspicuous longitudi-
nal parietal callus; area posterior to parietal
tooth excavated, with anterior border weakly
raised to plait-shaped; in the latter case, cor-
responding riblet of outer lip becomes stron-
ger, forming wall of narrow anal canal; outer
lip sharp, rarely smooth within, with one
strong riblet opposite columellar tooth, usu-
ally followed posteriorly by as many as eight
riblets that do not reach the edge of the lip,
gradually becoming smaller towards poste-
rior end of aperture. Inner partition of whorls
occupying entire body whorl; connection of
posterior visceral mass space with aperture
very narrow (Fig. 302). Protoconch trans-
lucent, whitish to slightly brown (Figs. 303-
305).

Animal bluish gray; foot dirty white; top of
neck dark brown to black; tentacles subcy-
lindric, pointed, translucent in first quarter,
abruptly changing to dark gray or black;
mantle skirt light gray. Pallial cavity long;
mantle organ dark brown, well developed,
forming very conspicuous pouch; kidney
long.

Radula (Figs. 306-310) having formula [24
+ (1 + 14) + 1 + (14 + 1) + 24] x 100. Base
of central tooth twice width of base of later-
al teeth, triangular, with very conspicuous
prominences on inner surface of arms; crown
smaller than that of lateral teeth, with medial
depression at posterior edge; mesocone
small, sharp; ectocones not defined. Lateral
teeth ten to 18; crown strong, broadly trian-

MARTINS

278

FIGS. 290-305.

WESTERN ATLANTIC ELLOBIIDAE 279

gular, half total length of tooth; mesocone
sharp, pointing laterally; no distinct en-
docone or ectocone. Transitional tooth with
base partly reduced, crown weakly project-
ing posteriorly, with small ectocone or serrate
edge on ectocone site. Marginal teeth 20 to
31 with reduced base and elongated crown;
mesocone strong and sharp, gradually be-
coming rounded toward the edge of the rad-
ular ribbon; appearance of ectocones incon-
sistent from row to row.

Stomach (Fig. 311) as in subfamily.

Reproductive system (Fig. 312) with ovo-
testis conical, dark brown; mucous gland
spiral, conical; ргеуадта! caecum very con-
spicuous; bursa duct thick, enters vagina op-
posite to exit of posterior vas deferens; bursa
large, approximately oval; vagina thin, about
one and four-fifths times length of body whorl;
penis thin, long; penial retractor inserting with
columellar muscle.

Nervous system (Fig. 313) having cerebral
commissure about as long as width of cere-
bral ganglion; left parietovisceral connective
twice the length of right one.

Remarks: Melampus (D.) bullaoides originally
was stated to belong to the British fauna al-
though Montagu (1808) doubted its origin. He
reported it as from Lincoln, because this was
the locality given on the lot label in the Port-
land collection, from which the shell had
been purchased. Lowe (1832) repeated the
original information when, for no apparent
reason, he renamed Montagu's species
Melampus bulla. Beck (1837) erroneously re-
ported Lowe's species from “Atlantic Ocean,
Boreal Africa.” The shell, however, became
well known in European collections, allowing
Gray (1840: 21) to remark, “it is one of the
most common shells in the small boxes from
the West Indies.” Jeffreys (1869: 109) also
noted that ‘‘Voluta bullaoides of Montagu (my
Auricula multivolvis) [Fig. 290] is a rather
common West Indian species.” He also

stated that a specimen had been found at
Scarborough and that the species had been
reported from Croisic in the Loire-Inferieure.
Morrison (1951a) explained those odd occur-
rences as inclusions in ballast picked up by
ships in the West Indies and dumped along
the coast of England.

The high degree of variability that charac-
terizes the genus Melampus is also notice-
able in Melampus (D.) bullaoides and caused
the introduction of most of the names here
considered synonyms. In Pfeiffer's three ma-
jor revisions of the family (1854b, 1856a,
1876), the name Melampus bullaoides ap-
pears only in the last two, indeed with a query
in the last one. Pfeiffer (1856a) noted that
Forbes & Натеу (1852) had treated his
Melampus cingulatus from Cuba as a junior
synonym of Montagu's species. Although he
admitted that they were very closely related,
Pfeiffer was reluctant to synonymize them
solely on the basis of Montagu’s figure, leav-
ing the problem, in his words, to the consid-
eration of the experts. Almost at the same
time Pfeiffer described Melampus cingulatus,
Orbigny (1841) described and nicely illus-
trated Auricula oliva from Cuba (Fig. 291).
Presumably Orbigny was not aware of Pfeif-
fer's publication, because the two descrip-
tions are very similar. The second of Pfeiffer's
supposed species, Melampus poeyi, also
from Cuba, is intermediate between Melam-
pus bullaoides and Pfeiffer's Auricula cingu-
lata, according to Pfeiffer’s observation
(1856a). The last of Pfeiffer’s names, Melam-
pus oblongus, was applied to a Bermudian
specimen (Fig. 292) that Pfeiffer (1856b)
thought was more closely allied to Melampus
angiostomus (Deshayes, 1831) than to his
Auricula cingulata. The Bermudian speci-
mens have a quasi-smooth outer lip, and
some populations have only the characteris-
tic strong riblet opposite the columellar tooth
(Figs. 293-296). Comparison of individuals
from Bermuda with specimens from Florida

FIGS. 290-305. Melampus (D.) bullaoides (Montagu). (290) Auricula multivolvis Jeffreys, holotype (USNM
55308), Scarborough, Englana, sl 7.5 mm. (291) Auricula oliva Orbigny, lectotype (BMNH 1854.10.4.109),
Cuba, sl 11.0 mm. (292) M. oblongus Pfeiffer, lectotype (BMNH 1968848), Bermuda, sl 11.1 mm. (293)
Somerset Bridge, Bermuda, sl 8.1 mm. (294) Hungry Bay, Bermuda, sl 10.6 mm. (295) Hungry Bay,
Bermuda, sl 10.3 mm. (296) Hungry Bay, Bermuda, sl 11.2 mm. (297) Crawl Key, Florida (R.B.), sl 9.0 mm.
(298) Big Pine Key, Florida, sl 9.4 mm. (299) South Mastic Pt., Andros Island, Bahamas, sl 9.1 mm. (300)
Detracia roquesana Gibson-Smith & Gibson-Smith, holotype (USNM 784718), Isla de los Roques, Vene-
zuela, sl 10.6 mm. (301) Long Key, Florida, sl 2.4 mm. (302) Grassy Key, Florida, sl 13.7 mm. (303) Lateral
view of spire and protoconch, Somerset Bridge, Bermuda. (304) Lateral view of spire and protoconch, Long
Key, Florida. (305) Top view of spire and protoconch, Somerset Bridge, Bermuda. Scale 1 mm.

280 MARTINS

FIGS. 306-309. Melampus (D.) bullaoides, radular teeth, Somerset Bridge, Bermuda, sl 9.0 mm. (306)
Central and lateral teeth. (307) Transitional and lateral teeth. (308) Marginal teeth. (309) Lateral view of

marginal teeth in preceding figure. Scale 50 um.

WESTERN ATLANTIC ELLOBIIDAE 281

1L 2L

ii

11L 12L T1M2M

Pace th

В 8M9M 15M 16M21M22M

A D ee _

FIG. 310. Меатриз (D.) bullaoides, radula. A,
Somerset Bridge, Bermuda; B, Big Pine Key, Flor-
ida. Scale 10 um.

15L T 1M 7M8M 12M 13M 17M 18M

FIG. 311. Melampus (D.) bullaoides, stomach, Flor-
ida. Scale 1 mm.

and Bahamas, however, revealed overlap in
characters of the apertural dentition (Figs.
297-299, 301, 302). One must conclude that,
owing to its variable expression, the apertural

FIG. 312. Melampus (D.) bullaoides, reproductive
system, Grassy Key, Florida. Scale 1 mm.

dentition in this species 15 not a reliable tax-
onomic character and that Melampus oblon-
gus Pfeiffer from Bermuda is conspecific with
Melampus (D.) bullaoides.

Very recently Gibson-Smith 4 Gibson-
Smith (1982), on the basis of six beach spec-
imens, described Detracia roquesana from
Islas de los Roques, off Venezuela (Fig. 300).
The Bermudian specimens in my collection,
which | refer to Pfeiffer's Melampus oblon-
gus, as well as those from the Bahamas, fit
the description of Gibson-Smith 4 Gibson-
Smith. The brown protoconch, so common in
Bermudian shells, also occurs in certain
specimens from Florida, although in the latter
specimens the aperture is narrower and more
dentate. Analysis of protoconch, radula and
anatomy of the Bermudian forms did not
yield any differences between specimens
from Florida and those from the Bahamas.
These facts led me to consider Detracia ro-
quesana Gibson-Smith & Gibson-Smith a
junior synonym of Melampus (D.) bullaoides
(Montagu).

Melampus (D.) bullaoides is very easily dis-
tinguished from Melampus (D.) floridanus by

282 MARTINS

"plpre

prvc prg

FIG. 313. Melampus (D.) bullaoides, central nervous system, Grassy Key, Florida. Scale 1 mm.

the presence of ribs on its spire and by its
protruding, mucronate apex. The spire is
constricted toward the apex and broadens
suddenly toward the base. The juveniles have
a crown of periostracal hairs. The last whorl
shows a wide range of color patterns, and
frequently young specimens are brightly col-
ored. In Melampus (D.) floridanus the spire 1$
regularly conical, glabrous, and the body
whorl has as many as three chestnut-brown
bands.

Habitat: Melampus (D.) bullaoides is а com-
mon inhabitant of the mangroves and can be
very abundant in some localities. The species
prefers the supralittoral zone and it frequently
lives on the edges of inland tidal lagoons,
sometimes in relatively dry places, in which
they aggregate under rocks, pieces of wood,
cardboard and other decaying trash.

Range: Bermuda; Florida, West Indies to
Suriname (Fig. 314).

Specimens Examined: FLORIDA (USNM
27914, 39833, 39838, 39839, 1524268,
492459): Fernandina (USNM 492544); Lake
Worth at Boynton (ANSP 194770); Miami
(ANSP 91284; USNM 153399, 492460); Co-
conut Grove (MCZ 82497, 291238, 291258);
Virginia Key (MCZ 46880); Bear’s Cut, Key
Biscayne (MCZ 153116); Soldier Key (MCZ
174459); Third Ragged Key, above Sands
Key (USNM 462736); Sands Key, Biscayne
Bay (MCZ 291269); Elliot Key (ANSP
160894); Key Largo (ANSP 56813; MCZ
56473, 291255, 291264; USNM 68130,
492546, 597456); Tavernier (MCZ 153121); N
of Tavernier Key (A.M.); Tavernier Key
(USNM 492550); Snake Creek (MCZ 291058);
Plantation Key (MCZ 199343, 291057); S of

WESTERN ATLANTIC ELLOBIIDAE 283

FIG. 314. Melampus (D.) bullaoides, geographic
distribution.

Ocean Drive, Plantation Key (A.M.); Indian
Key (USNM 462895, 492547); Indian Key Fill,
N of Indian Key Channel (A.M.); Little Duck
Key (MCZ 291061); Lower Matecumbe Key
(USNM 700771); N end of Long Key (A.M.);
Bonefish Key (MCZ 291059); Upper Grassy
Key (MCZ 291051); Grassy Key (MCZ
291064; A.M.); Crawl Key (MCZ 174470,
199342, 291046, 291047; A.M.); Key Vaca
(ANSP 181137); Marathon (MCZ 153258);
Knight Key (A.M.); Bahia Honda Key (ANSP
88030, 88132, 104098, 189562; USNM
269780); Howe Key (USNM 681640, 706760);
Big Pine Key (ANSP 89549, 104099, 189559;
MCZ 250733, 291048, 291049, 291517;
USNM 597454); Long Beach Drive, Big Pine
Key (A.M.); W end of Kohen Avenue, Big Pine
Key (A.M.); Big Torch Key (ANSP 104100;
A.M.); Little Torch Key (MCZ 291053); Ram-
rod Key (MCZ 291050, 291247); Sugarloaf
Key (ANSP 9635, 22475, 89550, 189560);
Lower Sugarloaf Key (USNM 672440); Sum-
merland Key (USNM 270318); West Summer-
land Key (MCZ 291084; A.M.); Military Key
(MCZ 291055); Pavilion Key (ANSP 93434);
Boca Chica Key (ANSP 104097, 152502,
189561; MCZ 162638; USNM 270327); Cow
Key (USNM 596786); Stock Island (ANSP
149994; USNM 270280); Key West (ANSP

56812, 100848, 174635, 264540, 294310;
USNM 36015, 36965, 61101, 153076,
270363, 338357); Snake Key (ANSP 105461);
Seminole Point (ANSP 105437); Boca
Grande Key (USNM 272834); Flamingo Key
(MCZ 291060); Cape Sable (MCZ 291062);
Marco Key (USNM 381333); mouth of Hend-
erson Creek, 5 km N of Marco (МСА 294215);
Little Marco (ANSP 93435); Bonita Springs
(MCZ 291063); Carl E. Johnson Park, Little
Carlos Pass (A.M.); Mound Key (MCZ
291270); Punta Rassa (MCZ 13705, 291056);
Sanibel Island (ANSP 179352; MCZ 291052);
Tarpon Bay, Sanibel Island (MCZ 13704,
55961); Turner's Pond, Sanibel Island (MCZ
232526); E of St. James, Pine Island (ANSP
93433); Captiva Island (ANSP 149406; MCZ
60186, 236852, 291054); Osprey (ANSP
88078); Mullet Key (USNM 652407; A.M.);
Pinellas Point (MCZ 294209); Bayou off Gulf-
port (MCZ 138942); St. Petersburg (MCZ
291242; USNM 343843, 366191, 466193);
Maximo Point, St. Petersburg (ANSP
167540); Shell Bay, off St. Petersburg (USNM
466207, 466289); Sand Key (ANSP 129249;
USNM 338365); Harts Bayou, Boca Ciega
Bay (MCZ 291266); Indian Rocks (ANSP
167541); Clearwater Island (ANSP 189558);
Clearwater (USNM 611785); Cedar Key (MCZ
291246; USNM 36895, 37611, 37612). BER-
MUDA: (ANSP 85590; MCZ 24246, 291240,
291253, 291260, 291262, 292267; USNM
6529, 94435, 98153, 173651, 492543);
Hamilton (ANSP 182551; USNM 152145,
171960); Fairyland (ANSP 99058, 111095); N
end of Long Bird Bridge (A.M.); Hungry Bay
(ANSP 88581; A.M.); S end of Ely's Harbour
(A.M.); W side of Somerset Bridge (A.M.);
Mangrove Bay (A.M.); Ireland Island (USNM
712379); Pond W of Evans Bay (A.M.); Rid-
dell's Bay (USNM 621666). ВАНАМА IS-
LANDS (MCZ 24141): GRAND ВАНАМА IS-
LAND: Dead Mans Reef [Sandy Bevan's Cay]
(ANSP 371222); Riding Point (ANSP 371520,
375562); 4 km NW of Sweetings Cay Light
(ANSP 307628); GREAT ABACO ISLAND:
Witch Point (ANSP 299481, 359153; USNM
4925800); Crossing Bay (ANSP 173189);
McLeans Town (ANSP 369066); Running
Mon Canal (ANSP 369777); North Hawksbill
Creek (ANSP 370565); BIMINI ISLANDS: Al-
icetown, North Bimini (USNM 598841); oppo-
site Cat Tail Pond, South Bimini (ANSP
325782); BERRY ISLANDS: Chub Cay (ANSP
359148); Frazier, Hog Cay (ANSP 194182,
195213); ANDROS ISLAND (ANSP 226713;
USNM 269844); Stafford Creek (ANSP

284 MARTINS

151848, 151930); Mangrove Cay (MCZ
24102; USNM 180518); Bastion Point, Man-
grove Cay (USNM 269226, 269252); Rocky
Point, Mangrove Cay (USNM 270214,
270215); Solomon Pond, Mangrove Cay
(USNM 269968); 5 km from mouth of Lisbon
Creek, Lindsey Creek (USNM 270234); First
island off Mintie Bar, SE of South Bight
(USNM 271785); Long Bay Key (USNM
269323); PARADISE ISLAND (А.М.); NEW
PROVIDENCE ISLAND (ANSP 18485,
299646; USNM 124376): Nassau (MCZ
107498; USNM 160767, 467111); Bar Point
(A.M.); Delaporte Point (A.M.); W of Rock
Point (A.M.); Clifton Point (A.M.); Clifton Pier
(A.M.); shore off Millars Road (A.M.); Millars
Sound by Bacardi Road (A.M.); Bonefish
Pond (A.M.); South Beach (MCZ 291268);
Malcolm Creek (A.M.); Dick’s Point (MCZ
291518); ROYAL ISLAND (MCZ 184098,
280395; USNM 343844, 366190, 468116);
ELEUTHERA ISLAND (USNM 465988): Tar-
pon Bay (MCZ 135934, 175921); Great Oys-
ter Pond (MCZ 291265); BRIGADINE KEY
(USNM 270034); CAT ISLAND: Arthurstown
(MCZ 291237); LONG ISLAND: Simms (MCZ
291251); Galloway’s Landing (MCZ 291241);
Pinders (MCZ 113328); AKLINS ISLAND:
Pinnacle Point (USNM 390857); ROOKERKEY
(USNM 390663, 390674a); GREAT INAGUA:
Matthewstown (MCZ 291263); 5 km SE of
Matthewstown (MCZ 190050); CAY SAL
BANK: Cay Sal, (MCZ 291256); Salt Lagoon,
Cay Sal (USNM 513426). TURK’S & CAICOS:
CAICOS ISLAND: Bell Cay (USNM 391323).
CUBA (ANSP 56810, 567800; MCZ 31418,
291257, 294214; USNM 10966, 39840,
55727, 336072, 492461): Cayo Juan Garcia
(MCZ 291271); La Habana (MCZ 291259);
Cayo Birricu, N. of Habana (ANSP 362823);
Cayo Blanco, Cärdenas Bay (ANSP 157955);
Playa del Bellamar (MCZ 291239, 291243);
Cayo Cristo (MCZ 292559); Caibarién (MCZ
291248, 294213); Dimas (USNM 492559b);
Cayo Romano (MCZ 291272); Punta de Pie-
dra (MCZ 291252); Santa Cruz del Sur (MCZ
131939, 291254); Santa Maria Key (MCZ
291261); Cochinos Bay (USNM 492548);
Cayo de las Cinco Léguas (ANSP 158053);
Finca, Sabanalmar (MCZ 294210); Isla de Pi-
nos (MCZ 48081). JAMAICA (ANSP 56811;
MCZ 291245; USNM 94746, 374270a,
492462, 492551): Montego Bay (ANSP
359146); Kingston (USNM 442736); Harbor
Head, Kingston (USNM 617127); Hunt's Bay
(USNM 441719); Cow Bay (USNM 440985);
Palisadoes (USNM 442465); Port Royal

(USNM 442419); Rock Fort (USNM 374243);
Great Goat Island (ANSP 359156). HAITI
(ANSP 146738): St. Louis (USNM 439392);
Gonave Island (MCZ 82118; USNM 380256);
near Port-au-Prince (USNM 403034, 403035,
440610a); lle-a-Vache, Soulette Bay (USNM
439169a, 439169b, 442850a); Port Salut
(USNM 403760); Aquin (USNM 403256,
403573, 440170); Bizoton (USNM 439828).
DOMINICAN REPUBLIC: Monte Cristi (MCZ
291249). PUERTO RICO: Punta Arenas, N of
Joyuda (A.M.). VIRGIN ISLANDS: ST. CROIX
(ANSP 56809). LESSER ANTILLES: AN-
GUILLA BANKS (MCZ 294216); ANTIGUA:
Fitches Creek (USNM 809737); BARBUDA
(USNM 735816). CARIBBEAN ISLANDS:
CAYMAN ISLANDS: Cayman Brac (MCZ
294212); Georgetown Barcadero, Grand
Cayman (ANSP 209770). COLOMBIA (MCZ
291273). VENEZUELA: Islas de los Roques
(USNM 784718). SURINAME: Paramaribo
(MCZ 274063).

Melampus (Detracia) floridanus
Pfeiffer, 1856
Figs. 315-318, 321-332

Melampus (Tralia) floridianus Shuttleworth. H.
8 A. Adams, 1854: 11 [nomen nudum].

Melampus floridanus Shuttleworth. Pfeiffer,
1854b: 147 [nomen nudum].

Tralia (Tifata) floridana Shuttleworth. H. 4 A.
Adams, 1855b: 245 [nomen пиаит].
Melampus floridanus “Shuttleworth” Pfeiffer,
1856a: 35 [Florida, herein restricted to
Myakka River; location of type un-
known]; Binney, 1860: 4; Nevill, 1879:
219; Dall, 1885: 281, pl. 18, fig. 2; Dall,
1889: 92, pl. 47, fig. 2; Simpson, 1889:
68; Kobelt, 1898: 213, pl. 24, fig. 14; Hin-
kley, 1907: 71; Maury, 1922: 55; C. W.
Johnson, 1934: 159; M. Smith, 1937:
146, pl. 55, fig. 5, pl. 67, fig. 2 [pl. 67 from

Dall (1885)].

Melampus floridianus Shuttleworth. Binney,
1859: 165 [error for floridanus; pl. 75, fig.
30 is of Melampus (M.) bidentatus Say
(Fig. 265)].

Tralia floridana (Shuttleworth) (Pfeiffer). Bin-
ney, 1865: 16 [fig. 17 is of Melampus (M.)
bidentatus]; Tryon, 1866: 9 [pl. 18, fig. 11
copied from Binney (1859) shows
Melampus (M.) bidentatus].

Detracia floridana (Pfeiffer). Morrison, 1951a:
17, figs. 4, 7 [description, habitat]; Mor-
rison, 1954: 15-16 [egg masses]; Morri-
son, 1959: 25 [early life history]; Burch,

WESTERN ATLANTIC ELLOBIIDAE 285

1960a: 182, pl. 1, figs. 2, 91 [chromo-
somes]; Abbott, 1974: 332 [Fig. 4093
copied from Binney (1859) is of Melam-
pus (M.) bidentatus]; Emerson & Jacob-
son, 1976: 191, pl. 26, fig. 23; Heard,
1982: 20, fig. 16.

Detracia floridana (Shuttleworth). Morrison,
1951b: 8.

Melampus floridanus Pfeiffer. Holle 8 Dineen,
1959: 50 [systematics].

Description: Shell (Figs. 315-318, 321-323)
to 7 mm long, globose to fusiform, thin,
smooth to corrugated, dark brown with gray-
ish tones, with as many as three chestnut-
brown bands on upper half of body whorl.
Spire moderately high, mucronate, with as
many as 10.25 flat, compressed whorls, with
fine, spirally arranged pits. Body whorl 70%
of total length, oval to subcylindric, lacking
carina on shoulder, smooth or with very faint
spiral lines. Aperture narrow, about 90% of
length of body whorl, weakly canaliculate at
base; inner lip with oblique columellar tooth,
conspicuous parietal callus and, above it and
hidden inside, small horizontal parietal tooth;
outer lip sharp, with as many as ten subequal
riblets, not reaching edge. Inner partition of
whorls occupying entire body whorl (Fig.
316). Protoconch raised, smooth, translu-
cent, dark brown (Figs. 321-323).

Animal bluish gray; foot paler; tentacles
subcylindric, pointed, darker toward tip;
mantle skirt grayish. Mantle organ dark
brown, well developed, forming conspicuous
pouch.

Radula (Figs. 324-328) having formula
[14+ (1 +12) +1 + (12 +1) + 14] x 100. Central
tooth with base wider than that of lateral
teeth, with conspicuous medial promi-
nences; crown half length and width of that of
lateral teeth, rounded posteriorly; mesocone
sharp; ectocones well marked, small. Lateral
teeth 11 to 16; crown strong, half total length
of tooth; mesocone sharp, pointing laterally;
ectocone well developed in all lateral teeth.
Transitional tooth with very weak endocone.
Marginal teeth 13 to 17, with reduced base
and elongate crown; mesocone becoming
shorter and thinner; endocone and second
cusp of ectocone in first marginal tooth; outer
edge of base assuming configuration of den-
ticle around third marginal tooth, first cusp of
ectocone becoming smaller and third cusp of
ectocone appearing; second cusp of en-
docone appearing on fifth to sixth marginal
tooth; eighth to tenth marginal teeth without

additional cusps; last two marginal teeth ru-
dimentary.

Digestive system as in Melampus s. S.;
stomach elongate, muscular band thick (Fig.
329).

Reproductive system (Fig. 330) with
ovotestis shallow-conical, dark brown; albu-
men gland spiral, conical; fertilization cham-
ber forming double pouch; vagina and asso-
ciated vas deferens muscular, long; bursa
duct entering vagina opposite exit of poste-
rior vas deferens; penis muscular, long;
length of anterior vas deferens 75% that of
penis.

Nervous system (Fig. 331) having cerebral
commissure about as long as width of cere-
bral ganglion; left pleuropedal connective
very short; left parietovisceral connective
longer than right one.

Remarks: The name Melampus floridanus ap-
peared in published lists for several years be-
fore Pfeiffer (1856a) validated it with a de-
scription in his Monographia. Holle & Dineen
(1959) stated that the original specimens had
been collected in Florida by a Mr. Rugel for
Shuttleworth, who deposited them in the
Cuming collection under his manuscript
name. Pfeiffer (1856a) probably had seen
these specimens marked as Auricula floridana
by Shuttleworth, and also the specimens in
the Albers collection marked as Auricula rugeli
by Charpentier. Both names are referred to as
manuscript names in Pfeiffer's description.
He gave the measurements of the specimen
he used for the description, however, men-
tioning that it was Nr. 15 of his collection. The
type material therefore should include only
the specimen in Pfeiffer’s collection, because
there is no assurance that Pfeiffer used other
collections in writing the description. Most au-
thors have given credit erroneously to Shut-
tleworth for introduction of Melampus (D.) flo-
ridanus but in accord with the ICZN Pfeiffer
must be credited with this name, for it is he
who validly introduced it.

According to Morrison (1951a), Binney
(1859), using shells collected by Bartlett from
the Florida Keys, wrongly illustrated this spe-
cies by using an example of a dwarf Melam-
pus (M.) bidentatus (Fig. 265). Several subse-
quent authors, including Abbott (1974),
perpetuated Binney's mistake by copying
that figure. Morrison (1951a), Emerson & Ja-
cobson (1976) and Heard (1982) correctly il-
lustrated Melampus (D.) floridanus, however.

Melampus (D.) floridanus can be distin-

286 MARTINS

FIGS. 315-323.

WESTERN ATLANTIC ELLOBIIDAE 287

FIGS. 324-327. Melampus (D.) floridanus, radular teeth. (324-326) Myakka River, Florida, sl 6.7 mm. (327)
Woodville, North River, North Carolina, sl 5.3 mm. Scale, Fig. 324, 50 um; all others, 100 um.

guished from Melampus (M.) bidentatus, with
which it commonly associates, by its smaller
size, stronger columellar tooth, the callus оп
the site of the first parietal tooth, its narrower
aperture and more numerous whorls. The
specimens of Melampus (D.) floridanus from
southern and western Florida are smooth and
sometimes brightly colored, and usually re-
tain all the whorls of the spire, whereas north-
ern specimens are thinner and corrugated,
with the apex greatly eroded.

Habitat: Melampus (D.) floridanus lives in salt
marshes and freshwater riverbanks on which
it often occurs with Melampus (M.) bidenta-
tus. The Floridian salt-marsh snail prefers
that zone of the marsh rarely flooded by

spring tides. The animals frequently live half-
buried in the sediment, against the base of
the stems of Spartina, Juncus and other
marsh plants. Very common in some places,
they were estimated by Morrison (1951a) to
attain a density of about four billion individu-
als in a square mile.

Range: New Jersey to Florida, and along the
Gulf Coast to Vera Cruz, Mexico (Fig. 332). |
have not observed specimens from the Flor-
ida Keys.

Specimens Examined: NEW JERSEY: Divid-
ing Creek (A.M.); Newport (ANSP 294331).
DELAWARE: Woodland Beach (USNM
522268); Bombay Creek (USNM 473356,

FIGS. 315-323. Melampus (Detracia). (315) M. (D.) floridanus Pfeiffer, Woodville, North River, North Caro-
lina, sl 5.2 mm. (316) M. (D.) floridanus, Woodville, North River, North Carolina, sl 5.5 mm. (317) M. (D.)
floridanus, Myakka River, Florida, sl 6.7 mm. (318) М. (D.) floridanus, Myakka River, Florida, sl 5.7 mm. (319)
Detracia parana Morrison, holotype (USNM 594591), Pará, [Belém], Brazil, sl 6.5 mm. (320) Auricula glob-
ulus Orbigny, lectotype (ВММН 1854.12.4.243), Guayaquil, Ecuador, sl 8.1 mm. (321-323) М. (D.) florida-
nus, lateral and top views of spire and protoconch, Myakka River, Florida. Scale 1 mm.

288 MARTINS

C1L2L3L 12L Т 1М 2M3M4M

ARO dre

8M 9M 10M 11M 12M 13M

ЦИИ

FIG. 328. Melampus (D.) floridanus, radula, Cres-
cent, Georgia. Scale 10 um.

A
EA
NE AS}
ES > —
Fa

FIG. 329. Melampus (D.) floridanus, stomach,
Georgia. Scale 1 mm.

628864, 628865, 628866); Augustine Pier
(ANSP 89556; MCZ 294219; USNM 492587).
MARYLAND: Morgan Creek, Charlestown
(MCZ 200469); Mayo Beach (USNM 522289);
Galesville (USNM 595601); Chesapeake
Beach (USNM 473812); Parkers Creek
(USNM 536708); N of Benedict, Patuxent
River (USNM 473459, 473460, 473461);

Benedict (USNM 473463); Kepplers,
Broomes Island (USNM 473466); Helen
Creek, N of Solomons Island (USNM

600764); Solomons Island (USNM 4240410;
Millstone Landing, mouth of Patuxent River
(USNM 521877); Cobb Island, Potomac River

FIG. 330. Melampus (D.) floridanus, reproductive
system, Sapelo Island, Georgia. Scale 1 mm.

1
Г р

ре pc pg pig сре cpc cg cbc

FIG. 331. Melampus (D.) floridanus, central nervous
system, Sapelo Island, Georgia. Scale 1 mm.

(USNM 473565, 473566, 499520, 522920);
Bretton Bay, Potomac River (USNM 628901);
Chapel Point, Potomac River (USNM
758317); opposite Chestertown (ANSP
106973); Town Point (MCZ 46419); Head of
Little Choptank River, Cambridge (USNM
348955); Cambridge (MCZ 52355, 291275);
Dailsville (ANSP 1332468, 303357; MCZ

WESTERN ATLANTIC ELLOBIIDAE 289

FIG. 332. Geographic distributions, Melampus (D.)
floridanus (circles), Melampus (D.) paranus (stars).
Open symbols, localities from literature.

55922); Chambers Farm, Dailsville (ANSP
65100); Whitehaven (MCZ 291276); East of
Dames Quarter (USNM 618923). VIRGINIA:
Colonial Beach, Potomac River (USNM
473890, 473891); Potomac Beach (USNM
473901, 473902, 473903); Poropotank River
(USNM 679376); 2 mi NE of Bartlett (USNM
595938); Yorktown (USNM 474078); Norfolk,
Lafayette River (USNM 667496). NORTH
CAROLINA: Cedar Island (A.M.); Williston
(A.M.); Woodville, North River (A.M.); Beau-
fort (MCZ 294261; USNM 678946); Morton's
Hill, near Beaufort (USNM 621431). SOUTH
CAROLINA: Yemassee (А.М.). GEORGIA:
Crescent (A.M.); Fort King George, at Darien
(USNM 628867). FLORIDA: Jacksonville
(ANSP 132461); Clear Lake (MCZ 294218;
USNM 30210); Miami (ANSP 77039; USNM
153403); Everglades fork of Miami River
(ANSP 82852); Biscayne Вау (USNM
492586); Seminole Point (ANSP 293554);
Flamingo Key (MCZ 291041); Cape Sable
(MCZ 291039); Turner River, near Chokolos-
kee (ANSP 93436); Everglades City (MCZ
291040, 294262); Naples (MCZ 291041); Fort
Myers (ANSP 62805; MCZ 291277; USNM
492585); Little Gasparilla Island (ANSP
142169); Myakka River (A.M.); Sarasota Bay

(ANSP 294332; USNM 30624); Big Bend
Road, Tampa Bay (A.M.); Tampa (ANSP
76114; MCZ 70562; USNM 37608, 504488);
Ballast Point, Tampa (MCZ 13815); Hudson
(A.M.); Tributary to Hudson Bayou (USNM
487336); Aripeka (ANSP 73901; USNM
149953); Little Blind Creek, below mouth of
Chassahowitzka River (ANSP 148526); Tar-
pon Springs (MCZ 291274); Suwannee River
(ANSP 189567); St. Marks (ANSP 56815,
56816). ALABAMA (USNM 492588): SE of
Heron Bay (ANSP 315714); Mobile (MCZ
68065); Coden Beach (USNM 422365).
MISSISSIPPI: Point Cadet, Biloxi (USNM
518640); Davis Bayou, Ocean Springs
(USNM 778280); Escatawpa River (ANSP
315718). LOUISIANA: New Orleans (USNM
119495). MEXICO: Tampico (ANSP 46584);
Rio Vinazco (USNM 675265); SE of Tuxpan,
Vera Cruz (USNM 675272).

Melampus (Detracia) paranus
(Morrison, 1951)
FIGS. 319, 332, 333

Detracia parana Morrison, 1951a: 19, fig. 3
[Amazon River at Pará, Brazil; holotype
USNM 594591 (Fig. 319); three para-
types USNM 32090]; Morrison, 1951b: 9;
Marcus 8 Marcus, 1965a: 42-51, figs.
19-21, 23-25 [distribution, ecology,
anatomy]; Rios, 1970: 138; Rios, 1975:
158, pl. 48, fig. 766.

Melampus (Detracia) paranus (Morrison). Al-
tena, 1975: 86.

Description: Shell (Fig. 319) to 7 mm long,
globose to fusiform, thin, smooth, yellowish
brown with one to three darker brown bands
on body whorl, the one nearest suture more
conspicuous. Spire low, with as many as ten
flat whorls, dark brown with lighter band.
Body whorl 80-90% of shell length, fusiform
to subcylindric, without hint of carina on
shoulder. Aperture 85-90% of length of body
whorl, narrow, weakly canaliculate at base;
inner lip with strong, oblique columellar tooth
and small, horizontal parietal tooth hidden in-
side aperture; outer lip sharp, usually with
one riblet opposite columellar tooth, some-
times with none, rarely with two or three.

Animal with tentacles bulbous at base;
eyes surrounded by unpigmented skin. Vis-
ceral mass separated from foot by one whorl,
with corresponding extension of mantle cav-
ity; mantle organ forming funnel-shaped
pouch.

290 MARTINS

a UE el 12L 4M a и
ANA ee Gor
Mm ae | N (hoe Cyd

о \ LIA

FIG. 333. Melampus (D.) paranus (Morrison), rad-
ula, Cananeia, Brazil; redrawn from Marcus & Mar-
cus (1965a). Scale 10 um.

Radula (Fig. 333) having formula
[16 + 16 + 1 + 16 + 16] x 100. Base of central
tooth weakly concave, not emarginate;
crown rounded posteriorly; mesocone small,
rounded; ectocones absent. Mesocone of
lateral teeth about half of length of tooth, with
conspicuous ectocone. Endocone appearing
on second marginal tooth; as many as five
ectocones on marginal teeth.

Remarks: Melampus (D.) paranus was de-
scribed by Morrison (1951a) on the basis of
four specimens in the United States National
Museum of Natural History, collected in Bra-
zil before 1885. Only the type specimens
were available to me and they constitute the
basis for my description of the shell. All data
on the animal and its anatomy were taken
from Marcus 4 Marcus (1965a). In their study
of 174 specimens from Cananeia, Brazil,
Marcus 8 Marcus observed the aperture
length to be barely 75% of the length of the
body whorl, an important difference from the
few specimens that constitute the type ma-
terial. | observed similar variation in the North
American companion species Melampus (D.)
floridanus.

According to Morrison (1951a) the strong
similarities between Melampus (D.) paranus
and Melampus (D.) floridanus suggest that
both species underwent closely parallel evo-
lution. The former differs from the North
American species by its lack of а callosity
above the columellar tooth and by the num-
ber of riblets inside the outer lip. Marcus &
Marcus (1965a) observed that Melampus (D.)
paranus commonly had one riblet, seldom
none and rarely two, and only one of the 174
specimens examined had three riblets inside
the outer lip. Melampus (D.) floridanus has
four to ten riblets inside the outer lip.

Habitat: According to Marcus & Marcus
(1965a), Melampus (D.) paranus is a supralit-
toral estuarine species that lives in man-
groves together with Melampus (M.) coffeus.

Range: Suriname (Altena, 1975), south to
Cananeia, Brazil (Marcus & Marcus, 1965а)
(Fig. 332).

Specimens Examined: BRAZIL: Pará [Belém],
on the Amazon River (USNM 32090, 594591).

Melampus (Detracia) monile
(Bruguiere, 1789)
Figs. 334-354

Bulimus monile Bruguiére, 1789: 338 [West
Indies, herein restricted to San Juan, Pu-
erto Rico; location of type unknown];
Dillwyn, 1817: 506 [erroneously stated
as a probable variety of Voluta flava
Gmelin, 1791]; Cuvier, 1817: 414.

Melampa monile (Bruguiére). Schweigger,
1820: 739.

Auricula monile (Bruguière). Férussac, 1821:
105; Lamarck, 1822: 141; Kuster, 1844:
30, pl. 4, figs. 7-9.

Auricula monile Lamarck. Menke, 1830: 36;
Gould, 1833: 67; Jay, 1839: 59; Reeve,
1842: 106, pl. 187, fig. 8.

Melampus monile Schweigger. Lowe, 1832:
292.

Conovulus monile (Bruguiere).
1836: 71, pl. 27, 165. 55а

Melampus (Melampus) топйе (Bruguiere).
Beck, 1837: 108.

Auricula monile Férussac. Potiez & Michaud,
1838: 202.

Melampus coronatus C. B. Adams, 1849: 41
[Jamaica; lectotype chosen by Johnson
8 Boss (1972) MCZ 186029 (Fig. 342); C.
B. Adams, 1851: 186; Pfeiffer, 1854b:
147; Pfeiffer: 1856a: 51; Johnson 4
Boss, 1972: 196, pl. 41, fig. 5 [lectotype
figured].

Melampus flavus Gmelin of authors. C. B. Ad-
ams, 1849: 42; C. B. Adams, 1851: 186;
H. & A. Adams, 1854: 9; Pfeiffer, 1854b:
147; Pfeiffer, 1856a: 21; Binney, 1859:
167, text fig.; Binney, 1860: 4; Binney,
1865: 12, fig. 14; Tryon, 1866: 8, pl. 18,
fig. 6; Poey, 1866: 394; Pfeiffer, 1876:
303; Mórch, 1878: 5; Arango у Molina,
1880: 59; Dall, 1883: 322; Dall, 1885:
281, pl. 18, fig. 1; Dall, 1889: 92, pl. 47,
fig. 1; Simpson, 1889: 68; Crosse, 1890:
258; Davis, 1904: 126, pl. 4, fig. 5; Peile,
1926: 88; Maury, 1922 55, (GW:
Johnson, 1934: 159; M. Smith, 1937:
146, pl. 55, fig. 12, pl. 67, Ig Piper
copied from Dall (1885)]; Holle 8 Dineen,
1959: 28-35, 46-51. Non Gmelin, 1791.

Deshayes,

WESTERN ATLANTIC ELLOBIIDAE 291

Melampus torosa Mórch, 1852: 38 [Antilles;
location of type unknown].

Melampus fusca Mórch, 1852: 35 [Antilles;
location of type unknown].

Melampus coronulus С. В. Adams. H. & A.
Adams, 1854: 10 [error for coronatus].

Melampus monilis Lamarck. Shuttleworth,
1854b: 102; Shuttleworth, 1858: 73 [un-
justified emendation of monile].

Melampus monile Lamarck. Mórch, 1878: 5.

Melampus flavus (Gmelin?) Binney. Dall 8
Simpson, 1901: 368, pl. 54, fig. 9. Non
Gmelin, 1791.

Melampus flavus var. purpureus Davis, 1904:
126, pl. 4, fig. 6 [Bermuda, herein re-
stricted to South Shore; lectotype se-
lected by Baker (1964) ANSP 86922 (Fig.
336).

Melampus flavus var. albus Davis, 1904: 126,
pl. 4, fig. 7 [Hungry Bay, Bermuda; lec-
totype selected by Baker (1964) ANSP
86924 (Fig. 337)].

Pira monile (Bruguière). Morrison, 1951b: 8;
Morrison, 1958: 118-124 [ecology];
Nowell-Usticke, 1959: 88; Morrison,
1964: 119-121 [systematics].

Melampus monile (Bruguiere). Warmke 8 Ab-
bott, 1961: 153, pl. 28, fig. p; Rios, 1970:
138; Morris, 1973: 273, pl. 74, fig. 9; Em-
erson & Jacobson, 1976: 192, pl. 26, fig.
27; Rehder, 1981: 647, fig. 363.

Melampus (Pira) monilis (Bruguiere). Abbott,
1974: 332, fig. 4090: Rios, 1975: 158, pl.
48, fig. 765; Gibson-Smith & Gibson-
Smith, 1982: 116, figs. 2, 3; Vokes 4
Vokes, 1983: 60, pl. 22, fig. 14.

Melampus (Pira) monile (Вгидшеге). Hum-
phrey, 1975: 196, pl. 22, fig. 23 [shell fig-
ured seems to be Melampus coffeus].

Melampus monilis (Bruguière). Cosel, 1978:
216; Mahieu, 1984: 314; Jensen 4 Clark,
1986: 457, figured.

Description: Shell (Figs. 334-345) to 16 mm
long, ovoid to fusiform, solid, shiny, usually
a uniform purplish brown or with as many
as three narrow white bands, rarely uniform-
ly white or yellowish. Excavated umbilical
groove sometimes present in gerontic spec-
imens. Spire low to moderately high, with as
many as 11.25 flat whorls, with two or three
well-marked spiral grooves on first two
whorls, one or two rows of elongated pits on
remaining whorls; spiral row of short, laterally
compressed periostracal hairs, often running
along spiral row of pits in adult specimens;
location of hairs does not correspond to that

of pits. Body whorl averaging 83% of shell
length, conic to ovoid, smooth or with pitted
and sometimes carinate shoulder. Aperture
about 90% length of body whorl, narrow, an-
teriorly angulate; inner lip with strong, ob-
lique, twisted columellar tooth, conspicuous
parietal callus and, just posterior to it, deep
parietal tooth; outer lip sharp, with as many
as ten subequal riblets not reaching edge.
Inner partition of whorls occupying two-
thirds of body whorl (Fig. 340). Protoconch
translucent, brownish (Figs. 343-345).

Animal bluish gray; foot whitish; top of
neck blackish; tentacles subcylindric,
pointed, translucent in first quarter, changing
sharply to dark gray or black; mantle skirt
light gray. Pallial cavity elongate; mantle or-
gan forming conspicuous pouch; kidney very
elongate.

Radula (Figs. 346-350) having formula
29 + (1 + 15) + 1 + (15 + 1) + 29] x 113.
Central tooth base twice width of lateral teeth
base, triangular, with conspicuous promi-
nences on inner surfaces of arms; crown nar-
rower and smaller than that of lateral teeth,
with posterior edge straight or with weak me-
dial depression; mesocone small, pointed;
very weak ectocones sometimes present.
Lateral teeth 14 to 17; crown broadly trian-
gular, half total tooth length; mesocone
sharp, pointed laterally; first lateral tooth with
medial posterior part of crown elongate, with
weak endocone; remaining lateral teeth with-
out endocone, posterior edge of crown with
medial prominence, medial posterior part of
base flaring, cusp-shaped; no ectocone.
Transitional tooth with weak ectocone. Mar-
ginal teeth 24 to 32; base reduced, crown
very elongate in first teeth, gradually becom-
ing smaller; mesocone strong, sharp, gradu-
ally becoming rounded at tip.

Digestive system as in Melampus s. $.;
stomach (Fig. 351) as in subfamily.

Reproductive system (Fig. 352) with
ovotestis leaf-like, round, dark brown; albu-
men gland spiral; prevaginal caecum con-
spicuous; bursa duct connecting with vagina
opposite exit of posterior vas deferens; bursa
large, oval-elongate; vagina thin, long, about
same size as posterior vas deferens; penis
thin, long; anterior vas deferens about 65%
of penis length; penial retractor attaching to-
gether with columellar muscle.

Nervous system (Fig. 353) having cere-
bral commissure narrower than width of ce-
rebral ganglion; left parietovisceral connec-
tive twice the length of right one.

292

MARTINS

FIGS. 334-345.

WESTERN ATLANTIC ELLOBIIDAE 293

FIGS. 346-349. Melampus (D.) monile, radular teeth, Shelly Bay, Bermuda, sl 11.2 mm. (346) Central and
lateral teeth. (347) Transitional and marginal teeth. (348,349) Marginal teeth. Scale 50 um.

Remarks: Bruguiere (1789) clearly stated
that his Bulimus monile was from the West
Indies, but the works he cited refer to both
the West Indies (Lister, 1770: pl. 834, figs. 60,
61, Barbados) and East Indies (Martini, 1773:
2, р. 126, pl. 43, fig. 444, East Indies). It was
probably this discrepancy that led Dillwyn
(1817) to suggest that Bruguière’s Bulimus
monile was only a variety of Voluta flava
Gmelin, 1791. This, in turn, led to the general
confusion of Bulimus monile with Voluta flava
and the general use of the latter name for the
West Indian species. Gmelin (1791: 3431),
however, under Voluta flava referred only to
figure 444 of Martini (1773), which definitely
represents an East Indian species.

Another explanation for the confusion of

the two species, besides common reference
to Martini's fig. 444, might reside in the vari-
able color pattern of the West Indian species.
The juveniles of Melampus (D.) monile, like
those of Melampus flavus, are often golden or
golden brown, as seen in C. B. Adams'
Melampus coronatus [= Melampus (D.) mo-
nile, juvenile]. Bruguière (1789) in the original
description stated that his specimens of
Melampus (D.) monile were not fully grown,
because they lacked the inner dentition о the
outer lip, a feature reported by Lister (1770)
and Martini (1773). This might explain his
statement about the “very light yellow” color.

Binney (1859), using the name Melampus
flavus Gmelin, 1791, for the West Indian spe-
cies, listed Melampus torosa Mörch and

FIGS. 334-345. Melampus (D.) monile (Bruguiére). (834) Specimen perhaps figured by Binney (1859:167,
fig. IV) (USNM 39827), sl 12.8 mm. (335) Shelly Bay, Bermuda, sl 14.4 mm. (336) М. flavus purpureus Davis,
lectotype (ANSP 86922), South Shore, Bermuda, sl 10.0 mm. (337) М. flavus albus Davis, lectotype (ANSP
86924), South Shore, Bermuda, sl 8.6 mm. (338) San Juan, Puerto Rico, sl 12.6 mm. (339) San Juan, Puerto
Rico, $1 12.5 mm. (340) San Juan, Puerto Rico, sl 12.2 mm. (341) Juvenile, Maravén, Venezuela, sl 1.67 mm.
(342) M. coronatus C. B. Adams, lectotype (MCZ 186029), Jamaica, sl 3.0 mm. (343,344) Lateral and top
views of spire and protoconch, Indian Key Fill, Florida. (345) Detail of spire and protoconch of specimen of

Fig. 341. Scale 500 um.

294 MARTINS

11M 13M 18M 20M 26M 28M

Aaa

FIG. 350. Melampus (D.) monile, radula, Shelly Bay,
Bermuda. Scale 10 um.

FIG. 351. Melampus (D.) monile, stomach, Ber-
muda. Scale 1 mm.

Melampus (D.) monile (Bruguiere) as syn-
onyms. Under Melampus torosa, Mörch
(1852) cited figure 444 of Martini (1773), in-
cluded Voluta flava Gmelin in the synonymy
and mentioned the Antilles as the locality. It
appears, then, that Mórch also confused
Melampus flavus (Gmelin) with Melampus (D.)
monile (Bruguière) and Mórch's name must
be treated as a synonym of Melampus (D.)
monile (Bruguière, 1789).

Another name listed in Mórch's (1852)

|

FIG. 352. Melampus (D.) monile, reproductive sys-
tem, Clifton Pt., New Providence, Bahamas. Scale
1 mm.

рус prg pig cple сре cc cg

y LA
\ A у \

eee

1 tha
vg plpre pipe

FIG. 353. Melampus (D.) monile, central nervous
system, Clifton Pt., New Providence, Bahamas.
Scale 1 mm.

Yoldi Catalogue is Melampus fusca from the
Antilles. In the synonymy he cited Martini
(1773, pl. 43, fig. 445), Voluta minuta Gmelin,
1791, which prompted Binney (1859) to in-
clude Melampus fusca as a synonym of
Melampus (M.) coffeus, and Voluta monile.
Figure 445 of Martini, upon which Voluta

WESTERN ATLANTIC ELLOBIIDAE 295

minuta Gmelin was based, has already been
shown to be unidentifiable [see the remarks
for Melampus (M.) coffeus]. Thus, the only
citation under Melampus fusca that can sup-
port the name is Voluta monile, of which
Melampus fusca Mórch must be considered
a synonym.

Dall & Simpson (1901) found it difficult to
separate Melampus (D.) monile; erroneously
listed as Melampus flavus, from Melampus
(M.) coffeus, owing to similarities in shape
and color. They noted that the apertural den-
tition is the most reliable distinguishing char-
acter. Bruguiere (1789) for Melampus (D.
monile mentioned two teeth, a small, oblique
columellar tooth and a smaller parietal tooth.
Usually Melampus (М.) coffeus has one
small, more or less oblique columellar tooth,
and two readily visible parietal teeth, the pos-
terior one the largest of the three. Some-
times, however, the anterior parietal tooth is
either very small or absent, hence the source
of confusion. The twisted columellar tooth,
the much smaller, hidden parietal tooth and
the hairs (noticed by Shuttleworth in 1858) or
pits on the shoulder of the body whorl and
spire unmistakeably distinguish Melampus
(D.) monile from Melampus (M.) coffeus,
Melampus (M.) bidentatus and Melampus (D.)
morrisoni. It differs from Melampus (D.) bul-
laoides in its more conical shape, longer ap-
erture and evenness of the riblets on the
outer lip. The hairs and general aspect of the
spire are similar to those of the latter species,
but Melampus (D.) monile has a more regular,
lower conical spire.

Melampus (D.) monile was placed by Mor-
rison (1951b) within the genus Pira H. 8 A.
Adams. The reasons that | do not accept that
decision are discussed in the remarks for De-
tracia. Melampus (D.) monile is placed in the
subgenus Detracia on the basis of shell, rad-
ular and anatomical characters. The strong,
twisted columellar tooth, the medial promi-
nences on the base of the central radular
tooth, the pouch-like mantle organ and the
greater separation between foot and visceral
mass are all typical characters of Detracia.

Habitat: Melampus (D.) monile is unique
among species of Detracia in its preference
for living much closer to the high-tide mark
than do any other species of the subgenus,
which usually live farther inland. Melampus
(D.) monile commonly lives under boulders
above the high-tide mark along open rocky
shores, together with Tralia (T.) ovula and Pe-

90 75 60 45 30

FIG. 354. Melampus (D.) monile, geographic distri-
bution. Open circle, locality from literature.

dipes mirabilis. It can also occur in man-
groves, but always near the high-tide mark.

Range: Bermuda; Florida; West Indies, Cen-
tral America to Guanabara Bay, Brazil (Rios,
1975) (Fig. 354).

Specimens Examined: FLORIDA: Indian
River (MCZ 201670); Miami (ANSP 47603,
294316; USNM 492589); Coconut Grove
(MCZ 201093, 291380, 294258); Brickell
Hammock, Biscayne Bay (MCZ 291382); 6
km S of Tavernier, Key Largo (MCZ 291385);
Tavernier Key (USNM 492549a); Plantation
Key (MCZ 294255); S of Ocean Drive, Plan-
tation Key (A.M.); Tea Table Key (MCZ
291009); Indian Key Fill, N of Indian Channel
(А.М.); Long Key (MCZ 291010; A.M.); Grassy
Key (A.M.); Crawl Key (MCZ 294256; А.М.);
Knight Key (A.M.); West Summerland Key
(MCZ 291388; A.M.); Big Torch Key (ANSP
189590); Key West (ANSP 56804; 174363;
USNM 36062a, 596785); Boca Grande, Gas-
parilla Island (ANSP 142273). TEXAS: Port
Maria (USNM 711207). BERMUDA (ANSP
56790, 85587; MCZ 24155, 24249, 24250;
USNM 11421a, 94432b, 173648, 173649,
173650, 228688): Fairyland (ANSP 99082); N
of Shelly Bay Beach (A.M.); Coney Island

296 MARTINS

(A.M.); Ferry Reach Park (R.B.); N end of
Long Bird Bridge (A.M.); St. George's Island
(USNM 621572); Castle Harbor (ANSP
143319); Hungry Bay (USNM 171939,
492555; A.M.); Agar's Island, Hungry Bay
(MCZ 48124, 106521; A.M.); South Shore
(ANSP 86922, 86924; MCZ 53063, 53064;
USNM 109560, 109561); Boat Bay (USNM
621691); S end of Ely's Harbour (A.M.); W
side of Somerset Bridge (A.M.); Mangrove
Bay (A.M.). BAHAMA ISLANDS (ANSP
56799; MCZ 9946; USNM 37607): GRAND
BAHAMA ISLAND (ANSP 374527): Running
Mon Canal (ANSP 369779); Eight Mile Rock
(ANSP 173262; MCZ 116712); Caravel Beach
[John Jack Point], Freeport (ANSP 370226);
Dead Mans Reef [Sandy Bevan's Cay] (ANSP
371225); McLeans Town (ANSP 369067);
GREAT ABACO ISLAND (MCZ 24140; USNM
492580): Hope Town Harbor (ANSP 299391);
Little Harbor (USNM 180520); Witch Point
(ANSP 299483, 359150); Matt Lowes Cay
(ANSP 299248); Marsh Harbor (MCZ
275572); BIMINI ISLANDS: Alicetown, North
Bimini (MCZ 144132); opposite Cat Tail
Pond, South Bimini (ANSP 325784); AN-
DROS ISLAND (MCZ 66755, 71633): Mor-
gan's Bluff (A.M.); Mastic Point (USNM
359884); South Mastic Point (A.M.); First is-
land off Mintie Bar, SE end of South Bight
(USNM 271785b); Mangrove Cay (USNM
2699680); Lisbon Point, Mangrove Cay
(USNM 269599a); Bastion Point, Mangrove
Cay (USNM 269260); Long Bay Key (USNM
269304); PARADISE ISLAND: (A.M.); NEW
PROVIDENCE ISLAND (USNM 603913): Nas-
sau (USNM 160765a); Culbert Point, 10 km
ESE of Nassau (MCZ 107793); Bar Point
(A.M.); Delaport Point (A.M.); W of Rock Point
(A.M.); Clifton Point (A.M.); Clifton Pier (A.M.);
shore off Millars Road (A.M.); Malcolm Creek
(A.M.); Coral Harbor (USNM 679136); ROYAL
ISLAND (USNM 468115a); CAT ISLAND:
Arthurstown (MCZ 107833); 6 km E of Arthur-
stown (MCZ 107825); RUM CAY (MCZ
87849); LONG ISLAND: 3 km NE of O’Neill’s
(ANSP 173265; MCZ 113102); Simms (MCZ
294259); Clarencetown (MCZ 113339);
GREAT INAGUA: Matthewstown (MCZ
291384). TURK’S 8 CAICOS: TURK'S IS-
LAND (MCZ 201098; USNM 492474,
509960). CUBA (ANSP 56801, 56802; USNM
492478): Jaimanitas (MCZ 294198); Habana
(ANSP 93653); Cayo Birricu, N of Habana
(ANSP 362824); Cojimar (ANSP 45089; MCZ
131921, 131955); Matanzas (ANSP 87896;
MCZ 294192); La Playa (MCZ 92045,

131950, 189818, 294260); Versalles (MCZ
291381); Cayo Cristo (MCZ 291389); Vara-
dero (ANSP 110604; MCZ 201677; USNM
598261); Cayo Galindo, Cärdenas Вау
(ANSP 157578); Cayo Frances (MCZ 42106,
131951); Siboney (USNM 533912); Agua-
dora, Santiago (USNM 391879); Guantán-
amo (ANSP 313059); Cabo Cruz (MCZ
87887); Rancho Alma, Cienfuegos (MCZ
291386). JAMAICA (ANSP 56803; MCZ
186029, 201094, 294254, 294257; USNM
6385, 94744, 492475, 492477, 492480):
Montego Bay (USNM 441488); Falmouth
(ANSP 397268); Robin’s Bay (USNM 442026,
442092); Jack's Bay (USNM 441926); Buff
Bay (USNM 441196); Port Antonio (USNM
440855); Priestmans River (USNM 492479);
Manchioneal (R.B.); Rock Fort (USNM
374242); Kingston (USNM 374270, 442730);
Harbor Head, Kingston (USNM 375579); Pal-
isadoes (USNM 442466); Runaway Bay
(USNM 202658); Little River (USNM 128046).
HAITI: Gonave Island (492531 a); lle-a-Vache,
Soulette Bay (USNM 439191a); Port Salut
(ANSP 226701; MCZ 183912); Les Cayes
(USNM 439742); Bale Anglaise, near Aquin
(USNM 439548a); Saltrou (USNM 439342,
442819); Bizoton (USNM 439832a). DOMIN-
ICAN REPUBLIC: Monte Cristi (MCZ 57591,
57750, 291383); Puerto Plata (MCZ 291379,
291387); Santa Bárbara de Samaná (ANSP
173263; MCZ 57757); Cayo Chico, E of Santa
Bárbara de Samaná (MCZ 57776); Cayo de
Tamiso (MCZ 57812); Isla La Matica, Playa
Boca Chica, E of Santo Domingo (R.B.). PU-
ERTO RICO: E of San Juan (USNM 683107);
Puerta de Tierra, San Juan (A.M.); Punta
Cerro Gordo (USNM 683012); Punta Agu-
jereada (MCZ 233338); Arecibo (MCZ
291391); Punta Arenas, N of Joyuda (A.M.);
Lighthouse, Cabo Rojo (MCZ 294194;
Ensenada Honda, Culebra Island (USNM
161161). VIRGIN ISLANDS: ST. THOMAS
(ANSP 56789; MCZ 294196; USNM 256035);
ST. CROIX (ANSP 56788; USNM 621396).
LESSER ANTILLES: ANTIGUA (MCZ 71511):
off Falmouth (USNM 502116); North Bay,
Guana Island (MCZ 88870; ANSP 351799);
GUADELOUPE (MCZ 294197; USNM
492481): Anse-Dumont, Gosier (USNM
758065); ST. VINCENT: (USNM 492473); Villa
(USNM 487000); BARBADOS: (ANSP 56797;
MCZ 291390; USNM 502108, 502109);
Bridgetown (USNM 502115); Pelican Island
(USNM 502110, 502112); Needham Point
(USNM 502113); off Telegraph Station
(USNM 502114); Maxwell’s Coast (USNM

WESTERN ATLANTIC ELLOBIIDAE 297

603784); San Blas (ANSP 56796); GRENADA:
Prickly Bay (ANSP 297189); off Hardman Bay
(ANSP 296483); TRINIDAD (MCZ 294193).
CARIBBEAN ISLANDS: SWAN ISLAND (MCZ
22938, 36611); CAYMAN ISLANDS: Cayman
Brac (ANSP 296178; MCZ 294195); OLD
PROVIDENCE ISLAND: N of Ironwood Point
(USNM 678831); ST. ANDREWS ISLAND
(ANSP 154359; MCZ 88689); СУВАСАО:
Port Marie 8 Daaibooi Ваа! (В.В.). MEXICO:
Isla Mujeres, Yucatán (ANSP 284638); As-
cension Bay, Quintana Roo (USNM 736381,
736718). BELIZE: North Spot (ANSP
281604); Belize (USNM 150281). HONDU-
RAS: Utila Island (USNM 61185); Roatan Is-
land (USNM 364701). COSTA RICA: Portete,
Limón (USNM 702853, 706403). PANAMA:
Limón Bay (USNM 732871, 734073); Fort
Sherman, Devil's Beach, 9 km N of Colón
(USNM 620529). COLOMBIA: Sabanilla
(USNM 103467, 193611). VENEZUELA: Cayo
Punta Brava, Parque Nacional de Morrocoy,
Tucacas (A.M.); El Palito (A.M.); Borburata
(USNM 784776); Maravén, Borburata, E of
Puerto Cabello (A.M.).

Melampus (Detracia) morrisoni
new name
Figs. 355-376

Detracia clarki Morrison, 1951a: 18, figs. 2, 6
[Key West, Florida; holotype USNM
594588 (Fig. 355)]; Morrison, 1951b: 9;
Morrison, 1958: 118-124 [habitat]; Ab-
bott, 1974: 332; Emerson & Jacobson,
1976: 191, pl. 26, fig. 24 [dubious Шиз-
tration]; Vokes & Vokes, 1983: 60, pl. 22,
fig. 16. Non Melampus clarkii White,
1895.

Description: Shell (Figs. 355-367) to 17.7
mm long, ovate-conic to subcylindric, solid,
shiny; uniformly white to dark brown or with
as many as five spiral brown bands or with
irregular yellowish axial markings on body
whorl. Spire short to moderately high, with as
many as 13.25 spirally grooved or pitted
whorls. Body whorl about 80% of total
length, with incised spiral lines on shoulder.
Umbilical excavation sometimes present.
Aperture about 90% of body whorl length,
very narrow; inner lip with very strong, ob-
lique, upcurved columellar tooth; posterior
parietal tooth strong, sometimes upcurved,
anterior parietal tooth sometimes fused with
posterior one; outer lip with as many as 18
uneven internal riblets, not reaching edge;

when numerous, not more than five riblets
extend inside aperture. Partition of inner
whorls occupying about 75% of body whorl
(Fig. 361). Protoconch smooth, translucent,
brownish (Figs. 365-367).

Animal whitish, mottled with irregular
brown spots, or uniformly black becoming
lighter toward yellowish gray foot; tentacles
subcylindric, pointed, discolored at base,
dark toward tip; mantle skirt yellowish gray.

Radula (Figs. 368-372) having formula [29
+ (1 + 20) + 1 + (20 + 1) + 29] x 110. Base of
central tooth wider than that of lateral teeth,
triangular, weakly constricted laterally, with
small medial prominence on inner surface of
arms; crown length half of lateral teeth; me-
socone small, sharp; ectocones very small or
lacking. Lateral teeth 15 to 28; crown strong,
cuneiform, half total length of tooth; meso-
cone sharp, pointed laterally; no distinct ec-
tocone or endocone. Transitional tooth with
elongate crown, with either weak ectocone or
serrated edge at site of ectocone. Marginal
teeth 23 to 42, with reduced base; crown
high; mesocone very strong, sharp, rapidly
becoming round at tip; first marginal tooth
with ectocone, becoming bicuspid on sec-
ond marginal tooth and tricuspid on third;
fourth cusp appears on seventh or eighth
marginal tooth; endocone visible on fourth
marginal tooth; outer edge of base gradually
shortening posteriorly, fusing with crown and
assuming shape of denticle stronger than ec-
tocone cusps.

Digestive system as in Melampus $.5.;
stomach (Fig. 373) as in subfamily.

Reproductive system (Fig. 374) with
ovotestis leaf-like, circular, dark brown; albu-
men gland spiral, conical; prevaginal caecum
very conspicuous; bursa duct connecting
with vagina just before exit of posterior vas
deferens; bursa elongate; vagina thin, with
length corresponding to nearly one and one-
fifth the length of body whorl; penis thin,
slightly longer than associated vas deferens.

Nervous system (Fig. 375) with cerebral
commissure about as long as width of cere-
bral ganglion; left parietovisceral connective
slightly larger than to twice size of right one.

Remarks: Morrison (1951a) described this
species as Detracia clarki. The anatomy of
Detracia, however, does not justify generic
rank and this taxon is here considered a sub-
genus of Melampus (see the remarks for
Melampus s. |.). The name Melampus clarki
was used by White (1895) for a fossil shell,

MARTINS

298

FIGS. 355-367.

WESTERN ATLANTIC ELLOBIIDAE 299

À

FIGS. 368-371. Melampus (D.) morrisoni, radular teeth, Plantation Key, Florida. (368) Central and lateral
teeth, sl 12.9 mm. (369) Marginal teeth, sl 12.9 mm. (370) Central and lateral teeth of juvenile, sl 1.8 mm.
(371) Marginal teeth of juvenile, sl 1.8 mm. Scale 50 um.

later chosen as the type species of the genus
Melampoides Yen, 1951. The inclusion of De-
tracia as a subgenus of Melampus creates а
case of secondary homonymy. A new name
is necessary and | hereby rename the species
Melampus (Detracia) morrisoni, in honor of J.
P. E. Morrison and in appreciation for his
work on the Ellobiidae.

Melampus (D.) morrisoni is unusual among
members of this subgenus in having a con-
spicuous parietal tooth. In the other mem-
bers of the group this tooth is not readily
visible owing to its being deep within the ap-
erture. The strong columellar tooth, the
greater length of the pallial gonoducts, the
pouch-like mantle organ and the medial
nodes on the base of the central tooth of the
radula justify removing this species from

С 11 2 19L 1T 2T 1M 2M

FIG. 372. Melampus (D.) morrisoni, radula, Planta-
tion Key, Florida. Scale 10 um.

FIGS. 355-367. Melampus (D.) morrisoni, new name. (355) Detracia clarki Morrison, holotype (USNM
594588), Key West, Florida, sl 12.5 mm. (356) Grassy Key, Florida, sl 13.6 mm. (357) Grassy Key, Florida,
$1 14.0 mm. (358) Grassy Key, Florida, sl 17.7 mm. (359) Grassy Key, Florida, sl 15.7 mm. (360) Grassy Key,
Florida, 14.2 mm. (361) Plantation Key, Florida, sl 12.8 mm. (362) Key Largo, Florida, sl 12.7 mm. (363)
Juvenile, Long Key, Florida, sl 2.15 mm. (364) Millars Sound, New Providence, Bahamas, sl 13.8 mm.
(365-367) Lateral and top views of spire and protoconch, Long Key, Florida. Scale 500 um.

300 MARTINS

FIG. 373. Melampus (D.) morrisoni, stomach, Flor-
ida. Scale 1 mm.

FIG. 374. Melampus (D.) morrisoni, reproductive
system, Grassy Key, Florida. Scale 1 mm.

Melampus s. s. and placing it in Detracia,
however.

Melampus (D.) morrisoni can be confused
with Melampus (M.) bidentatus because they
converge in shape and color and are similar

in their variability. In its typical form, Melam-
pus (D.) morrisoni is readily distinguished by
its very narrow shell aperture, well-devel-
oped, upcurved columellar tooth, numerous
whorls and its more globose shape. Some
populations from inner lagoons, in which they
occur with Melampus (M.) bidentatus, show
gradation to an average-sized columellar
tooth and a common ovoid shape (Fig. 364).
In such cases anatomical studies are helpful.
The combination of a greater value of vagina
length/body whorl length and the slightly
higher number of whorls/shell length charac-
terizes Melampus (D.) morrisoni. The latter
species also can resemble the morphs of
Melampus (M.) coffeus that have a less pro-
nounced carina on the shoulder of the body
whorl. The strong, curved columellar tooth,
the presence of striae on the shoulder of the
body whorl, the narrow aperture and the un-
even outer lip riblets of Melampus (D.) morri-
soni clearly separate this species from
Melampus (M.) coffeus.

As do other members of the genus,
Melampus (D.) morrisoni undergoes a change
in radular morphology with age (Figs. 370,
371). The central tooth has weak but distinct
ectocones. The first lateral tooth, deeply tri-
cuspid in very young individuals, becomes
bicuspid, then unicuspid, with serrated
edges on the sites of the endocone or ecto-
cone, or both. The transitional tooth develops
an endocone, which remains through the
marginal teeth. The marginal teeth have more
ectocone cusps in the juvenile than in the
adult. The radula of juveniles very strongly
resembles in tooth count and morphology
that of the adult Melampus (D.) floridanus
(Figs. 324-327). This similarity suggests
some degree of neoteny in radular develop-
ment within the genus Melampus.

Habitat: Melampus (D.) morrisoni lives in as-
sociation with Melampus (M.) bidentatus and
Melampus (D.) bullaoides. It prefers sheltered
inland places in which the mangrove is thin
and reached only by very high tides. Individ-
uals aggregate under rocks and in old bur-
rows of fiddler crabs.

Range: South Florida [the St. Augustine
record is dubious, according to Morrison
(1951а)]; Bahama Islands south to Cuba and
Yucatán, Mexico (Fig. 376).

Specimens Examined: FLORIDA: St. Augus-
tine (USNM 492529a); Grant (MCZ 291233);
Miami (ANSP 189594, 294333; USNM

WESTERN ATLANTIC ELLOBIIDAE 301

prc PO P

1
\ \ |
i]

lprc plg

~ 90 75 60 _ 45 30

FIG. 376. Melampus (D.) morrisoni, geographic dis-
tribution.

82844a); Brickell’s Hammock, Miami (ANSP
294334); Virginia Key (ANSP 189591; USNM
82859a); Biscayne Bay (MCZ 291230); Flor-

cpc

ida City (ANSP 294335); Middle Key, Barnes
Sound (USNM 338339a); Pumpkin Key, Card
Sound (USNM 355114); Key Largo (MCZ
291232; USNM 603120); N of Tavernier
Creek, Key Largo (A.M.); 6 km S of Tavernier
(MCZ 291234); Tavernier Key (USNM
492552a); S of Ocean Drive, Plantation Key
(A.M.); Windley Key (USNM 603105); Indian
Key (USNM 462894); Indian Key Fill, М of In-
dian Key Channel (A.M.); Lower Matecumbe
Key (USNM 492554, 700774); Long Key
(A.M.); Grassy Key (MCZ 291236; А.М.);
Crawl Key (MCZ 291042); Marathon, Key
Vaca (ANSP 294336; MCZ 294264); Knight
Key (A.M.); Bahia Honda Key (ANSP 189576;
USNM 269777, 269980); Spanish Harbor Key
(USNM 667407); Newfound Harbor (USNM
272688, 338376); Little Pine Key (USNM
681638); Big Pine Key (ANSP 189583; MCZ
294270; USNM 104092a); end of Kohen
Avenue, Big Pine Key (A.M.); Howe Key
(USNM 681639); Big Torch Key (ANSP
189579; A.M.); Ramrod Key (MCZ 291043,
294263); Geiger's Key (ANSP 189582); Sug-
arloaf Key (ANSP 189577; USNM 104094a);
Porpoise Point, Big Coppit Key, 5 km N of
Key West (MCZ 275573); Boca Chica Key
(ANSP 189593; USNM 270329); Stock Island
(ANSP 189581; USNM 594589); Key West
(ANSP 180089, 189578; MCZ 291235;

302 MARTINS

USNM 36062, 668245); Chokoloskee Key
(ANSP 93430). BAHAMA ISLANDS: GRAND
BAHAMA ISLAND (ANSP 374526): North
Riding Point (ANSP 371539); GREAT ABACO
ISLAND: Witch Point (ANSP 359152; USNM
594592); Angel Fish Point (MCZ 294265);
NEW PROVIDENCE ISLAND: Millars Sound
by Bacardi Road (A.M). CUBA: Cape Cajón
(USNM 492571); Cayo Perro (ANSP 189575;
USNM 594590). Cayo Juan Garcia (MCZ
291231); Cayo Maja, near Cayo Santa Maria
(MCZ 294217). MEXICO: Isla Mujeres, Quin-
tana Roo, Yucatán (R.B.).

Genus Tralia Gray, 1840

Tralia Gray, 1840: 21. Type species by mono-
typy: Tralia pusilla (Gmelin, 1791) [= Vo-
luta ovula (Bruguiere, 1789)].

Tralica Gray. Reeve, 1877, pl. 1 [in synonymy
of Auricula; error for Tralia].

Description: Shell thick, oval-elongate; aper-
ture moderately long, widest part anterior to
columellar tooth; inner lip with three white
teeth, first parietal strongest; outer lip thick-
ened, slightly reflected, with distinct anal
groove.

Animal white; tentacles flat dorso-ventrally.
Internal edge of arms of central radular tooth
with prominent medial nodes; first lateral
tooth with conspicuous endocone; transition
to marginal teeth marked. Salivary glands at-
tached lateroventrally to esophagus; esoph-
agus white. Ovotestis granular; mucous gland
very convoluted, not clearly spiral; bursa duct
connects at some distance from proximal end
of vagina; penis very long, muscular.

Remarks: The genus Tralia was created by
Gray (1840) for the West Indian Tralia pusilla
(Gmelin) [= Tralia ovula (Bruguiere)] on the ba-
sis of the peculiar, simple outer lip with its
distinct anal groove. H. & A. Adams (1855b:
244) separated Tralia Gray from Melampus
Montfort on the basis of the incorrect obser-
vation, “the foot was posteriorly acute, en-
tire.” The Adams brothers were confused,
however, for they commented, “This group,
which appears to have a simple, undivided
tail, .. . perhaps, when the animals are better
known, will be found to be merely a subge-
nus of Melampus.”

H. & A. Adams recognized four subgenera
of Tralia, Pira, Tifata, Signia and Persa. | have
commented on Pira and Tifata under the re-
marks for Detracia Gray. Signia, the third

subgenus of Tralia introduced by H. 8 A. Ad-
ams (1855b), was later treated as a subgenus
of Melampus by Thiele (1931) and Zilch
(1959). The fourth subgenus, the Pacific
Persa, was recognized by Thiele and Zilch as
belonging to the genus Tralia; it is character-
ized by the very short spire and by the con-
vex, distinctly ribbed whorls. Another subge-
nus also placed in Tralia was introduced by
the Adams brothers in 1855 as Siona, a ge-
nus of Cassidula, just prior to the publication
of their Genera of Recent Mollusca. In this
latter publication they changed the name to
Sarnia without specifying the reason. Thiele
(1931) recognized Siona as belonging to Tra-
lia; Zilch (1959) noted that the name Siona
was preoccupied and that Sarnia should be
the correct name. The type of the subgenus,
Tralia (Sarnia) frumentum (Petit, 1842), from
South America, is an Eastern Pacific species
conchologically very similar to the eastern
North Atlantic Pseudomelampus exiguus
(Lowe, 1832) [Pedipedinae] and to the genus
Microtralia (Figs. 174-181). Keen (1971) con-
sidered Pseudomelampus to be synonymous
with Sarnia and placed it within the Ellobii-
nae. According to Marincovich (1973) the
radula of Tralia (Sarnia) frumentum is similar
to that of Ellobium, and he considered Sarnia
to belong in the Ellobiinae. It appears, then,
that Sarnia cannot be considered a subgenus
of Tralia. | have considered Sarnia, on the
basis of the conchological resemblances
with Pseudomelampus, to belong in the Pe-
dipedinae (see the remarks under the Ellobi-
idae). A study of the reproductive and ner-
vous systems is needed to ascertain its
phylogenetic position, however.

Binney (1865: 16, fig. 16) figured the animal
of an alleged Tralia, but admitted in a foot-
note that he did not know which species it
represented. Simpson drew from nature a
figure of an animal from Charleston, South
Carolina, a locality that is not in the range of
Tralia (T.) ovula. | concur with Dall (1885) that
Simpson's drawing probably represents My-
osotella myosotis (Draparnaud).

The genus Tralia is readily distinguished
from Melampus by its wide anterior aperture,
its strong first parietal tooth and its thickened
outer lip with a subposterior internal groove.

The apertural dentition of Tralia is unusual
for a member of the Melampinae and, after
anatomical research is carried out, it is pos-
sible that some Indo-Pacific species pres-
ently put in this subgenus will be found to
belong to other subfamilies.

WESTERN ATLANTIC ELLOBIIDAE 303

Habitat: The only information available about
the habitat of the genus refers to the West
Indian Tralia (T.) ovula and, for this reason,
comments on habitat will be made under that
species.

Range: Tralia 1$ a tropical group, living
mostly in the Indo-Pacific. It is represented in
the West Indies by one species, Tralia (T.)
ovula, which seems to have been introduced
in West Africa.

Subgenus Tralia $. $.

Description: Shell to 16 mm long; spire mod-
erately high, with pitted lines; body whorl
smooth or weakly marked with punctate spi-
ral lines.

Remarks: see the remarks under Tralia $. I.

Tralia (Tralia) ovula (Bruguiere, 1789)
Figs. 377-387, 389-400

Bulimus ovulus Bruguiere, 1789: 339 [Guade-
loupe, West Indies; location of type un-
known]; Cuvier, 1817: 414.

Voluta pusilla Gmelin, 1791: 3436 [locality un-
known, herein designated to be Guade-
loupe, West Indies; location of type un-
known]; Dillwyn, 1817: 507; Wood, 1825:
91, pl. 19, fig. 20; Hanley, 1856: 98, pl.
19 Mg. 20:

Voluta triplicata Donovan, 1802, pl. 138 [lo-
cality unknown, herein designated to be
Guadeloupe, West Indies; location of
type unknown]; Montagu, 1808: 99; Dill-
wyn, 1817: 507; Wood, 1825: 91, pl. 19,
fig. 19; Hanley, 1856: 98, pl. 19, fig. 19.

Auricula (Conovulus) ovula (Bruguiere). Fér-
ussac, 1821: 104; Rang, 1829: 173.

Auricula nitens Lamarck, 1822: 141 [Guade-
loupe, West Indies; type in the MHNG
(Mermod, 1952)]; Menke, 1830: 36;
Gould, 1833: 67; Jay, 1839: 59; Küster,
1844: 18, pl. 2, figs. 11-13.

Melampus ovulum Schweigger. Lowe, 1832:
289.

Pythia triplicata (Donovan). Beck, 1837: 104.

Pythia ovulum (Bruguiere). Beck, 1837: 104.

Auricula ovula (Bruguiere). Potiez & Michaud,
1838: 204, pl. 20, figs. 13, 14.

Auricula pusilla (Gmelin). Deshayes. 1838:
332.

Auricula (Conovulus) pusillus Deshayes. An-
ton, 1839: 48.

Tralia pusilla Gray. Gray, 1840: 21.

Auricula ovula Férussac. Orbigny, 1841: 186,
pl. 13, figs. 1-3.

Tralia pusilla (Gmelin). Gray, 1847a: 179; H. &
A. Adams, 1855b: 244, pl. 82, fig. 8; Bin-
ney, 1865: 17, fig. 18; Tryon, 1866: 9, pl.
18, fig. 9; Dohrn, 1866: 133 [first record
from Eastern Atlantic]; Dall, 1885: 276,
pl. 18, fig. 5; Dall, 1889: 92, pl. 47, fig. 5;
Dall in Simpson, 1889: 69; Dall & Simp-
son, 1901: 369, pl. 59, fig. 13; Odhner,
1925: 5, pl. 1, fig. 8B, pl. 2, fig. 18 [radula
and reproductive system figured]; Peile,
1926: 88; C. W. Johnson, 1934; 159;
Coomans, 1958: 103, pl. 10; Franc,
1968: 525.

Melampus pusillus (Gmelin). C. B. Adams,
1849: 42; C. B. Adams, 1851: 186;
Pfeiffer, 1854b: 147: Pfeiffer, 1856a: 46
[erroneously stated as also inhabiting
Hawaii]; Binney, 1859: 168, pl. 75, fig.
29; Binney, 1860: 4; Poey, 1866: 394;
Jeffreys, 1869: 109; Pease, 1869: 61;
Pfeiffer, 1876: 317; Arango y Molina,
1880: 59; Crosse, 1890: 258.

Melampus (Tralia) pusillus (Gmelin). H. 4 A.
Adams, 1854: 10.

Melampus nitens (Lamarck). Shuttleworth,
1854b: 101; Shuttleworth, 1858: 73;
Morch, 1878: 5; Nevill, 1879: 219.

Tralia pusilla Linnaeus. Fischer & Crosse,
1880: 22.

Tralia (Tralia) pusilla (Gmelin). Thiele, 1931:
466.

Tralia ovula (Bruguiere). Morrison, 1951b: 9;
Nowell-Usticke, 1959: 88; Coomans,
1969: 82; Warmke & Abbott, 1961: 153,
pl. 28, fig. m; Morris, 1973: 274, pl. 74,
fig. 10; Abbott, 1974: 332, fig. 4095;
Emerson & Jacobson, 1976: 193, pl. 26,
fig. 29; Gibson-Smith & Gibson-Smith,
1982: 117; Vokes & Vokes, 1983: 60, pl.
22, fig. 17; Mahieu, 1984, 314 pp.

Tralia (Tralia) ovula (Bruguiére). Zilch, 1959:
67, fig. 215.

Tralia ovula sculpta Nowell-Usticke, 1959: 88
[St. Croix-by-the-Sea, Cane Bay, St.
Croix; lectotype herein selected AMNH
220313 (Fig. 384); listed on page VI as
Tralia ovulata sculpta].

Tralia cf. ovula (Bruguiere). Gibson-Smith &
Gibson-Smith, 1979: 22 [Cantaure For-
mation, Venezuela (Miocene)].

Tralia venezuelana Gibson-Smith 4 Gibson-
Smith, 1982, figs. 7-9 [Borburata, Falcón
State, Venezuela; holotype USNM
784719 (Fig. 377).

304 MARTINS

Description: Shell (Figs. 377-387, 389-391)
to 16 mm long, oval-elongate, solid, shiny,
uniformely chestnut brown to dark purplish
brown, sometimes with one or two paler
bands on body whorl. Umbilicus present.
Spire low to moderately high, with as many
as nine flat whorls sculptured with four or five
spiral rows of deep pits. Body whorl averag-
ing 85% of shell length, oval-elongate, with
striated, uncarinate shoulder; striations visi-
ble over entire body whorl of most young,
commonly only on anterior region in adults.
Aperture averaging 85% length of body
whorl, posteriorly angulate, widely rounded
anteriorly, white to dark purple inside; inner
lip with three evenly spaced, large white
teeth; columellar tooth and posterior parietal
tooth of same size, reversely oblique, col-
umellar tooth inclined toward base of aper-
ture; first parietal tooth strongest, perpendic-
ular to columellar axis; excavation posterior
to second parietal tooth, bordered outside by
more or less conspicuous callus, continuing
inwards, commonly with small irregularities,
sometimes with prominent denticle; outer lip
sharp in juveniles, thick and weakly reflected
in gerontic individuals, weakly sinuous pos-
teriorly and with thick callous denticle inside,
opposite second parietal tooth; outer lip den-
ticle ridge-like, continuing inside aperture, to-
gether with second parietal tooth delimiting
relatively wide canal. Inner partition of whorls
occupying less than half of the body whorl
(Fig. 379). Protoconch smooth, yellowish to
brown, with nucleus visible (Figs. 389-391).

Animal white; foot with transverse groove,
whitish, with minute brown spots over bifid
posterior end; tentacles dorsoventrally flat-
tened, spatulate, with first quarter bulbous,
white, abruptly changing to dark grey or
black toward tip; seminal groove unpig-
mented; mantle skirt with very small brown
spots over light brown background. Kidney

rectangular, elongate; mantle organ well de-
veloped, not pouch-like.

Radula (Figs. 392-396) having formula [38 +
(1 + 15) + 1 + (15 + 1) + 38] x 115. Base of
central tooth with conspicuous medial prom-
inences on inner edge of arms; crown length
about half of that of lateral teeth, broadly tri-
angular anteriorly, elongate posteriorly; ecto-
cones absent. Lateral teeth 13 to 19; crown
broadly triangular; conspicuous endocone on
first lateral tooth; posterior medial portion of
base of remaining lateral teeth flaring at junc-
ture with crown, simulating endocone; last lat-
eral tooth sometimes with very weak ecto-
cone. Transitional tooth with lateral portion of
crown posteriorly elongate, with tricuspid ec-
tocone; base almost straight. Marginal teeth
35 to 43; crown very elongate and irregularly
pointed posteriorly; mesocone sharp, long,
becoming rounded, spatulate, almost as long
as remaining denticles, but much stronger;
first marginal tooth with conspicuous en-
docone and a tricuspid ectocone; as many as
nine ectocone cusps on last ten marginal
teeth.

Digestive system with salivary glands at-
tached close together by fine thread on ven-
tral side of white esophagus; posterior crop
very dilated, forming pouch before entering
stomach; stomach (Fig. 397) as in subfamily;
digestive gland pale yellow to bright orange.

Reproductive system (Fig. 398) having
ovotestis dark brown with whitish spots, gran-
ular; mucous gland and albumen gland inter-
penetrating at base; spermoviduct cylindrical,
very muscular; bursa duct connecting at a
point about one-fourth of total length from
proximal end of long, muscular vagina; pos-
terior vas deferens about 80% of vagina
length; penis long, very muscular, with pos-
terior half sometimes wrapped in membra-
nous sheath; anterior vas deferens of variable
length.

FIGS. 377-391. Tralia. (377) T. venezuelana Gibson-Smith & Gibson-Smith, holotype (USNM 784719),
Borburata, Falcón State, Venezuela, sl 12.7 mm. (378) Т. (T.) ovula (Bruguiere), El Palito, Venezuela, sl 13.6
mm. (379) Т. (T.) ovula, El Palito, Venezuela, sl 13.5 mm. (380) 7. (T.) ovula, San Juan, Puerto Rico, sl 12.8
mm. (381) 7. (T.) ovula, Rock Pt., New Providence, Bahamas, sl 14.9 mm. (382) 7. (T.) ovula, Ilha do Principe,
Gulf of Guinea (MCZ 73375), sl 9.0 mm. (383) “Voluta triplicata Donovan,” West Indies (USNM 442093),
from Turton's Cabinet, Jeffreys collection, sl 14.8 mm. (384) Т. ovula sculpta Nowell-Usticke, lectotype
(AMNH 220313), St.-Croix-by-the-Sea, Cane Bay, St. Croix, sl 12.0 mm. (385) 7. (T.) ovula, Haiti (MCZ
18392), sl 10.4 mm. (386) 7. (T.) ovula, Robin's Bay, Jamaica (USNM 712378), sl 6.4 mm. (387) Т. (T.) ovula,
juvenile, Maravén, Venezuela, sl 2.3 mm. (388) Т. vetula Woodring, holotype (ANSP 12506), Bowden,
Jamaica, sl 5.5 mm. (389) Т. (T.) ovula, lateral view of spire and protoconch, Tucacas, Venezuela. (390) 7.
(T.) ovula, lateral view of spire and protoconch, Rock Pt., New Providence, Bahamas. (391) 7. (Т.) ovula, top
view of spire and protoconch, Haiti (USNM 439659). Scale 1 mm.

WESTERN ATLANTIC ELLOBIIDAE 305

RENEE 6
A ena =

FIGS. 377-391.

306 MARTINS

FIGS. 392-395. Tralia (T.) ovula, radular teeth. (392-394) El Palito, Venezuela, sl 14.7 mm. (395) Central
tooth and adjacent lateral teeth, with articulation between base of one tooth and crown of next tooth, Bar
Pt., New Providence, Bahamas, sl 14.1 mm. Scale 50 um.

10M 11M 12M 20M 21M 22M 30M31M32M

ADO AE

FIG. 396. Tralia (T.) ovula, radula, El Palito, Vene-
zuela. Scale 10 um.

Nervous system (Fig. 399) with cerebral
commissure as long as width of cerebral
ganglion; left parietovisceral connective two
to three times longer than right one; left FIG. 397. Tralia (T.) ovula, stomach, Venezuela.
parietal ganglion half size of right one; vis- Scale 1 mm.

WESTERN ATLANTIC ELLOBIIDAE 307

FIG. 398. Tralia (T.) ovula, reproductive system. A, Clifton Pt., New Providence, Bahamas; B, Puerto Rico,
$1 13.8 mm; С, Tucacas, Venezuela, sl 10.9 mm; D, El Palito, Venezuela, sl 13.8 mm. Scale 1 mm.

308 MARTINS

spc pc pg pe ple coo 0 bg

FIG. 399. Tralia (T.) ovula, central nervous system,
San Juan, Puerto Rico, sl 13.8 mm. Scale 1 mm.

ceral ganglion as large as right parietal gan-
glion.

Remarks: The names Bulimus ovulus Bru-
guiére, 1789, and Voluta pusilla Gmelin,
1791, were thought to have appeared in the
same year (Dall, 1885) and early authors were
more inclined to use the latter. In fact Bru-
guiére's name antedates Gmelin’s and it has
now been accepted as the correct name by
most authors.

Gmelin’s description, although mentioning
the tridentate columella, is brief and omits the
geographic origin of the specimen. The fact
that both Gmelin and Bruguiere referred to
Martini (1773, fig. 446) indicates that these
authors were describing the same species.

Donovan (1802) introduced Voluta tripli-
cata (Fig. 383) based on material of unknown
origin, although Montagu later stated (1808)
that the specimens were from Guernsey, En-
gland. As in the case of Melampus (D.) bul-
laoides (Montagu) and of its junior synonym
Auricula multivolvis Jeffreys, Donovan’s ma-
terial might have reached England in the bal-
last of ships coming from the West Indies.
Auricula (Conovulus) triplicatus Anton (1839)
should not be confused with Donovan’s spe-
cies. Anton referred to the highest (posterior)
parietal tooth as the strongest, a character-
istic of most Pedipedinae. Connolly (1915)
considered Anton’s species a junior synonym
of Marinula pepita King, 1832.

Bruguière (1789) and Dillwyn (1817) both
mentioned Martini’s (1773) reference to fine,
axial striations on the shell; this can refer only
to the very fine growth lines that are some-
times visible. The shell can be sculptured

with well-marked spiral striae, however (Figs.
387, 389-391). It was on the basis of the spi-
ral striations that Nowell-Usticke (1959: 88)
described Tralia ovula sculpta in two words,
“spire lined” (Fig. 384). Shuttleworth (1858)
was the first to mention the five deeply pitted
spiral lines on the early whorls, erroneously
adding that they were “ciliated” in juveniles.
Juveniles and some adults of Melampus (D.)
monile, which at a first glance can be con-
fused with Tralia (T.) ovula, have a crown of
hairs on the spire (Fig. 341), but neither the
adults nor the young of Tralia (T.) ovula have
hairs.

Gibson-Smith & Gibson-Smith (1982) de-
scribed Tralia venezuelana from Borburata,
Venezuela (Fig. 377), which they distin-
guished from Tralia (T.) ovula on the basis of
its pitted spire and the presence of a fourth
denticle in the aperture. | have observed a
pitted spire, more or less pronounced, on all
well-preserved specimens and on all young
shells of Tralia (T.) ovula. It is particularly
marked in some thin-shelled, elongate, dwarf
morphs from Cuba, Jamaica, Haiti and St.
Croix. The fourth denticle, in the shallow pos-
terior parietal excavation, also occurs in
specimens from Cuba, Jamaica, Haiti, Puerto
Rico and St. Croix. Specimens from the Ba-
hamas have an irregular surface on the pos-
terior portion of the inner lip, but no distinct
denticle. The fourth denticle seems to be a
variable character, not associated with differ-
ences in radula, anatomy, or shell. Presence
of extra parietal teeth has been reported in
the Pacific Melampus fasciatus, Melampus
luteus and Melampus nucleolus and it is not
considered a reliable taxonomic character for
those species (Jickeli, 1872). Extra denticles
also occur in Melampus (M.) coffeus and
Melampus (M.) bidentatus (Martins, personal
observation). | therefore have placed Tralia
venezuelana Gibson-Smith & Gibson-Smith
in the synonymy of Tralia (T.) ovula.

In some Venezuelan specimens the poste-
rior half of the very muscular penis was
folded and wrapped in a membranous sheath
(Fig. 398D). Such specimens also had an un-
usually muscular spermoviduct and a shorter
anterior vas deferens. Except for larger size
(average 14 vs. 12 mm) no other shell and
radular characters were associated with the
phenomena. They were not associated with
the presence of a fourth denticle on the ap-
erture. It is possible that such phenomena
are anomalies of gerontic specimens of that
population.

WESTERN ATLANTIC ELLOBIIDAE 309

There are two reports of fossil West Indian
Tralia. Woodring (1928) described Tralia (T.)
vetula from the Pliocene Bowden Formation
of Jamaica (Fig. 388). After examining several
lots of Recent material from Jamaica, | com-
pared Woodring's example with the thinner-
shelled, slender, dwarf specimens of Tralia
referred to above; | found that these recent
specimens show all gradations of thickness.
Tralia (T.) vetula 1$ considered a distinct spe-
cies on the basis of the less pronounced den-
tition of the inner lip, however. Gibson-Smith
& Gibson-Smith (1979) reported a Tralia ?
ovula from the Early Miocene Cantaure For-
mation of Paraguaná, Venezuela, and also
from the Late Pliocene Mare Formation of
Cabo Blanco, Venezuela (Gibson-Smith &
Gibson-Smith, 1982). In the latter publication
the Gibson-Smiths identified those speci-
mens with their Tralia venezuelana, which |
consider synonymous with Tralia (T.) ovula.

Habitat: Tralia (T.) ovula lives along the high-
tide mark. The animals prefer piles of boul-
ders on open rocky shores, but they also live
in the less-protected mangroves.

Range: Bermuda; Florida Keys; West Indies
to Trinidad; Central America to Venezuela;
IIha do Principe, Gulf of Guinea, Africa (Fig.
400).

Specimens Examined: FLORIDA (USNM
37597, 39873): Tavernier Key (USNM
492519); Lower Matecumbe Key (USNM
492595); Long Key (A.M.). BERMUDA (MCZ
304151; USNM 6531, 94434). BAHAMA IS-
LANDS (USNM 492465): GRAND BAHAMA
ISLAND (ANSP 173482, 375528): Eight Mile
Rock (MCZ 294268); GREAT ABACO IS-
LAND: Little Harbor (USNM 180486); AN-
DROS ISLAND (MCZ 58507, 66744, 66756);
NEW PROVIDENCE ISLAND: Nassau (USNM
534924); Rock Point (A.M.); Clifton Point
(A.M.); RUM CAY (MCZ 304157); LONG IS-
LAND (ANSP 173483): 3 km NE of O’Neill’s
(MCZ 304152); GREAT INAGUA ISLAND:
Matthewstown (MCZ 304153). TURK’S & CA-
ICOS: TURK’S ISLAND (MCZ 304150; USNM
492469, 509960а). CUBA (ANSP 56807;
USNM 10965, 492472): Habana (MCZ
294794); Cayo Birricu (ANSP 362825); Jai-
manitas (MCZ 294199); Matanzas (ANSP
167243; MCZ 304168); La Playa (MCZ
304159, 304165); Versalles (MCZ 304154);
Varadero (MCZ 304164); Caibarién (MCZ

FIG. 400. Tralia (T.) ovula, geographic distribution.

304163, 304175); Cayo Francés (MCZ
294200); Siboney (USNM 533913); Punta de
Piedras (MCZ 304166); Bahia de Santiago
(MCZ 304161); Cabo Cruz (MCZ 304156).
JAMAICA (ANSP 66964; MCZ 294269,
304158, 304167, 304169, 304174; USNM
49744a, 94745, 492467, 492471, 492593):
Montego Bay (ANSP 359145); Robin’s Bay
(USNM 442092a, 442093); Jack’s Bay
(USNM 441834); Port Maria (USNM 711209);
Buff Bay (USNM 441195); Stoney Cove
(USNM 440762); Port Antonio (ANSP 62022;
USNM 712147); Priestman’s River (USNM
492468); Manchioneal Bay (ANSP 61883;
MCZ 9950; USNM 127359; R.B.); Port Royal
(USNM 395452a, 442419a); Runaway Bay
(USNM 202657); Little River (USNM 128047,
492463). HAITI: Yuma River (ANSP 60950);
St. Louis (USNM 439390); St. Marc (USNM
492470); Port Salut (ANSP 226694; MCZ
183922; USNM 440024); lle-a-Vache (USNM
401874, 401875, 439169); Les Cayes (USNM
439746); Torbeck (USNM 402261, 439659);
Aquin (USNM 403256a, 440170a); Baie Ang-
laise, near Aquin (USNM 439548); between
Vieux Bourg and Bale des Flamands (USNM
403425); N of Metesignix (USNM 404149);
Saltrou (ANSP 387078; USNM 439342а,
442813); Bizoton (USNM 439828a). DOMIN-

310 MARTINS

ICAN REPUBLIC (MCZ 304172; USNM
151297): Santo Domingo (ANSP 62910);
Monte Cristi (MCZ 304162); Samaná (MCZ
281639). PUERTO RICO: Puerta de Tierra,
San Juan (A.M.); Rifle Range Beach, Punta
Agurejeada (MCZ 233299); Arecibo (MCZ
304160); Cabo Rojo Lighthouse (МСА
294202); Ensenada Honda, Culebra Island
(USNM 169886). VIRGIN ISLANDS: ST.
CROIX (ANSP 56806; MCZ 200448, 304155;
USNM 621395): Prosperity Beach (MCZ
304173); St. Croix-by-the-Sea, Cane Bay
(AMNH 192356, 220313; ANSP 231952); ST.
THOMAS (ANSP 56805, 359147; USNM
250034, 530175): Sapphire Beach (ANSP
306673); Water Bay (ANSP 56808); ST.
JOHN'S (MCZ 304171); GUANA ISLAND
(MCZ 294203): North Bay (ANSP 351790).
LESSER ANTILLES: ANGUILLA ВАМК$ (MCZ
294201); ANTIGUA (ANSP 109155; USNM
215048): off Falmouth (USNM 502098);
GUADELOUPE (USNM 492466, 492518):
Anse-Dumont, Gosier (USNM 758066); BAR-
BADOS: (MCZ 304170, 304177, 304178;
USNM 502104, 502105); Bridgetown (USNM
502102); Needham Point (USNM 502101);
Maxwell’s Coast (USNM 603783); Pelican Is-
land (USNM 502100); GRENADA: Prickly Bay
(ANSP 297184); TRINIDAD: South Coast
(ANSP 363992). CARIBBEAN ISLANDS:
SWAN ISLAND (MCZ 36612, 294267); CAY-
MAN ISLANDS: Little Cayman (MCZ 294204);
ST. ANDREWS ISLAND (ANSP 159360); CU-
ВАСАО: Port Marie and Daaibooi Baai (R.B.).
MEXICO: Ascension Bay, Quintana Roo
(USNM 736380). BELIZE: Belize (USNM
151050). HONDURAS: Roatan Island (USNM
364701а). COSTA НСА: Portete, Limön
(USNM 702826, 706404). PANAMA: Fort
Sherman, Devil's Beach (USNM 620530);
Toro Point, Fort Sherman (USNM 732868,
734071). COLOMBIA: Sabanilla (USNM
193612). VENEZUELA: Cayo Punta Brava,
Parque Nacional de Morrocoy, Tucacas
(A.M.); El Palito (A.M.); Borburata (USNM
784719, 784772); Maravén, Borburata (A.M.).
EASTERN ATLANTIC: Ilha do Principe, Gulf of
Guinea (MCZ 73375).

CONCLUSIONS

Phylogeny and Classification

Gastropods have long been divided into
prosobranchs, opisthobranchs and pulmo-
nates. Ihering’s (1876, 1877) anatomical re-
search led him to conclude that the Gas-

tropoda were polyphyletic. He derived the
prosobranchs from the annelids, and the
opisthobranchs and the pulmonates from
the platyhelminths. Ihering’s view has not
been accepted and there is consensus that
the gastropods are in fact monophyletic. A
difference of opinion arises, however, about
the way in which the generally more ad-
vanced euthyneurans (opisthobranchs and
pulmonates) are related to the more primitive
streptoneurans (prosobranchs). Pelseneer
(1894a) and Hubendick (1945) thought that
the euthyneurans arose from the archaeo-
gastropods. Pelseneer based his decision
upon the similarities of the rhipidoglossan
radula of the trochids with that of the ceph-
alaspideans and basommatophorans. Mor-
ton (1955c) slightly modified this view by
proposing a pre-archaeogastropod as the
ancestor of the archaeogastropods and eu-
thyneurans. A different view was held by
Fretter (1946, 1975), Boettger (1954) and
Gosliner (1981), who considered the meso-
gastropod stock as the ancestor of the eu-
thyneurans. For the first two authors this
origin would be located near the Rissoacea,
on the basis of the size and habitat of the
Recent species of that superfamily, which
live in marine, estuarine, freshwater and ter-
restrial habitats. Their small size could ex-
plain the loss of the ctenidium, and the inva-
sion of the terrestrial habitat would favor the
secondary development of an air-breathing
pallial cavity. In addition, the mesogastropod
lineage explains the absence of the right kid-
ney throughout the Euthyneura and the sim-
ilarity of the reproductive tract with that of a
female mesogastropod (Fretter, 1975). Gos-
liner (1981), however, on the basis of the re-
productive system, together with the fossil
record, considered the Littorinacea as the
stock that produced the euthyneurans.

More recently emphasis has been put on
the Pyramidelloidea Gray, 1840, a зирейат-
ily of mostly small streptoneurans assembled
within the suborder Allogastropoda Haszpru-
nar, 1985. Separation of this superfamily
from the Opisthobranchia has favored the hy-
pothesis of a common ancestry for the Pyra-
midelloidea and euthyneurans (Haszprunar,
1985, 1988; Salvini-Plawén 8 Haszprunar,
1987).

Traditionally the ellobiids have been con-
sidered the living representatives of the prim-
itive pulmonates and simultaneously they
have been associated with the primitive
opisthobranchs. Mórch (1865: 11) was the

WESTERN ATLANTIC ELLOBIIDAE 311

first to recognize the affinities of pulmonates
and opisthobranchs when, on the basis of
their hermaphroditism, he included both
groups within his subclass Androgyna.
Pelseneer (1894a) broadened that relation-
ship by also calling attention to their detorted
(euthyneurous) nervous system. This euthy-
neurous condition led some authors to con-
sider both groups members of the subclass
Euthyneura Spengler, 1881 (Boettger, 1954;
Taylor & Sohl, 1962; Burch, 1962; Haszpru-
nar, 1985). Others, while recognizing the
close similarities between the two groups,
still prefer the traditional terminology, consid-
ering the Pulmonata and the Opisthobran-
chia as separate subclasses (Morton, 1955с;
Fretter & Graham, 1962; Robertson, 1973;
Fretter, 1975; Hubendick, 1978; Salvini-Pla-
wén, 1980).

Pelseneer (1893, 1894a), Thiele (1935) and
Boettger (1954) derived the pulmonates from
the cephalaspidean Acteon and considered
the ellobiids to be the link between the two
groups. Pelseneer found support for his de-
cision in the earlier fossil record of the
acteonids and on the similarities of the shell
apertures of the acteonids and ellobiids.
Morton (1955c) suggested that these con-
chological characters might be adaptive only
and, as such, their taxonomic value 1$ not
significant. As Harry (1951) pointed out, the
heterostrophy common to the ellobiids and
the opisthobranchs constitutes a much
stronger taxonomic character. More com-
mon is the opinion that the opisthobranchs
and pulmonates arose from the same proso-
branch stock, not one from the other (Mor-
ton, 1955c; Fretter, 1975; Gosliner, 1981).

Within the pulmonates, the relationship
between the basommatophorans and the
stylommatophorans also has been the object
of several hypotheses. Apart from the direct
line “opisthobranch-ellobiid (basommatopho-
ran)-stylommatophoran” scheme advocated
by Pelseneer (1893, 1894a), Hedley (1917)
and Boettger (1954), the more commonly
held opinion is that the basommatophorans
are too diverse to be considered collectively
as ancestors to the stylommatophorans.
Burch (1962), based on the number of chro-
mosomes, proposed that a hypothetical pre-
basommatophoran (called Ur-Basommato-
phora) with opisthobranchiate ancestry might
have given rise to Morton's (1955c) Archae-
opulmonata and Branchiopulmonata. The
first group includes the Ellobiidae, Amphibol-
idae and Siphonariidae, commonly consid-

ered the “lower basommatophorans.” The
second group includes the freshwater pul-
monates, or “higher limnic basommatopho-
rans.” According to this view the Stylom-
matophora are polyphyletic, although having
all originated from the Archaeopulmonata.
Harry (1964) slightly modified Burch’s phyl-
etic tree by adding a “pre-pulmonate” an-
cestor to the Urbasommatophora [sic]. A
similar framework of classification, also fol-
lowed here, was adopted by Van Mol (1967),
Hubendick (1978), Salvini-Plawén (1980),
Boss (1982) and Tillier (1984), who restricted
the term Basommatophora to Burch’s
“higher limnic basommatophorans.”’

Quite a different view was proposed by
Starobogatov (1976), mostly on the basis of
the reproductive system. He reversed the di-
rection of the evolutionary relationships
within the pulmonates and considered the
basommatophorans as derived from the sty-
lommatophoran stock. Starobogatov did not
consider the widespread fusion of the ganglia
in the stylommatophorans, a condition that 1$
considered derived. Reversal to a more dis-
persed condition of the ganglia is very un-
likely and has not been reported for other
groups.

Phylogenetic relationships within the Ello-
biidae, although obscured by the presence in
each subfamily of primitive and derived char-
acters, can be elucidated by comparing the
reproductive and nervous systems. Diauly,
nonglandular condition of the pallial gono-
ducts and ganglionic concentration on the
nervous system are derived conditions (Gos-
liner, 1981; Haszprunar, 1985, 1988; Salvini-
Plawén 8 Haszprunar, 1987); | consider these
features decisive in the interpretation of the
degree of departure from the ancestral plan.
A tentative phylogenetic tree elaborated on
that basis is presented in Figure 401 and Ta-
bles 5 and 6 (Appendix).

Consistent patterns of organization al-
lowed a clear delimitation of subfamilial
boundaries. Five types of organization of the
reproductive system were identified, corre-
sponding to Morton's (1955c) subfamilial di-
visions, for which they are named. First is the
Pythiinian type, monaulic, with the anterior
mucous gland and prostate gland running
parallel to each other and covering the pallial
gonoduct as far as the vaginal atrium. Sec-
ond is the Ellobiinian type, diaulic, with the
anterior mucous gland and prostate gland
covering the pallial gonoducts for all their
length. Third is the Carychiinian type, monau-

312 MARTINS

Ringicula
Pythia
Myosotella

Ovatella
Laemodonta
assidula
Pedipes
Creedonia
Cremnobates?

Microtralia

OF Of Lee Mierotralia
Pseudomelampus
(2) LY Marinula
ome ES

Auriculodes
Ellobium
Auriculinella

FIG. 401.

WESTERN ATLANTIC ELLOBIIDAE 313

lic, with the pallial gonoduct glandular and
the prostate gland concentrated distally in
the gonoduct. Giusti (1975), in a sketchy rep-
resentation of the reproductive system of
Zospeum, left doubt whether the system is
monaulic or diaulic, but the glandular appear-
ance resembles the pythiinian or ellobiinian
types; however, dissections of Carychium cf.
tridentatum from the Azores using superficial
staining with methylene blue (Martins, per-
sonal observation) indicated an agglomera-
tion of what appeared to be a prostate gland
near the distal portion of the pallial gonoduct,
thus substantiating Morton’s (19550) inter-
pretation. Fourth is the Pedipedinian type,
monaulic ог incipient semidiaulic, with the
anterior mucous gland and the prostate
gland covering only the proximal half of the
pallial gonoduct. Fifth 1$ the Melampinian
type, advanced semidiaulic, with a very
short, nonglandular spermoviduct and sepa-
rated, nonglandular, long vagina and poste-
rior vas deferens.

Although not so discrete as in the repro-
ductive system, patterns were found in the
organization of the central nervous system,
concerning the relative lengths of the various
connectives. Three types were identified. The
Pythiinian type has a wide visceral nerve ring
and a long right parietovisceral connective.
The Ellobiinian-Carychiinian type has a wide
visceral nerve ring, sometimes with marked
chiastoneury, and a very short right parie-
tovisceral connective. The Pedipedinian-
Melampinian type has a short visceral nerve
ring; the connectives between the cerebral
ganglia and the visceral nerve ring are longer
and symmetrical in the Melampinae, whereas
they are generally shorter in the Pedipedinae,
with the right ones longer than the left ones.

The phylogenetic relationships within the
different subfamilies of halophilic Ellobiidae
are better explained if one considers the
Pythiinae as the representatives of the prim-
itive core from which the Ellobiinae, the Cary-
chiinae and the remaining taxa as a third
group independently radiated. First, the
Pythiinae have a monaulic, glandular pallial
gonoduct and a wide visceral nerve ring.
Pythia still has a very weak remnant of chia-

stoneury and retains a functional open sper-
matic duct. The presence of a pallial gland in
species of Pythia, Cassidula, Ovatella and
Laemodonta suggests a relationship of that
group of the Pythiinae with the terrestrial
Carychiinae. Secondly, the Ellobiinae must
have separated from the primitive stock very
early, for they retain the most primitive ner-
vous system. The reproductive system is di-
aulic; the pallial gonoducts, although glandu-
lar in their entirety, separate immediately
anterior to the seminal vesicle, eliminating the
spermoviduct. Thirdly, the Pedipedinae and
the Melampinae might have arisen from the
same stock. In both subfamilies the visceral
nerve ring is very short, concentrating the
ganglia in the cephalopedal region. In the Pe-
dipedinae there 1$ some proximal concentra-
tion of the anterior mucous gland and pros-
tate gland, giving rise to а partly nonglandular
pallial duct that is monaulic in most genera. In
Leuconopsis and Pseudomelampus the vas
deferens separates some distance before the
female genital opening, giving rise to what 1$
called here an incipient semidiaulic repro-
ductive system [Visser's semidiaulic system],
vaguely resembling a rudimentary step to-
ward the condition in the Melampinae. In this
subfamily the reproductive system 1$ here
called advanced semidiaulic. There is a very
short spermoviduct, on which the inconspic-
uous prostate gland 1$ located, and the an-
terior mucous gland has completely disap-
peared. In Tralia the bursa duct inserts more
distally in the vagina, a fact that suggests that
there might have been a proximal migration
of that structure and of the spermoviduct.
The combination of ganglionic concentration
and nonglandular, advanced semidiaulic pal-
lial gonoduct indicates that the Melampinae
are the least primitive ellobiids.

The subfamilies, listed in order of increas-
ingly derived characters, are Pythiinae, Ello-
biinae, Pedipedinae and Melampinae. As
stated above, however, primitive and derived
characters occur in each subfamily. The
Pythiinae, for example, live farther inland than
any other halophilic ellobiid (Morton, 19550);
this habit is seen as a derived condition.
Some Pedipedinae (Pedipes, Creedonia) do

FIG. 401. Cladograms for Ellobiidae generated by PAUP from data in Tables 5, 6 (Appendix). A, Consensus
of 703 trees, all characters included; B, Consensus of 1396 trees, excluding character G, status of sperm
groove. a, Outgroup; b, Pythiinae; с, Carychiinae; а, Pedipedinae; e, Melampinae; +, Ellobiinae. O, Plesio-
morphies (monauly, pallial ducts entirely glandular, wide visceral nerve ring); 1, Apomorphy diauly; 2,
Apomorphy concentration of visceral nerve ring; 3, Apomorphy pallial ducts partly glandular; 4, Apomorphy
incipient semidiauly; 5, Apomorphies advanced semidiauly and pallial ducts nonglandular.

314 MARTINS

not resorb their inner whorls and are consid-
ered primitive in this respect. The Melampi-
nae retain a free-swimming veliger larva and,
consequently, have a highly heterostrophic
protoconch, which is a primitive feature. The
occurrence of such a variety in the expres-
sion of the different characters within the
Ellobiidae obscures the tracing of a linear
phylogenetic relationship for the family. |
conclude with Morton (1955c) that the evolu-
tion among the Ellobiidae is better under-
stood as following a mosaic pattern, in which
the organs and the mode of life evolve at dif-
ferent rates in the various taxa.

Zoogeography of the Ellobiidae

The Recent Ellobiidae are a primarily trop-
ical family, distributed in three centers.

The first is the Indo-Pacific center, extend-
ing from the East African coast to Polynesia.
This center is characterized by large ellobi-
ids, such as Ellobium, Cassidula and Pythia.
Only four of the 21 genera of halophilic ello-
biids are not represented in this center, the
Mediterranean Ovatella and Auriculinella,
the Eastern Atlantic Pseudomelampus and
the newly created West Indian Creedonia.
Besides Ellobium s.s. and the other two gen-
era that characterize this center, four others
are endemic in the Indo-Pacific, Cylindrotis,
known from the Philippines and Thailand
(Brandt, 1974), Ophicardelus, from the Aus-
tralian region, Allochroa, recorded from the
Pacific Islands and from the Red Sea, and the
widely distributed Auriculastra.

The second is the West Indian center,
which includes the Neartic and Neotropic
regions and Ascension Island. The genus
Melampus characterizes this center. Ten
genera are present in the Western Atlantic, of
which only the new genus Creedonia is en-
demic. Of the 18 western Atlantic species
seven belong to the genus Melampus.

Wallace (1876) assigned with difficulty an-
other mid-South Atlantic island, St. Helena,
to his Ethiopian region, but he did not even
mention Ascension Island. Rosewater (1975)
noted that Ascension Island is very poor in
endemic marine mollusks (only one subspe-
cies and the new ellobiid species Leuconop-
sis manningi) and that the malacofauna of the
island contains even numbers of species
from both sides of the Atlantic. The inclusion
of Ascension Island in the West Indian center
is justified by the presence of the Western
Atlantic Pedipes mirabilis and of the new spe-

cies Leuconopsis manningi. These are the
only ellobiids reported from that South Atlan-
tic island.

The third is the Mediterranean center,
which includes the Macaronesian Islands
(Azores, Madeira, Canary Islands and Cape
Verde Islands), is characterized by the en-
demic Auriculinella and Ovatella, and also by
the more widely distributed Pseudomelam-
pus and Myosotella. Pseudomelampus is
reported from South Africa [Melampus acino-
ides (Morelet, 1889)] and Myosotella, repre-
sented by the extremely variable and equally
overnamed Myosotella myosotis, has be-
come cosmopolitan.

The tropical character of the ellobiids 1$
well exemplified in their Western Atlantic dis-
tribution. Of the 18 recorded species only
three are reported from the American coast
north of southern Florida, Melampus (М.) bi-
dentatus, Melampus (D.) floridanus and the
introduced European Myosotella myosotis.

Bermuda was included by Wallace (1876)
in his Alleghenian subregion of the Neartic,
but it appears that the island should belong
rather in the Antillean subregion of the Neo-
tropical. Eight (67%) of the ellobiid species
not represented on continental shores north
of southern Florida were recorded from Ber-
muda.

Another interesting note is the record of
Tralia (T.) ovula from the African coast. It ap-
pears to be an isolated report, for the genus
has not been reported from elsewhere in Af-
rica; however, the 49 specimens collected by
Dohrn in 1866 and deposited in the Museum
of Comparative Zoology indicate that the
species was not rare at llha do Príncipe, in
the Gulf of Guinea. This West Indian species
might have been transported to Africa in the
ballast of ships, which | think was important
in the dispersal of Myosotella myosotis as
well.

It is worth noting that both species of Pe-
dipes, although broadly overlapping in the
West Indies and Bermuda, overlap very little
in Florida; | could not substantiate in any mu-
seum collection or in my extensive collec-
tions any record of Pedipes mirabilis from the
Florida Keys.

As noted in the remarks under the family,
the fossil record of the Ellobiidae 1$ relatively
poor. It is interesting that the oldest known
fossil ellobiids are the European Carychiopsis
from the Paleozoic of France. This genus re-
sembles the Recent terrestrial Carychium,
which is known from the Jurassic of Asia,

WESTERN ATLANTIC ELLOBIIDAE 315

Europe, America and West Indies (Zilch,
1959). The Paleozoic of Europe contains fos-
sils of the heavily dentate Traliopsis and the
high-spired Stolidoma, which resemble some
Recent examples of the Pedipedinae or the
Pythiinae. They were most probably halo-
philic, as were Rhytophorus and Melam-
poides [Melampinae] from the Cretaceous of
North America. It appears, then, that the el-
lobiids had already invaded the terrestrial
habitat through the Carychiinae during the
Paleozoic, which implies that the group had
separated very early from the prosobranch
stem.

Another interesting note on the paleogeog-
raphy of the Ellobiidae is the presence of the
Recent Indo-Pacific genera Ellobium and
Cassidula in the Eocene of Europe, which
suggests a Tethyan distribution.

The Tertiary and Quaternary ellobiid gen-
era of the West Indies are representatives of
the Recent fauna both in their taxonomy and
geographic boundaries. Marinula from the
Pacific coast of Costa Rica (Dall, 1912), Pe-
dipes from Venezuela (Gibson-Smith 8 Gib-
son-Smith, 1979, 1985), Tralia from Venezu-
ela (Gibson-Smith 4 Gibson-Smith, 1982)
and Jamaica (Woodring, 1928), and Melam-
pus (Detracia) from Virginia (Conrad, 1862)
reflect the modern distribution of the respec-
tive genera.

ACKNOWLEDGMENTS

| wish to express my gratitude to Dr. Rob-
ert С. Bullock, of the University of Rhode Is-
land, for his patient advice and stimulating
enthusiasm during the development of this
work as a dissertation for the Ph.D and for
allowing me to use material in his personal
collection. | thank Dr. Ruth D. Turner and Dr.
Kenneth J. Boss, Curators of the Department
Mollusks of the Museum of Comparative
Zoology, Harvard University, for their gener-
ous help and their kindly allowing me to use
the collection and library of that Department
and never sparing their precious, expert ad-
vice on the intricacies of the nomenclatorial
problems. Also, | wish to thank the late Dr.
Joseph Rosewater and the late Dr. Joseph
Houbrick of the United States National Mu-
seum in Washington, Dr. George M. Davis
and Dr. Arthur Bogan of the Academy of Nat-
ural Sciences of Philadelphia, Dr. William K.
Emerson of the American Museum of Natural
History in New York, Dr. John D. Taylor and

Dr. Fred Naggs of the British Museum (Nat-
ига! History), Dr. Simon Tillier, Annie Tillier
and Dr. J.-P. Hugot of the Muséum National
d'Histoire Naturelle de Paris for kindly pro-
viding access to their collections and for al-
lowing me to use their computers for working
with PAUP. | am indebted to Prof. Brian S.
Morton of the University of Hong Kong and to
Dr. A. Sasekumar of the University of Malay-
sia, who provided me with preserved material
of great importance for anatomical compari-
sons, as well as to Dr. Claude Vaucher of the
Muséum d’Histoire Naturelle de Geneve for
lending me Bourguignat’s type material. The
histological work was greatly simplified and
enormously improved by the generosity and
expertise of Dr. Paul Yevich and Mrs. Carolyn
Barcsz Yevich of the E.P.A. Laboratory, Nar-
ragansett Bay, Rhode Island, and by the con-
tinuous assistance of Ms. Esther Peters; | am
immensely grateful to them. | am most in-
debted to the Rev. Joseph D. Creedon, Pas-
tor of Christ the King Community, Kingston,
Rhode Island, for his unreserved friendship,
warm hospitality and exemplary patience in
following and supporting the development of
my research. | am also grateful to the Univer-
sity of the Azores, to the Fundacáo Calouste
Gulbenkian and to the Instituto Nacional de
Investigacáo Científica for their support in the
various trips to the United States, England
and France to prepare the revision of this
work. Lastly, | wish to express my apprecia-
tion for the loving support and continuous
encouragement of my wife, Micéu.

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NESTERN ATLANTIC ELLOBIIDAE 329

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Revised Ms accepted 6 November 1995

APPENDIX

TABLE 1. Number of specimens used for various morphometric comparisons. G, reproductive organs;

R, radula; S, shell.

Species S R G
M. coffeus 9 5 5
M. coffeus 16 9 —
М. coffeus 5 5 5
M. coffeus 42 — —
М. coffeus 14 5 5
M. coffeus 17. —
М. coffeus 5 5 5
M. coffeus 2 —

M. coffeus 17 — —
M. coffeus 5 5 5
М. coffeus 5 5 5
М. coffeus 5 5 5
M. coffeus 25 5 5
M. bidentatus 10 10 10
М. bidentatus 5 5 5
M. bidentatus 50 — —
M. bidentatus 5 5 5
M. bidentatus 38 23 15
M. bidentatus 5 5 5
M. bidentatus 5 5 5
M. bidentatus 5 5 5
M. bidentatus 5 5 5
M. bidentatus 5 5 5
M. bidentatus 5 5 5
M. bidentatus 18 12 12
M. bidentatus 38 — 5
м. bidentatus 20 20 20
М. bidentatus 5 is)
M. bidentatus 5 5 5
M. bidentatus 39 21 2]

LOCALITY

Hungry Bay, Bermuda

Grassy Key, Florida, U.S.A.

Knight Key, Florida, U.S.A.

Big Pine Key, Florida, U.S.A.

Mullet Key, Florida, U.S.A.

Hawksbill Creek, Eight Mile Rock, Grand Bahama Island,
Bahamas

South Mastic Pt., Andros Island, Bahamas

Shore of Millars Road, New Providence Island, Bahamas

Anegada, Virgin Islands

Piñones, Boca de Cangrejos, San Juan, Puerto Rico

Punta Arenas, Puerto Rico

Laguna Rincón, Bahía de Boquerón, Puerto Rico

Tucacas, Venezuela

Jamestown, Rhode Island, U.S.A.

Narrow River, Wakefield, Rhode Island, U.S.A.

Stonington, Connecticut, U.S.A.

Bivalve, New Jersey, U.S.A.

Cedar Island, North Carolina, U.S.A.

Woodville, North Carolina, U.S.A.

Yemassee, South Carolina, U.S.A.

Crescent, Georgia, U.S.A.

Valona, Georgia, U.S.A.

New Smyrna Beach, Florida, U.S.A.

Grassy Key, Florida, U.S.A.

Knight Key, Florida, U.S.A.

Big Pine Key, Florida, U.S.A.

Big Torch Key, Florida, U.S.A.

Big Bend Road (Rt. 672), Hillsborough Co., Florida, U.S.A.

Hudson, Florida, U.S.A.

Hungry Bay, Bermuda

(continued)

330

MARTINS

TABLE 1. Number of specimens used for various morphometric comparisons. G, reproductive organs;

В, radula; $, shell (Continued).

Species

M.
M.
M.

[ke

E. dominicense

B.

ооо оо Ula dictation <= SIS ISS

. bidentatus
. bidentatus
. bidentatus
. bullaoides
. bullaoides
. bullaoides
. floridanus
. floridanus
. floridanus
. paranus

. morrisoni
. morrisoni
. morrisoni
. morrisoni

morrisoni
monile
monile
monile
ovula
ovula
ovula
ovula
ovula
ovula
ovula
ovula
ovula
mirabilis
mirabilis
mirabilis
mirabilis
mirabilis
mirabilis
mirabilis
mirabilis
ovalis
ovalis
ovalis
ovalis
ovalis
ovalis
ovalis

. succinea
. occidentalis
occidentalis

novimundi
novimundi
novimundi
novimundi
novimundi
novimundi
novimundi
manningi
myosotis
myosotis
myosotis
cubensis

heteroclita

S

—
NN © ND © — м № — фр O1 O1 N

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SNPOONN-

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LOCALITY

Mangrove Cay, Andros Island, Bahamas
Bonefish Pond, New Providence Island, Bahamas
Millars Sound, New Providence Island, Bahamas
Hungry Bay, Bermuda

Somerset Bridge, Bermuda

Big Pine Key, Florida, U.S.A.

Dividing Creek, New Jersey, U.S.A.

Woodville, North Carolina, U.S.A.

Myakka River at Rt. 41, Sarasota Co., Florida, U.S.A.
Pará [Belém], Amazon, Brazil

Key Largo, Florida, U.S.A.

Long Key, Florida, U.S.A.

S of Ocean Dr., Plantation Key, Florida, U.S.A.
Grassy Key, Florida, U.S.A.

Knight Key, Florida, U.S.A.

Long Bird Bridge, Bermuda

Shelly Bay, Bermuda

Hungry Bay, Bermuda

Rock Pt., New Providence Island, Bahamas
Clifton Pt., New Providence Island, Bahamas
Havana, Cuba

Jamaica

Port Salut, Haiti

San Juan, Puerto Rico

Maravén, Borburata, Venezuela

El Palito, Venezuela

Ilha do Principe, Gulf of Guinea

Shelly Bay, Bermuda

Morgan’s Bluff, Andros Island, Bahamas
Paradise Island, Bahamas

Clifton Pt., New Providence Island, Bahamas
San Juan, Puerto Rico

Punta Arenas, Puerto Rico

Е Palito, Venezuela

Puerto Cabello, Venezuela

Ely's Harbour, Bermuda

Big Pine Key, Florida, U.S.A.

Mullet Key, Florida, U.S.A.

Morgan’s Bluff, Andros Island, Bahamas
Clifton Pt., New Providence Island, Bahamas
Punta Arenas, Puerto Rico

Puerto Cabello, Venezuela

Long Key, Florida, U.S.A.

Hungry Bay, Bermuda

Crawl Key, Florida, U.S.A.

Indian Key, Florida, U.S.A.

Pigeon Cay, Bimini Islands, Bahamas

Mintie Bar, South Bight, Andros Island, Bahamas
Morgan’s Bluff, Andros Island, Bahamas
Mangrove Cay, Andros Island, Bahamas
Clifton Pt., New Providence Island, Bahamas
Jack's Bay, Jamaica

English Bay, Ascension Island

Jamestown, Rhode Island, U.S.A.

Newport River, Beaufort, North Carolina, U.S.A.
Hungry Bay, Bermuda

Grassy Key, Florida, U.S.A.

Big Torch Key, Florida, U.S.A.

Hungry Bay, Bermuda.

WESTERN ATLANTIC ELLOBIIDAE 331

TABLE 2. Shell morphometry of several species of Marinula and of Creedonia succinea. Measurements
of Creedonia succinea from specimens in my collection; all others from Connolly (1915). AL/BWL, ratio
aperture length/body whorl length; BWL/SL, ratio body whorl length/shell length; SL, shell length (mm);
W, number of whorls.

SPECIES SL W BWL/SL AUBWL
M. pepita 10.4 4.00 0.89 0.76
M. pepita 10.3 4.00 0.91 0.64
M. pepita 9.3 4.00 0.86 075
M. xanthostoma 9.9 5.00 0.88 0.71
M. tristanensis 10.4 3:19 0.92 0.92
M. velaini 9.0 3.50 0.93 0.79
М. parva 9.2 4.00 0.91 0.71
М. filholi 7.4 3.50 0.85 0.71
М. mandroni 4.0 — — —
C. succinea 3:3 4.50 0.80 0:72
C. succinea 3:3 4.25 0.79 0.69
C. succinea 3:0 4.25 0.79 0.76
C. succinea 3.0 4.20 0.81 0.72
С.

succinea 2.8 4.00 0.80 0.70

TABLE 3. Radular formulae of species of Marinula and of Creedonia succinea.
Transitional teeth were counted as lateral teeth; in some cases lateral teeth and
marginal teeth were pooled.

SPECIES RADULAR FORMULA SOURCE

M. xanthostoma (112 + 1 + 112) x 180 Odhner (1925)

M. parva (135 + 1 + 135) x 180 Odhner (1925)

M. juanensis (45 + 110 + 1 + 110 + 45) x 180 Odhner (1925)

M. tristanensis (75 + 35 + 1 + 35 + 75) Connolly (1915)

M. filholi (121 +1 +121) Powell (1933)

C. succinea (14 + 10 + 1 + 10 + 14) x 65 Martins, this paper
C. succinea (12 + 14 +1 + 14 + 12) x 82 Martins, this paper
C. succinea (13 + 12 + 1 + 12 + 13) x 62 Martins, this paper
C. succinea (12 +12 + 1 + 12 + 12) x 72 Martins, this paper

TABLE 4. Shell measurements and radular counts of young Melampus coffeus. LOC, localities: ВАН,
Millars Road, New Providence, Bahamas; PR, Bahía de Boquerón, Puerto Rico. L,, Lz, Lz, number of
lateral tooth first appearing unicuspid, bicuspid and tricuspid; M, number of marginal teeth; R, number
of horizontal rows; SL, shell length (mm); T, number of transitional teeth; W, number of teleoconch
whorls; 2-7, number of denticles first appearing on second through fourteenth marginal tooth.

LOC SL W ROUE nee LOU: 2 A a a 6 7
PR 266° 4550 E 2 2 5 8 9
PR 348 585 75 1 OF Aes Ao ae 7 40 №
BAH 268 Ге мо “F GS wet о 2 ES м м =
ВАН 476 750 70 — 6 5 1 16 2 6 8 12 14 —

332

MARTINS

TABLE 5. Characters used to generate the cladogram presented in Fig. 401. Characters (A-l) as
explained in the table; character states (0-3), the condition of each character, from primitive to
advanced when polarized; type, polarized (ordered) or nonpolarized (unordered) character; weight (1-4),
ascending assigned importance of the character.

CHARACTERS
A. Ацу (separation of pallial gonoducts)

В. Glandular cover of pallial gonoducts

C. Position of insertion of bursa duct

D. Origin of posterior vas deferens
(as a nonglandular duct)
E. Pallial gland

F. Status of sperm groove

С. Chiastoneury

H. Oesophageal/Visceral rings (ratio of
total length of connectives,

excluding ganglia)

|. Parietovisceral connectives
(ratio left/right)

TYPE
Ordered

Ordered

Ordered

Ordered

Ordered

Ordered

Ordered

Ordered

Unordered

WEIGHT
4

CHARACTER STATES

0.- Monauly

1.- Incipient semidiauly
2.- Advanced semidiauly

3.- Diauly

0.- Entirely covered
1.- Partly covered
2.- Naked

0.- Near female genital opening

1.- On anterior third of oviduct

2.- On posterior third of oviduct
0.- Opposite insertion of bursa duct
1.- Posterior insertion of bursa duct

0.- Present
1.- Absent
0.- Open

1.- Closed
0.- Present
1.- Absent
0.- <0.80

1.- 0.80-1.20
2.- 1.21-1.99
3.- >2.00

0.- <0.90

1.- 0.90-1.9
2.- 2.0-4.0

3.- >4.0

TABLE 6. Taxa and data matrix used to generate the cladogram presented in Fig. 401. Names in
boldface are the taxon and rank represented in the cladogram. Explanation of characters in Table 5.
Data on the outgroup Ringicula sp. from Fretter (1960) and Gosliner (1981); all others are original. ? no
information; * character reversed, owing to sinistrality of the taxon.

TAXON

Ringicula sp. [outgroup]

Pythia plicata (Férussac)

Myosotella myosotis (Draparnaud)

Ovatella aequalis (Lowe)

Laemodonta cubensis (Pfeiffer)

Cassidula mustelina (Deshayes)

Carychium cf. tridentatum (Risso)

Ellobium (Auriculodes) dominicense
(Draparnaud)

Ellobium (Ellobium) aurismidae (Linnaeus)

Auriculinella bidentata (Montagu)

Blauneria heteroclita (Montagu)

Pedipes mirabilis (Mühlfeld)

Creedonia succinea (Pfeiffer)

Marinula (Cremnobates) xanthostoma
(Н.& A. Adams)

Microtralia occidentalis (Pfeiffer)

Pseudomelampus exiguus (Lowe)

Marinula (Marinula) tristanensis Connolly

Leuconopsis novimundi Pilsbry & McGinty

Tralia ovula (Bruguiere)

Melampus (Melampus) coffeus (Linnaeus)

Melampus (Detracia) bullaoides (Montagu)

LOCALITY

North Atlantic
Thailand
Bermuda
Azores
Bermuda
Hong Kong
Azores
Florida, U.S.A.

Thailand
Azores
Bermuda
Bahamas
Florida, U.S.A.
Oman

Bermuda

Azores

Gough Id., $ Atlantic
Bahamas

Puerto Rico

Florida, U.S.A.
Florida, U.S.A.

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MALACOLOGIA, 1996, 37(2): 333-348

ASPECTS OF THE POPULATION DYNAMICS AND PHYSIOLOGICAL ECOLOGY
OF THE GASTROPOD PHYSELLA CUBENSIS (PULMONATA: PHYSIDAE) LIVING

IN А WARM-TEMPERATE STREAM AND EPHEMERAL POND HABITAT

Donald L. Thomas & James В. McClintock

Department of Biology, University of Alabama at Birmingham, UAB Station, Birmingham,

Alabama 35294, USA

ABSTRACT

Population density, size frequency, and reproduction of the pulmonate gastropod Physella
cubensis living in a central Alabama stream and ephemeral pond habitat were assessed over
a three-year period from January 1989 through December 1991. These parameters covaried
seasonally and from year to year with fluctuating environmental temperature and precipitation.
Population dynamics of ephemeral pond snails were also affected by episodic drying events.
Physella cubensis is able to survive habitat desiccation in ephemeral pond habitats by burrow-
ing into the hypopheric zone of the sediments. This behavior is displayed only by juvenile snails
(1-5 mm shell length). Overwintering in the sediments 1$ restricted to young adult snails (5-8
mm shell length). Food quality and particularly temperature were found to influence growth and
survivorship. Optimum temperature for growth and survivorship was 25°C (vs. 15°C and 35°C).
Snails raised at 15°C and 25°C exhibited a dramatic shift in the timing of first oviposition (60 vs.
18 days, respectively), but did not differ significantly in body size at first reproduction. Snails
raised at 35°С appeared thermally stressed and failed to oviposit. Food quality influenced
reproductive output, with only snails fed medium- and high-quality diets producing eggs. Both
field and laboratory studies indicate that P. cubensis living in a warm-temperate climate ex-
emplify an opportunistic life history strategy in which such traits as rapid juvenile growth and
attainment of maturity, shortened lifespan, high fecundity, and constant reproduction over the

duration of the adult lifespan are favored.

Key words: Physella cubensis, population dynamics, physiological ecology, temperature,

food quality.

INTRODUCTION

Studies of seasonal fluctuations in density,
size frequency, and reproductive activity of
organisms in a variety of habitats and across
diverse taxa are central to understanding
how fitness is maximized under changing
environmental constraints. Much of the liter-
ature on the population dynamics and life
histories of freshwater gastropods consists
of studies from northern climates (e.g., De-
Witt, 1955; McNeil, 1963; Eisenberg, 1966;
Clampitt, 1970; Eisenberg, 1970; McGraw,
1970; Eckbald, 1973; Hunter, 1975; Browne,
1978; Brown, 1979a,b, 1982; Diamond, 1982),
whereas studies in southern temperate and
subtropical climates are comparatively few
(Cridland, 1957; McMahon, 1975; Gray, 1987;
Brown et al., 1989; Crowl, 1990).

In his extensive review of the life cycles of
freshwater snails, Calow (1978) indicated that
many aquatic pulmonates studied in cool-
temperate environments have annual repro-
ductive cycles with a single spring breeding

333

period. Calow predicted that in warm-tem-
perate and subtropical environments subject
to stochastic seasonal drying events, pulmo-
nates might be expected to exhibit a short-
ened lifespan, higher fecundity, and increased
mortality. These environments would neces-
sitate rapid maturation and opportunistic,
seasonally plastic reproductive patterns. Sim-
ilar predictions have been advanced by Pi-
anka (1970), Giesel (1976), Browne 8 Russell-
Hunter (1978), Brown (1979a), Stearns (1980),
and Brown (1985), although few empirical
studies of pulmonates living in warm-temper-
ate and subtropical environments (e.g., Gray,
1987) have been conducted.

Temperature has been shown to have a
significant influence on growth, reproduction,
and microhabitat selection in a number of
freshwater gastropods (DeWitt, 1954; Dun-
can, 1959; Beames 4 Lindeborg, 1967; Clam-
pitt, 1970; McGraw, 1970; van der Schalie 8
Berry, 1973; McMahon, 1975; Brown, 1979a;
McMahon & Payne, 1980; Ross & Ultsch,
1980; Hernandez et al., 1981; Krkac, 1982;

334 THOMAS 8 MCCLINTOCK

Gray, 1987; Blandenier & Perrin, 1989; Brown
et al., 1989; Lam & Calow, 1990; Brackenbury
& Appleton, 1991). Thomas 8 McClintock
(1990) found that elevated water temperature
increased rates of embryonic and juvenile
growth and also influenced the body size at
which sexual maturity (first oviposition) was
attained in the pulmonate gastropod Physella
cubensis. It has been suggested that temper-
ature might be one of the most important fac-
tors influencing the life histories of pulmonate
gastropods occupying warm-temperate en-
vironments (DeWitt, 1954; Duncan, 1959;
Calow, 1978; Brown, 1979a; Thomas 8 Mc-
Clintock, 1990).

The frequent drying of ephemeral streams
and ponds is of particular importance in shap-
ing the life histories of non-migratory aquatic
invertebrates, including molluscs (Paterson &
Fernando, 1969; Eckbald, 1973; Calow, 1978;
Hornbach, 1980; Brown, 1983, 1985; Crowl,
1990; Wyngaard et al., 1991). Stochastic
drying events may affect age-specific survi-
vorship, timing of sexual maturity, length of
reproductive period, and fecundity of aquatic
pulmonates living in ephemeral habitats
(Cridland, 1957; Eckbald, 1973; Brown, 1985).
Several aquatic pulmonates use the hypo-
pheric zone of sediments as a refuge from
desiccation (DeWitt, 1955; Cridland, 1957;
Clampitt, 1970; Brown, 1979a, 1985; Crowl,
1990). This burrowing behavior may be
size specific (DeWitt, 1955; Cridland, 1957;
Brown, 1985; Crowl, 1990). Populations of
Physella cubensis occurring in small streams
and ponds in central Alabama are known to
survive extensive periods of desiccation dur-
ing summer months (Thomas, pers. obs.).
Size-specific survivorship during desiccation
events may be important in regulating popu-
lation dynamics and ultimately selecting for
life history traits of P. cubensis.

Food quality (availability of nutrients and
energy) may have significant effects on the
relationship between fecundity, egg size, and
age of sexual maturity in invertebrates
(Spight 8 Emlen, 1976; Quian 8 Chia, 1991).
К has been argued that for some lymnaeid
(Bovbjerg, 1968; Brown, 1979b) and physid
snails (Clampitt, 1970) food supply may be a
negligible factor in limiting growth and distri-
bution for these taxa due to the generalist
nature of the diet. The need for more detailed
studies of the effects of food quality on fe-
cundity, egg size, and adult maturation has
been emphasized by Calow (1978), Brown
(1983), and Brown et al. (1985).

The specific objectives of this study are to:
(1) document the population dynamics and
reproductive cycles of Physella cubensis in a
warm-temperate climate over a three-year
period and (2) measure the effects of temper-
ature and food quality on juvenile and adult
growth, onset of oviposition, egg size, and
fecundity under controlled laboratory condi-
tions.

METHODS
Aspects of Population Dynamics

Abundance of Physella cubensis and their
egg masses in Shades Creek, a shallow, low-
velocity stream in central Alabama (33°34’N;
86°53’W), as well as those in a nearby
ephemeral pond habitat, were assessed from
January 1989 through December 1991. Four
sampling sites were established, including
three in the stream and one in the ephemeral
pond. Each site was delineated by a ten-
meter transect marked at either end by steel
stakes. Stream sites | and Il extended in di-
rections parallel and perpendicular to the
current, respectively. Average water depth in
stream sites | and ll ranged from 2 to 30 cm,
and substrata consisted of a combination of
coarse silt and small cobble (Wentworth
scale). While sites | and ll were similar in wa-
ter depths and in substrata, site | extended
through relatively calm water (mean velocity
<0.003 m/sec.), whereas portions of site Il
extended into a channel which experienced
higher mean water velocities (0.25 m/sec.).
Stream site Ш, located approximately 150 т
upstream from sites | and II, extended paral-
lel to the current along a smooth bedrock
shelf at a distance of 1 m from the stream
bank. Water depth in stream site Ш ranged
from 2-18 cm, and average water velocity
was 0.75 m/sec. With the exception of vari-
ous filamentous algae, no conspicuous mac-
rophytes were present at any of the stream
sites. A fourth study site was established in
an isolated, ephemeral pond located 36 m
from the stream. The pond was a shallow
(maximum depth 20 cm) rain-fed depression
of approximately 25 m° in area. The substra-
tum was characterized by a 3-8 cm layer of
leaf litter, algae and detritus. Conspicuous
aquatic macrophytes in the pond included
Typha latifolia and Juncus effusus.

Sampling was conducted approximately
every two weeks. On each sampling date, a

PHYSIOLOGICAL ECOLOGY OF PHYSELLA CUBENSIS 335

1-m? quadrat was placed at three randomly
determined positions along each transect.
Snails were enumerated by hand collecting
all snails from each quadrat. All vegetation,
stones and other debris were examined thor-
oughly, with care being taken to minimize
habitat disruption. Sampling by dredging or
coring was deemed inappropriate due to the
rocky nature of the stream sites and to the
disturbance to the habitat incurred by these
methods. The long-axis shell length of up to
the first 50 randomly selected individuals
from each quadrat were measured to the
nearest 0.5 mm. Egg masses of Physella
cubensis, found attached to stones or other
solid debris, were counted. All snails and egg
masses were restored to their original quad-
rats following field measurement. Air temper-
atures as well as stream and pond water tem-
peratures were recorded. Precipitation data
(cm/week) was obtained from a National
Weather Service Station located 5.4 km from
the study area. Relationships between the
temporal abundance of stream and pond
snails and the environmental parameters of
temperature and precipitation were analyzed
statistically using the Spearman-Rank Corre-
lation (Zar, 1974). Size frequencies of stream
snails emerging from overwintering in the
sediments in the spring and of pond snails
emerging from aestivation in the sediments
following dry periods were statistically tested
for similarity (independence) using a Contin-
gency Table Analysis (G-test, Sokal 4 Rolf,
1981).

The size of aestivating snails was exam-
ined in the field. Core samples of substratum
from an ephemeral pond (separate from, but
with similar characteristics to the one in
which population sampling was conducted)
in which dense populations of Physella
cubensis had been noted were collected two
weeks following the complete drying of the
pond in late May 1992. Bottom sediments
were collected by using a 12-cm diameter
hollow steel cylinder, driven 15 cm deep into
the substratum at five randomly selected lo-
cations. Each sample consisted of 1.5 L of
sediment. Each sediment sample was im-
mersed in distilled water and allowed to
stand for a 24-h period, after which the sam-
ples were individually washed through a
0.25-mm mesh sieve and the snails col-
lected. Vital condition of aestivating snails
was confirmed by placing them in distilled
water and observing active emergence from
the shell over a 24-h period. The long-axis

shell length of all living snails was measured
to the nearest 0.5 mm.

Effects of Temperature and Food Quality

The effects of temperature and food quality
on growth, reproduction and survivorship of
Physella cubensis were examined in the lab-
oratory. Two-hundred and twenty-five labo-
ratory-hatched snails (12 h neonates) were
distributed into 45 glass finger bowls (12 cm
diameter; 5 cm depth) to yield five snails per
bowl. Each bowl received 150 ml of distilled
water, which was changed daily. Snails were
placed in three temperature treatments of
15°C, 25°C and 35°C (15 bowls/treatment) in
environmental chambers on a 12:12 h light:
dark photoperiod. Snails in each temperature
treatment were further divided into low, me-
dium, and high quality food treatments, each
consisting of five replicate bowls containing
five snails per bowl. For the purposes of this
study, different amounts of food embedded
in agar were considered to represent differ-
ent quality foods. We chose to define food
quality as the concentration of nutrients and
energy content per unit food mass (Bedding-
field 8 McClintock, 1993). The three different
quality diets were prepared from 5% gum
agar (Sigma) combined with a proteinaceous
powdered food compound originally devel-
oped for omnivorous sea urchins (J. M.
Lawrence, pers. comm.). Three different con-
centrations (qualities) of artificial food were
prepared (0.5%, 7%, and 15% dry food, re-
spectively) and fed to the snails ad libitum.

Snails were maintained in the nine temper-
ature and food quality treatments for 138
days. Every third day, the long-axis shell
length of each snail was measured. Egg
masses, if present, and numbers of eggs per
mass were recorded for each bowl. Individual
eggs were measured (long and short diame-
ters) for the first 70 days of oviposition using
a compound microscope equipped with an
ocular micrometer. Growth and survivorship
(arcsine transformed data) of snails were
compared statistically using a two-way
ANOVA followed by pairwise comparisons
using a Scheffé test (Zar, 1974). Statistical
analyses were applied on days 18 and 60
(dates at which first oviposition occurred for
25°С and 15°С temperature treatments, re-
spectively). Sizes of eggs produced were
compared statistically between different tem-
perature and food quality treatments using a
Student's t-test (Zar, 1974).

336 THOMAS 8 MCCLINTOCK

RESULTS
Aspects of Population Dynamics

Physella cubensis were active on the sed-
iment surface in the three stream sites 166.7
+ 37.3 (+ 1 SE) days/year over the three-year
period. Annual abundance patterns of stream
snails (Fig. 1) and egg masses (Fig. 2) were
significantly correlated (г. = 0.825, Р <
0.0001) with зеазопа! shifts in temperature
(Fig. 3), with snails less common or absent
from the surface of the substratum during
cold winter months. Overwintering via aesti-
vation in the sediments of stream snails oc-
curred from late September to late May. In
stream sites | and |, mean densities of snails
in 1990 and 1991 were twice those observed
in 1989. Abundance of stream snails during
the warm months of each year were inversely
correlated (г. = —0.345; Р < 0.01) with
marked seasonal precipitation events, such
as the heavy rains that occurred in the early
summer of 1989 (Fig. 4). Stream snails exhib-
ited multiple peak densities annually, which
varied in magnitude from year to year (Fig. 1).
Egg masses were present in all stream sites
for the duration of the presence of adult
snails. Peaks in egg-mass density (Fig. 2)
were synchronous with peaks in snail den-
sity. Patterns of snail and egg-mass abun-
dance at sites | and II were similar from year
to year. Both snail and egg-mass densities
were consistently lower at site Ш than those
at sites | and Il.

The size frequencies of stream _ snails
emerging in the spring indicate that overwin-
tering in the sediment was limited to young
adult snails (Mean + 1 SE: 6.5 + 1.2 mm shell
length, n = 213). The size frequencies of
emergent snails did not differ significantly be-
tween the three stream sites (G = 0.42; P =
0.94) or from year to year within sites (G =
0.41; P = 0.92). Changes in size distributions
(Figs. 5-7) suggest that periods of peak
growth for stream snails occurred during the
months of July and August. At this time of
year, growth from young juvenile (1-2 mm
shell length) to large adult (9-12 mm shell
length) appeared to require approximately
5-6 weeks.

Temporal periodicity of pond snail densi-
ties (Fig. 1) was primarily related to episodic
drying events, which occurred most fre-
quently between the months of March and
November (Figs. 5-7). Intra-annual multiple

peak densities of snails and egg masses (Fig.
1, 2) were significantly correlated (r, = 0.414;
P < 0.0001) with seasonal peaks in precipi-
tation which hydrated the pond. Pond snails
did not exhibit as constant a period of over-
wintering in the sediments as did stream snail
populations. Rather, pond snails were active
on the surface of the substratum during win-
ter on days when temperatures were mild.
Pond snails and egg masses were never ob-
served on sampling dates when the pond
temperature was below 14°C. Pond snails
emerging following desiccation events were
limited to juveniles (1-5 mm shell length;
Figs. 5-7). The size frequencies of emergent
pond snails did not differ significantly among
different intra-annual (G = 4.72; P = 0.53) re-
hydration events or from year to year (G =
4.53; P = 0.50). Size distributions suggest
that during warm seasons (e.g., May, 1990;
Fig. 6), growth from young juvenile (1-2 mm
shell length) to large adult (9-12 mm shell
length) requires approximately four weeks. In
contrast, during periods of cold temperatures
in winter and early spring months (e.g., 1989;
Fig. 5), growth from juvenile (2-4 mm shell
length) to young adult (6-8 mm shell length)
required approximately 12 weeks. In general,
peak growth for pond snails occurred in the
wet spring months, in contrast to the dryer
late summer months for stream populations.

Physella cubensis exhibited differential
survivorship in response to a desiccation
event in the field. Of 75 living snails collected
from the cores of dry pond sediments, 82.3%
were less than 4 mm in shell length. No living
snails that measured greater than 5 mm shell
length (mean + 1 SE shell length = 2.2 + 0.13
mm) were extracted from the pond sedi-
ments. The empty shells of medium- to large-
size snails were observed on the surface of
the dry pond sediments, but were not
counted.

Effects of Temperature and Food Quality

Growth rates and onset of oviposition of
Physella cubensis raised at 15°C, 25°C and
35°C and fed low, medium, and high quality
diets are shown in Figure 8. Physella cuben-
sis raised at 35°C grew at rates similar to
those raised at 15°C irrespective of food
quality (day 60 Sheffé S = 0.174; P = 0.858).
Growth of snails raised at 25°C was signifi-
cantly greater by day 18 (ANOVA, F = 137.5,
Р < .0001; Scheffé S = 2.02, Р < .0001) than
that of snails in 15°C and 35°C treatments,

PHYSIOLOGICAL ECOLOGY OF PHYSELLA CUBENSIS

100

Mean Density(#/m° )

1989

337

Stream site I

Stream site II

Stream site Ш

Pond

1990 1991

FIG. 1. Seasonal mean densities of stream and pond populations of Physella cubensis from January 1989
through December 1991. Note different scaling of vertical axis for stream site Ill.

regardless of food quality. First oviposition
occurred on days 18 and 60 for 25°C and
15°C treatments, respectively. Snails raised
at 35°C did not oviposit and experienced
high mortality, whereas snails raised at 15°C
experienced minimal mortality and attained
sexual maturity. Cumulative egg production
50 days after first oviposition occurred for
15°C and 25°C treatments was 678 and 659
eggs, respectively. Sizes of eggs (mean long-

axis diameter + 1 SE = 0.3 + 0.12 тт; п =
500) did not differ significantly between tem-
perature treatments (t = 0.939; P = 1.5). Tem-
perature had a significant influence on survi-
vorship (ANOVA, Е = 23.6; Р < 0.0001), with
the highest mortality occurring at 35°С
(Scheffé $ = —28.5, P < 0.0001) and lowest
mortality occurring in snails raised at 15°С
(Scheffé $ = - 16.23, Р < 0.002) as shown in
Figure 9.

338 THOMAS & MCCLINTOCK

120

Stream site I

40 Pond

Mean Egg Mass Density (#/m? )

20

0
Ws ВЕ

1969

We SD, 5 Е

Stream site Il

Stream site Ш

W 52 S Е

1990 1991

FIG. 2. Seasonal mean densities of egg masses laid by stream and pond populations of Physella cubensis
from January 1989 through December 1991. Note different scaling of vertical axes for stream site Ш and

pond site.

Within each of the three temperature treat-
ments, snails fed medium and high quality
diets did not differ significantly in growth (day
60 Scheffé S = 0.241, P = 0.612) or in the
timing of first oviposition. Snails fed low qual-
ity diets, however, grew significantly more

slowly (day 18 ANOVA, F = 0.33, P < 0.0001;
Scheffé S = 1.63, P < 0.0001) than snails fed
medium and high quality diets. Moreover,
snails fed low quality diets did not oviposit
regardless of temperature. Cumulative egg
production 50 days after first oviposition oc-

PHYSIOLOGICAL ECOLOGY OF PHYSELLA CUBENSIS 339

Stream

25

15

Pond

Temperature (°C)

1989 1990 1994

FIG. 3. Water temperatures of stream and pond from January 1989 through December 1991. Discontinu-
ities in the temperatures recorded for the pond site indicate periods when the pond was dry.

а
о

W Sp $ F W $ 5 F W E S F W
1989 1990 1991

FIG. 4. Weekly precipitation recorded at a weather station located 5.4 km from the study site from January
1989 through December 1991.

Precipitation (cm?)

curred for snails fed medium and high quality icantly between medium and high quality
diets was 675 and 655 eggs, respectively. food treatments (t = 0.35; P = 0.13). Food
Egg size (mean long-axis diameter + 1 SE = quality did not significantly influence survi-

0.3 + 0.12 mm; n = 500) did not differ signif- vorship at 15°C (day 60 Scheffé $ = —6.46, P

340 THOMAS 8 MCCLINTOCK

ы Site I Site II Site Ш Ропа
1989 100 J
100
130
125 M

Frequency (%)

ANA

0 2 4 6 8 10 12 14

Shell length (mm)

FIG. 5. Size frequencies (%) of Physella cubensis occurring in stream sites and pond in 1989. Discontinu-
ities in both stream and pond graphs represent sampling periods when snails were absent from the surface
of the substratum. Sample sizes are indicated to the left of each histogram.

PHYSIOLOGICAL ECOLOGY OF PHYSELLA CUBENSIS 341

Site I Site I Site HI Pond Е
80 30
1990 70 |

Frequency (%)

0 2 4 6 8 10 12 14

Shell length (mm)

FIG. 6. Size frequencies (%) of Physella cubensis occurring in stream sites and pond in 1990. Discontinu-
ities in both stream and pond graphs represent sampling periods when snails were absent from the surface
of the substratum. Sample sizes are indicated to the left of each histogram.

342
80
60
S
S 19
32
>
O 10
5
119
= 40
о 150
=
FL,

THOMAS & McCLINTOCK
Site I Site I Site Ш Pond

1991 à
7 M
9
115
45 A
140
150 M
150
1 5 ere

и

8 5 5;
13

150 J

150 ТЕ

2 4 6 8 10 12 14

Shell length (mm)

FIG. 7. Size frequencies (%) of Physella cubensis occurring in stream sites and pond in 1991. Discontinu-
ities in both stream and pond graphs represent sampling periods when snails were absent from the surface
of the substratum. Sample sizes are indicated to the left of each histogram.

PHYSIOLOGICAL ECOLOGY OF PHYSELLA CUBENSIS 343

10 15°С —t— Low
—— Medium
8 ===. Hiph /;
A
“+ 225 » : в
— 2 Gi |
Е o
— 0] asec
Е 8
—
OD 6
e
oD) 4
poi
== 2
4)
Rz 0)
2 10
= A
= 35°С
$)
= 6
4
2
0
0 50 100 150

Time (days)

FIG. 8. Growth of Physella cubensis raised at 15°С, 25°С and 35°С and fed low, medium, and high quality
diets. Arrows indicate the dates of first oviposition. Shown are means + 1 S.E. (n = 25 at time 0).

344 THOMAS & MCCLINTOCK

100 353

Mean survivorship (%)

0 50 100 150

Time (days)

FIG. 9. Mean survivorship of Physella cubensis raised at 15°C, 25°C, and 35°C and fed low, medium, and
high quality diets. Shown are means + 1 S.E. (п = 5 replicate bowls beginning with five individuals per bowl).

= 0.53) but caused significant differences in 25°С and 35°С by day 18 (Scheffé $ =
survivorship in snails fed low and high quality —14.69, Р < 0.005).
foods (ANOVA, Е = 6.4, Р < 0.004) raised at A significant interactive effect between

PHYSIOLOGICAL ECOLOGY OF PHYSELLA CUBENSIS 345

temperature and food quality on growth was
observed by day 18 (ANOVA, Е = 14.97, Р <
0.0001) but not at day 60. In contrast, signif-
icant interactive effects between temperature
and food quality on survivorship were ob-
served both on day 18 (ANOVA, F = 4.61, P <
0.005) and on day 60 (ANOVA, F = 5.57, Р <
0.001). Parasitism of snails was not observed
in any treatment.

DISCUSSION

Temporal abundance patterns of stream-
dwelling Physella cubensis varied from year
to year and appeared to be related primarily
to seasonal fluctuations in temperature and
precipitation. Seasonally low temperatures
appeared to restrict the presence of stream
snails on the sediment surface to a period of
approximately seven months per year. Young
adult stream snails overwintered in the sedi-
ments during the colder months of each year.
Similar over-wintering behavior has been
recorded for other freshwater gastropods
(Paterson & Fernando, 1969; Clampitt, 1970;
McGraw, 1970; Browne, 1978). Low popula-
tion densities and abbreviated periods of
abundance during warm months were corre-
lated with heavy precipitation events (e.g.
May-June 1989). High water velocities ac-
companying flooding of the stream may have
dislodged exposed individuals, particularly in
site Ш, which was located on a smooth rock
shelf. Flood-associated siltation may have
also restricted or temporarily eliminated ac-
cess to detrital and algal food sources. Snails
appeared to avoid regions of the stream that
regularly experienced relatively high water
velocities, limiting their distribution to pools
and riffles within the stream (Thomas, pers.
obs.). In conditions of high water velocity as-
sociated with heavy precipitation, P. cuben-
sis seek shelter by burrowing into the sub-
stratum or by moving under rocks or other
immobile debris (Thomas, pers. obs.). Similar
behaviors have been observed in P. virigata
(Crowl 8 Schnell, 1990). The low density of
snails consistently observed at stream site III
may have resulted partly from the lack of ref-
ugia from periodic water scouring in this site.

Size-distribution data indicate that peak
growth of Physella cubensis in the stream oc-
curred in the late summer months when wa-
ter temperatures were high and when precip-
itation was low. Based on size-distribution
analysis, growth of snails from young juve-

niles to large adults typically required five
weeks during warm months.

Temporal abundance patterns of the
ephemeral pond snails studied was consid-
erably more irregular than of that observed in
stream populations. The abundance of pond
snails seemed to be primarily regulated by
episodic drying events. Desiccation of the
pond resulted in extensive mortality, as evi-
denced by the abundance of empty shells.
No migration of snails from the pond was ob-
served. The interruption of the life cycle of
pond snails by desiccation events, and a
mandatory time period required for sexual
maturation, are likely factors responsible for
consistent low densities of egg masses. Per-
sistent precipitation events of late spring
1989 extended the time period during which
the pond contained water. During this period,
snails experienced a period of rapid growth,
with young juveniles (1-2 mm shell length)
attaining adult sizes (9-10 mm shell length) in
approximately four weeks. Reproductive out-
put was also elevated during this period.
Comparatively slow growth occurred in late
winter and early spring of 1989, during ex-
tended periods of low temperature.

Pond snails are capable of surviving drying
events by burrowing into the moist zone of
sediments. However, field observations indi-
cate that only juvenile snails (<5 mm shell
length) are able to escape desiccation in this
manner. This size-specific survivorship con-
curs with field observations of DeWitt (1955)
and Clampitt (1970) for Physa (= Physella)
gyrina and of Brown (1985) for Lymnaea elo-
des, but contrasts with the findings of McNeil
(1963), who observed greater survivorship in
larger (7-16 mm shell length) aestivating
Stagnicola palustris.

Adult Physella cubensis subjected to dry
conditions in the laboratory (Thomas, unpub-
lished data) were observed to produce a dry
mucous film across the shell aperture. How-
ever, snails in the laboratory survived in this
manner no longer than 24 hours in the ab-
sence of moisture, indicating that this means
of protection from desiccation is effective
only for a short time. This behavior may en-
hance survival in dry conditions in the field on
a short-term basis, perhaps allowing snails to
temporarily escape aquatic predators or wa-
ter pollution.

As adults do not survive desiccation, im-
mature Physella cubensis must replenish
pond populations following seasonal desic-
cation events. Immature snails must first sur-
vive a mandatory period of growth in order to

346 THOMAS & MCCLINTOCK

oviposit. Such conditions as these would ul-
timately be expected to favor selection for
rapid juvenile growth and maturation. More-
over, continuous reproduction following mat-
uration may ultimately be essential in main-
taining a significant number of juveniles in the
population capable of opportunistically sur-
viving episodic desiccation events. Both
rapid growth and attainment of sexual matu-
rity appear to be mediated further by more
proximate factors, such as water tempera-
ture and food quality.

Laboratory studies support the hypothesis
that temperature plays a central role in the
regulation of growth and reproduction of
Physella cubensis (Thomas & McClintock,
1990; present study). Snails raised at 25°C
grew faster over the first 50 days of the study
and began ovipositing 42 days earlier than
those raised at 15°C. Temperatures of 25°C,
which are frequently experienced by stream
and pond populations of Physella cubensis
during summer months in Alabama, appear
to be optimum for growth and reproduction.
Growth at 25°C was considerably reduced
following the onset of ovipostion, likely indi-
cating a shift in resource allocation from
growth to reproduction. In contrast to snails
raised at 15°C, those raised at 10°C (Thomas
& McClintock, 1990) never attained sexual
maturity during the study. Such characteris-
tics as rapid growth and attainment of matu-
rity as observed at 25°C are consistent with
those considered adaptive by Calow (1978)
and Brown (1985) for pulmonates living in
warm-temperate, seasonally unstable habi-
tats subject to rapid daily temperature shifts.

Physella cubensis exposed to a sustained
temperature of 35°C apparently suffer signif-
icant metabolic stress. It is likely that this
temperature is near the lethal level of temper-
ature tolerance for P. cubensis. Although
temperatures of 33°C have been recorded in
shallow ephemeral pond habitats (Thomas,
unpublished data), temperatures of this mag-
nitude are likely experienced for only brief pe-
riods by P. cubensis (Thomas, pers. obs.).
Van der Schalie and Berry (1973) found that
while physids had the widest range of tem-
perature tolerance of the three North Ameri-
can aquatic pulmonate genera, Physella gy-
rina raised at 34°C experienced 100%
mortality after only 49 days. Clampitt (1970)
reported that P. gyrina raised at 35°C expe-
rienced 50% mortality at 11-13 days.

When resources are limited, nutrients may
be differentially allocated towards growth

and reproduction (Spight & Emlen, 1976;
Rollo & Hawryluk, 1988). Whereas low qual-
ity food restricted growth and prevented
attainment of sexual maturity in Physella
cubensis, there were no significant differ-
ences in growth, fecundity, onset of oviposi-
tion, or egg size between snails fed medium
and high quality artificial foods. The medium
quality food may have provided adequate nu-
trients relative to the capacity of assimilation
efficiency in P. cubensis, rendering no addi-
tional benefit to the higher quality food. The
slow growth and lack of oviposition of snails
provided low quality food indicates nutrient
and/or energy limitations imposed by this
diet (see also Eisenberg, 1966; Beddiny,
1977). Interactive effects observed between
temperature and food quality indicate that
the combination of medium and high quality
foods and a temperature of 25°C produced
optimal conditions for growth.

In both medium and high quality food treat-
ments, reproductive output was continuous
over the duration of the life span. Rapid mat-
uration and constant reproduction when food
is not limiting has been considered a key life
history adaptation for organisms living in sea-
sonally unstable habitats (Pianka, 1970; Smith
& Fretwell, 1974; Wilbur et al., 1974; Giesel,
1976; Calow, 1978; Caswell, 1983; Brown,
1985). Given the unpredictable drying events
experienced by Physella cubensis and other
freshwater pulmonates, rapid maturation and
high fecundity for the duration of the life span
may be essential in maintaining an adequate
cohort of juveniles in the population capable
of aestivating. Food supply is clearly not
unlimited for many organisms, but may be
of reduced importance for many omnivorous
invertebrates. Periphyton, which encrusts
plants and rocks, and other organic and in-
organic debris has been suggested as pri-
mary food sources of many aquatic pulmo-
nates, including members of the Physidae
(DeWitt, 1955; Bovbjerg, 1968; Clampitt, 1970,
1973; Hunter, 1975; Lodge, 1985, 1986). Al-
though quantitative studies of natural diets
were not conducted, observations in the field
indicate that P. cubensis is a generalist grazer,
with a distribution, growth, and fecundity in
natural populations that may rarely be limited
by food quality or quantity. Instead, temper-
ature and the periodicity and magnitude of
precipitation events appear to be the most
important factors regulating the population
dynamics of P. cubensis in this central Ala-
bama stream and ephemeral pond habitat.

PHYSIOLOGICAL ECOLOGY OF PHYSELLA CUBENSIS 347

ACKNOWLEDGMENTS

We wish to thank Ginger Holt, Rebbecca
Seals, and Wendy Tipton, who assisted in
field data collection. Robert Angus provided
invaluable assistance in the statistical analy-
ses. This research was supported by the De-
partment of Biology, University of Alabama at
Birmingham.

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MALACOLOGIA, 1996, 37(2): 349-361

THE TAXONOMIC STATUS OF XEROAMANDA MONTEROSATO, 1892
(PULMONATA, HYGROMIIDAE)

Giuseppe Manganelli, Leonardo Favilli & Folco Giusti

Dipartimento di Biologia Evolutiva, Via P. A. Mattioli 4, 1-153100 Siena, Italy

ABSTRACT

The status of Xeroamanda Monterosato, 1892, is revised. Anatomical study showed that the
type species of this nominal genus, Helix amanda Rossmássler, 1838, is characterized by a
structure of the distal genitalia coinciding with that of the subfamily Hygromiinae sensu Schi-
leyko (dart sac complex with 0 + 2 stylophores) and more precisely with the group of genera
with 0 + 2 stylophores and right ommatophore retractor independent of the genitalia (Cernuella,
Xerosecta, Polloneriella, Microxeromagna, Xeromunda). The wide dart sac complex fused to the
walls of the vagina for a long tract and the penis joining the distal vagina level with the
stylophores enables it to be included in Cernuella. The latter has two subgenera: Cernuella $.
str. (penial papilla with three basal frenula, two tufts of digitiform glands on opposite sides of
vagina, proximal vagina very short or absent, proximal duct of bursa copulatrix flared), and
Xerocincta (penial papilla without frenula, digitiform glands around the vagina, proximal vagina
long, proximal duct of bursa copulatrix not flared). Xeroamanda (penial papilla with three basal
frenula, digitiform glands around the vagina, proximal vagina long, proximal duct of bursa
copulatrix not flared) is intermediate between Cernuella s. str. and Xerocincta and forms a third
distinct subgenus of Cernuella. At present, C. (Xeroamanda) includes two species, both en-
demic to the Sicilian area: the western Sicilian C. amanda (Rossmássler, 1838) and the Sicilian-

Maltese C. caruanae (Kobelt, 1888).

Key words: Hygromiidae, Cernuella, Xeroamanda, nomenclature, taxonomy, Sicily.

INTRODUCTION

Monterosato (1892: 22) established Xe-
roamanda, for a group of species from Tuni-
sia, Algeria, and Sicily, among which he cited:
Hlelix]. amanda and [Helix] usticensis. The
same year “Hel. amanda” was designated
type species in an anonymous review (1892:
151) of the Monterosato paper, usually attrib-
uted to Kobelt, editor of the Nachrichtsblatt
der Deutschen Malakozoologischen Gesell-
schaft.

Kobelt (1904) subsequently regarded Xera-
manda [sic] as a subgenus of Xerophila and
assigned thirteen species to it. Some years
later, Gude & Woodward (1921) regarded Xe-
roamanda as a junior synonym of Jacosta, one
of the subgenera in which they divided Heli-
copsis.

The taxonomy of Xeroamanda was more
recently considered by Sacchi (1955), Brandt
(1959), Zilch (1960), Forcart (1976), Manga-
nelli & Giusti (1988), and Nordsieck (1993).
Sacchi (1955) studied the genitalia of Alge-
rian specimens that he regarded as conspe-
cific with Sicilian Helix amanda, having been
unable to find live topotypical specimens in
the localities where the species had been re-

349

ported. He concluded that Xeroamanda must
be regarded as a separate subgenus of He-
licella. On the contrary, Brandt (1959), in his
revision of Cyrenaikan helicellids, regarded
the type species of Xeroamanda as a poly-
typical species comprising numerous sub-
species, including a new one described by
him from Cyrenaika. Because study of the
genitalia of this new species showed that it
belonged to Trochoidea, he considered Xe-
roamanda to be a subgenus of Trochoidea.
Zilch (1960) doubted Xeroamanda was a dis-
tinct subgenus of Trochoidea, regarding it as
a junior synonym of another subgenus of Tro-
choidea, Xeroplexa.

Later, Forcart (1976: 143) stated that the
drawing of the genitalia published by Sacchi
(1955: 4, fig. 1) showed clearly that Helix
amanda sensu Sacchi corresponded to Cer-
nuella. Nevertheless, he considered any con-
clusion premature, because neither Sacchi
(1955) nor Brandt (1959) had examined topo-
types of the type species.

Manganelli & Giusti (1988) examined a ju-
venile specimen of Helix usticensis, the other
species originally included in Xeroamanda by
Monterosato, finding a situation of the distal
genitalia externally similar to, but internally

350 MANGANELLI, РАМЕ & GIUSTI

different from, that of adult Cernuella. They
concluded that if their finding is confirmed in
adult specimens of the type species, Xe-
roamanda must be considered a separate
taxon. Nordsieck (1993) observed that the
features noted by Manganelli & Giusti (1988)
suggest affinity with the Tunisian and Maghe-
brian Xeroplana, which might turn out to be a
distinct Sicilian-Maghebrian genus.

Some years ago we asked some Sicilian
colleagues for living specimens of Helix
amanda, but they were unable to find any.
Fortunately, some specimens were found
among specimens of Cernuella rugosa (La-
marck, 1822) sent to us by Dr. V. E. Orlando
in 1987. This material was studied anatomi-
cally and led to the following revision.

MATERIALS AND METHODS

Whole shells were photographed under the
optical microscope (Wild M5A). All dimen-
sional parameters (shell height, maximum
shell diameter, aperture height and aperture
diameter) were measured using calipers.

Living specimens were drowned in water,
then fixed and preserved in 75% ethanol
buffered with СаСО.. The bodies were iso-
lated after crushing the shells and dissected
under the optical microscope (Wild M5A) us-
ing very thin, pointed watchmaker's forceps.
Anatomical details were drawn using a Wild
camera lucida. The dimensions of anatomical
tracts were measured using a graduated mil-
limetric lens on the same microscope.

Radulae were manually extracted from the
buccal bulbs, washed in pure 75% ethanol,
mounted on copper blocks with electron-
conductive glue, sputter-coated with gold,
and photographed using a Philips 505 SEM.

The material examined 1$ listed as follows:
locality, municipality and province names
in parenthesis, UTM reference, collector(s),
date, number of specimens in parenthesis.
Locality names and UTM references were ac-
cording to the official 1:50,000 scale map of
Italy (series M 792, sheet 593).

Explanation of symbols in Figures 3-9, 13-
23: BC, bursa copulatrix; BW, body wall; DBC,
duct of bursa copulatrix; DG, digitiform
glands; DP, distal penis; DSC, dart sac com-
plex; DGS, dart gun; E, epiphallus; F, flagel-
lum; FR, frenulum; FHD, first hermaphrodite
duct; FO, free oviduct; G, glans or penial pa-
pilla; GA, genital atrium; IS, inner stylophore;

LDL, left dorsal lobe; LLL, left lateral lobe; OS,
outer stylophore; P, penis; PN, pneumos-
tome; POS, prostatic portion of ovispermid-
uct; PP, proximal penis; PR, penial retractor
muscle; PV, proximal vagina; RCG, right ce-
rebral ganglion; RDL, right dorsal lobe; RLL,
right lateral lobe; ВОВ, right ommatophore
retractor; RPG, right pedal ganglion; SL, sub-
pneumostomal lobe; T, talon; UOS, uterine
portion of ovispermiduct; VD, vas deferens.

THE TYPE SPECIES OF XEROAMANDA

Cernuella amanda (Rossmássler, 1838)

Caracolla limbata Philippi, 1836: 137, pl. 8,
fig. 7. Type locality: “... prope Panor-
mus... ,” but incorrect according to
Calcara (in Benoit, 1862) and Benoit
(1862, 1882). Type series: holotype (Fig.
1) from “Panormus,” Philippi Collection,
Zoologisches Museum und Institut für
Spezielle Zoologie of the Humboldt-Uni-
versitat of Berlin, Germany.

Helix amanda Rossmássler, 1838: 10, pl. 32,
fig. 449. New name for Helix limbata
(Philippi, 1836), junior secondary hom-
onym of Helix limbata Draparnaud, 1801,
and permanently invalid according to
Art. 59 (b) of ICZN (1985).

Material Examined

“Panormus” (holotypus of Caracolla lim-
bata; Philippi Collection, Zoologisches Mu-
seum und Institut fur Spezielle Zoologie of
the Humboldt-Universitat of Berlin, Ger-
many); San Vito Lo Capo (San Vito Lo Capo,
Trapani), 33SUCO2, V. E. Orlando leg. 10.87
(3 shells + 2 spirit specimens); Mercato
Gnarosa (Custonaci, Trapani), 33STC91, 1.
Sparacio leg. 17.10.84 (1 shell); Semaforo
(San Vito Lo Capo, Trapani), 33SUC0028,
F.G. & G.M. leg. 7.10.94 (numerous shells).

Diagnosis

A species belonging to Cernuella, anatom-
ically similar to C. caruanae (Kobelt, 1888)
(penial papilla with three basal frenula; digiti-
form glands all around vagina; proximal
vagina long), but different by virtue of its
smaller, depressed, and carinate shell.

STATUS OF XEROAMANDA 351

Description
Shell (Figs. 1, 2)

Shell dextral, medium sized, robust, lentic-
ular, carinate, uniformly yellowish-grey or with
brown speckling and/or traces of a pale
brown band in upper half, and with traces of
5-6 bands, fragmented to form rows of small
brown spots in lower half, opaque; external
surface with fine ribs, densely packed, slightly
raised and rather irregularly positioned and
spaced; spire conical, depressed, consisting
of 5-5-1/2 clearly convex whorls, regularly
and slowly growing, last whorl large, barely
dilated, sometimes slightly descending near
aperture, with slightly raised, cord-like keel at
periphery; sutures moderately deep, shoul-
dered by keel of preceding whorls; umbilicus
open, rather small (about 1/6 of shell maxi-
mum diameter); aperture elliptical, its external
margin angled at keel; peristome interrupted,
simple, reflected only at columellar margin,
with upper margin sometimes starting at keel,
sometimes below keel, with well-developed,
yellowish, internal callous rib.

Dimensions of the holotype. Shell diame-
ter: 11.9 mm; shell height: 6.4 mm. Dimen-
sions of largest shell examined. Shell diame-
ter: 12.7-13.2 mm; shell height: 6.9-8.0 mm.

Body and General Anatomy (Fig. 3)

Data on colour of soft parts of living spec-
imens is not available. Retractor of right om-
matophore independent of penis and vagina;
kidney sigmurethrous; jaw odontognathous;
penial nerve from right pedal ganglion.

Genitalia (Figs. 4-9)

General scheme of semidiaulic monotrem-
atic type. Large hermaphrodite gonad (ovo-
testis) consisting of bunch of acini, ducts of
which converge into first hermaphrodite duct;
initial portion of first hemaphrodite duct very
slender, then widening to function as seminal
vesicle; first hermaphrodite duct ending in
club-like “talon” adhering to internal side of
large, bean-like albumen gland; talon consist-
ing of seminal receptacles (tree-like system of
tubules, ending in about three branches) and
fertilization chamber; second hermaphrodite
duct (ovispermiduct) arising from base of al-
bumen gland, and consisting of female chan-

nel (uterine portion of ovispermiduct contain-
ing seminal groove) and prostate gland (with
sperm groove) fused to define single cavity;
rather long free oviduct following female
channel; duct of bursa copulatrix arising from
where proximal vagina follows free oviduct,
long, slender (slightly shorter than epiphallus),
initially barely flared, ending in large, pyriform
bursa copulatrix (gametolytic gland); proximal
vagina proportionally long; digitiform glands,
far from apex of inner stylophore, subdivided
into two opposite groups, each of 3-4 units;
bases of units cover large portion of vagina
perimeter (Small portion of vagina perimeter is
free between the two groups); distal vagina
initiating from where dart-sac complex enters
one side of vagina; dart-sac complex 0 + 2, i.e.
consisting of one pair of stylophores, outer
fused to external side of inner for most of
its length; larger outer stylophore contain-
ing dart; smaller inner stylophore with small,
empty cavity; cavities of stylophores opening
independently one above other into groove
along side of conical structure named ‘‘dart-
gun” (Manganelli & Giusti, 1988); dart-gun
constituting basal appendix of dart-complex
projecting into distal vagina lumen; vaginal
pleats variable in number along internal sur-
face of vagina walls on both sides of dart-gun,
two of them fusing below dart-gun and giving
rise to sort of half-ring; vagina ending in gen-
ital atrium just after penis enters its far end;
long, slender vas deferens following sperm
groove (inside prostate gland of ovispermid-
uct) and ending in penial complex; penial
complex composed of flagellum, epiphallus
and penis; flagellum very short (about 1/9
epiphallus length), ending level with where vas
deferens enters penial complex and epiphal-
lus begins; epiphallus very long (about four
times penis length), ending where penial re-
tractor muscle contacts penial complex wall
and penis begins; penis short, entering distal
vagina level with apex of dart-gun and con-
taining penial papilla (glans); penial papilla
arising 1/3-1/4 of penis length from base,
long, slender, cylindrical, with apical opening
bordered by two-three “lips,” its base con-
nected to penial walls by three small, sym-
metrically disposed muscles (frenula); trans-
verse section of penial papilla with compact
walls with only few, small lacunae, and central
duct (ejaculatory duct) continuing directly
from proximal penis and epiphallus lumen;
penis opening into vagina bordered by sort of
annular pleat, possibly sphincter.

352 MANGANELLI, РАМЕ 8 GIUSTI

\
# НАХ »

A UNNE
Ue LES !

à 4
A

\ =

FIGS. 1, 2. Shells of Cernuella amanda (Rossmässler, 1838). 1, holotype of Caracolla limbata Philippi, 1836,
Philippi Collection, Zoologisches Museum und Institut für Spezielle Zoologie of Berlin, Germany. 2, a shell
from San Vito Lo Capo, Trapani, 33SUCO2, V. E. Orlando leg. 10.87.

STATUS OF XEROAMANDA 353

LDL

2mm

FIGS. 3, 4. Mantle edge (3) and talon (with transverse section) (4) of Cernuella amanda (Rossmässler, 1838)
from San Vito Lo Capo, Trapani, 33SUCO2, V. E. Orlando leg. 10.87.

Radula (Figs. 10-12)

Similar to that of other Cernuella species
(the radula of the various genera of the Hy-
gromiidae and Helicidae does not usually
have diagnostic characters), consisting of
many rows each of about 49-51 teeth; cen-
tral tooth with large tricuspid crown, meso-
cone long more than twice ectocone height;
first lateral teeth with bicuspid crown with
long, robust mesocone and small (about 1/2-
1/3 mesocone height) ectocone; last lateral
and latero-marginal teeth with bicuspid
crown, with long mesocone without small
protuberance or cusp on its inner side (only
exceptionally do a few lateromarginal teeth
have this small protuberance or cusp; in
other species, it is situated at about 2/3 of
mesocone height); extreme marginal teeth
with crown composed of very reduced me-
socone, its tip not split, and very reduced
ectocone, frequently split into two small,
sharp points.

Type Locality
Philippi claimed to have collected his sin-

gle specimen of Caracolla limbata in the
neighbourhood of Palermo: “Unicum speci-

men prope Panormus inveni.” According to
later students of Sicilian malacofauna, this lo-
cality was incorrect. The species was pre-
sumably collected by Calcara at Girgenti and
Catania in eastern Sicily and by Benoit at Ca-
latafimi in western Sicily (Calcara, reported in
Benoit, 1862: 188; Benoit, 1862: 188, 1882:
34-35). The presence of the species in the
localities reported by Calcara has not been
confirmed by subsequent research (Benoit,
1882: 34-35).

Type Series

The species was described from a single
shell collected by Philippi ‘ргоре Panormus”
(Philippi, 1836; Rossmássler, 1838). This shell
(Fig. 1) therefore has the status of holotype. lt
was illustrated by Philippi (1836: pl. 8, fig. 7)
and again by Rossmássler (1838: pl. 32, fig.
449) and is currently kept in the Philippi Col-
lection at the Zoologisches Museum of the
Humboldt-Universitát, Berlin, Germany.

Habitat

Xeroresistant species occurring in natural
habitats on calcareous substrata.

354 MANGANELLI, FAVILLI & GIUSTI

y ee

FIGS. 5-7. Genitalia (gonad to part of ovispermiduct excluded) (5, 6) and digitiform glands (7) of Cernuella
amanda (Rossmássler, 1838) from San Vito Lo Capo, Trapani, 33SUCO2, V. E. Orlando leg. 10.87. In the
specimen illustrated in Fig. 4, the outer stylophore has shifted and dart tip protrudes from the dart-sac
complex due to deformation in the course of sample preparation.

STATUS OF XEROAMANDA

355

FIGS. 8, 9. Details of distal genitalia of Cernuella amanda (Rossmássler, 1838) from San Vito Lo Capo,
Trapani, 33SUCO2, V. E. Orlando leg. 10.87. 8, distal penis opened to show penial papilla and frenula; 9,
vagina opened to show dart gun structure; penial papilla protrudes from distal penis.

Distribution

The species is endemic to Sicily and at
present is known from only one locality, San
Vito Lo Capo, north of Trapani. Other shells,
corresponding perfectly to those of San Vito
Lo Capo, were found in a nearby site, Mer-
cato Gnarosa, by |. Sparacio.

Nomenclature

Philippi (1836: 137, pl. 8, fig. 7) introduced
the nominal species Caracolla limbata for a
Sicilian xerophilous hygromiid described from
a single specimen collected near Palermo.
Two years later, when reporting Philippi’s
species in his /conographie, Rossmássler

356 MANGANELLI, РАМЕ & GIUSTI

FIGS. 10-12. Вааща of Сетие!а amanda (Ross-
mässler, 1838) from San Vito Lo Capo, Trapani,
33SUCO02, V. E. Orlando leg. 10.87.

(1838: 10, pl. 8, fig. 7) moved it to the genus
Helix and, to avoid secondary homonymy with
the older Helix limbata Draparnaud, 1801,
renamed Caracolla limbata, Helix amanda,
after the second personal name of Philippi,
Amandus.

This century, revision of the helicoids
caused splitting of the genus Helix into many
different genera. When Helix limbata Drapar-
naud, 1801, and Helix limbata (Philippi, 1836)
fell into two different genera, the secondary
homonymy between them ceased. This led

13

FIGS. 13, 14. Hygromia and Cernuella—group (in-
cluding Xeroamanda, Xerocincta, Cernuella and
the apomorphous Cernuellopsis) share many char-
acter states that except the structure of penial pa-
pilla derived from dart-sac complex, e.g.: 0 + 2
dart-sac complex (1), dart-sac complex adhering
to vagina (2), inner stylophore with thick, muscular
walls and narrow lumen (3), stylophores opening
independently of each other (4), dart-gun structure
(5).

Brandt (1959) to revive Philippi’s name over
Rossmássler's. The reintroduction of Philip-
pi's name passed unobserved (e.g., Alzona,
1971) or caused some confusion. In his Cat-
alogue of the species of Helicidae, Richard-
son (1980) lists amanda Rossmassler, 1838
(р. 246) and limbata Philippi, 1836 (р. 248), as
different species of Trochoidea!

The fate of junior secondary homonyms is
defined by Art. 59 (b) of ICZN (1985), which
states that “a junior secondary homonym re-
placed before 1961 is permanently invalid.”
Hence, the reintroduction of Philippi’s name
by Brandt (1959) is invalid.

Taxonomy

Pfeiffer (1853: 136) regarded the Sicilian
species as a junior synonym of the Algerian
Helix rozeti Michaud, 1833 (type locality: “еп-
tre Mostaganem et les marabouts de Mes-
rah’’). This synonymy was accepted by some
(Martens, in Albers, 1860; Kobelt, 1871;
Westerlund, 1876; Paulucci, 1878; Pfeiffer,
1878-1881; Tryon, 1887) and overlooked or
disregarded by others (H. Adams & A. Ad-
ams, 1855; Benoit, 1857, 1875, 1882; Bour-
guignat, 1864; Issel, 1880; Lallemant, 1881;
Westerlund, 1889; Pilsbry, 1895; Kobelt,
1904; Sacchi, 1995; Brandt, 1959; Alzona,
1971). Because no data is currently available
on topotypical Helix rozeti, this synonymy
seems completely speculative.

STATUS ОЕ XEROAMANDA 357

ROR RCG

KW NN o
у | \ %

с»

18 4

21

$ (

}

CL cl
Sas
+.

20
5
N FR i ps
V
OS ~ D Г
23

FIGS. 15-23. Character states used in cladistic analysis. For explanation, see Table 1.

At present, Cernuella amanda (Ross-
mässler, 1838) is endemic to Sicily and im-
mediately distinguishable from the species of
the two currently accepted subgenera of
Cernuella, Xerocincta and Cernuella s. str., by
virtue of features of the distal genitalia. It is
also easy to distinguish from the other spe-
cies belonging to Xeroamanda, the Sicilian-
Maltese Cernuella caruanae (Kobelt, 1888):
C. amanda has a small, depressed, carinate
shell (Figs. 1, 2), whereas C. caruanae has a
small-medium to medium, almost globular
shell (Kobelt, 1890: pl. 113, figs. 672-673;
1890: pl. 113, figs. 676, as Helix gattoi; Giusti
et al., 1995: figs. 553-557, 565). However,
because no substantial anatomical charac-

ters distinguish the two taxa, further research
is necessary to establish their relationship
(two distinct species or two subspecies of
the same species? the same species with a
variable shell?). This can only be done in the
framework of the revision of all the Sicilian
Cernuella, a difficult task because of the ex-
istence of more than 60 nominal specific taxa
(Favilli, 1994). The status of these taxa 1$
problematical and their revision is complex,
because it involves locating and studying of
the original descriptions, syntypes, topo-
types and obtaining much living material for
classical morphological (conchological and
anatomical) and genetic (allozyme polymor-
phism) studies.

358 MANGANELLI, РАМЕ & GIUSTI

TABLE 1. List of characters.

1—Retractor of right ommatophore (Fig. 15)
between penis and vagina = 0
independent of penis and vagina = 1
2—Penial nerve (Fig. 16)
from cerebral ganglion = 0
from pedal ganglion = 1
3—Joining of penis and vagina (Fig. 17)
atrial = 0
vaginal = 1
4—Frenula at base of penial papilla (Fig. 18)
absent = 0
present = 1
5—Proximal vagina (Fig. 19)
long = 0
very short or absent = 1
6—Digitiform glands (Fig. 20)
single units all around vagina or two barely distinct tufts, each consisting of various units on
opposite sides of vagina; small portion of vagina perimeter 15 left between the two tufts = 0
two distinct tufts, each consisting of a few units on opposite sides of vagina; large portion of
vagina perimeter 1$ free between the two tufts = 1
7—Proximal portion of bursa copulatrix duct (Fig. 21)

slender = 0
wide and flared = 1
8—Opening of stylophores (Fig. 22)
directly into vagina = 0
into a muff inside vagina = 1
9—Vagina and inner stylophore (Fig. 23)

Vagina adhering to side of inner stylophore opposite to that facing outer stylophore = 0
Vagina inserted in side of the inner stylophore facing outer stylophore = 1

THE STATUS OF XEROAMANDA

The revision of the many taxa of the genus
groups introduced by Monterosato (1892) for
the sections of Xerophila has posed serious
problems to everyone involved in the study of
these palaearctic helicoids in the last 100
years (for example, Pilsbry, 1895; Kobelt,
1904; Gude & Woodward, 1921; Hesse,
1926, 1934; Lindholm, 1927; Thiele, 1931,
Germain, 1929; Zilch, 1960). In fact, until the
old, conchological approach to gastropod
taxonomy was abandoned and replaced with
the more valid anatomical approach, revi-
sions were a matter of opinion and did not
produce natural classifications.

Unfortunately, the early approach to the re-
vision of Xeroamanda (Sacchi, 1955; Brandt,
1959) was vitiated by the analysis of non-
topotypical specimens (Algerian in the case
of Sacchi, 1955; Libyan in the case of Brandt,
1959), which did not resolve the problem and
produced two opposite interpretations of Xe-
roamanda, as Forcart (1976) realized: Xe-
roamanda as a distinct subgenus of Helicella
(Sacchi, 1955); Xeroamanda as a distinct
subgenus of Trochoidea (Brandt, 1959).

Research on the taxonomy and systemat-
ics of helicoids has gained new impetus
since the 1970s and 1980s thanks to Schi-
leyko’s study of the inner structure of the
distal genitalia (dart-sac complex and penis).
This work was innovative and produced
many revisions that permitted a new look at
the taxa involved in revision of Xeroamanda
(Schileyko, 1972, 1978, 1991; Hausdorf,
1988; Manganelli & Giusti, 1988, 1989; Giusti
et al., 1992, 1994; Nordsieck, 1993).

Anatomical study of the type species of
Xeroamanda showed that this taxon is char-
acterized by a distal genital structure that co-
incides with that of the subfamily Hygromii-
nae sensu Schileyko (dart зас complex with 0
+ 2 stylophores). The genus-group taxa of
the Hygromiinae sensu Schileyko with 0 + 2
stylophores and right ommatophore retractor
independent of the genitalia (Cernuella, Xero-
secta, Xeromagna, Polloneriella, Microxero-
magna, Xeromunda) were recently revised by
Hausdorf (1988) and Manganelli & Giusti
(1988, 1989). Xeroamanda is clearly charac-
terized by the structure of the distal genitalia
(dart sac complex wide and fused to inner
walls of vagina for a long tract, penis joining

STATUS OF XEROAMANDA 359

Hygromia

8(1) 9(1)

1(1) :2(1)

Cernuellopsis

Xerocincta

Xeroamanda

5(1) 6(1) 7(1)
Cernuella (s.str.)

FIG. 24. The most parsimonious phylogenetic hypothesis consistent with the data matrix showing evolution

of characters.

distal vagina level with stylophores), which
allowed it to be included in Cernuella. At
present, Cernuella includes two subgenera
Cernuella s. str. (penial papilla with three
basal frenula, two opposite tufts of digitiform
glands, proximal vagina very short or absent),
and Xerocincta (penial papilla without frenula,
digitiform glands around vagina, proximal va-
gina long). Xeroamanda 1$ intermediate be-
tween Cernuella s. str. and Xerocincta (pe-
nial papilla with three basal frenula, digitiform
glands around the vagina, proximal vagina
long) and comes to be a third distinct subge-
nus of Cernuella.

Table 1 lists the characters (Figs. 15-23)
considered for analyzing the relationships
between the Xeroamanda and the allied ge-
nus-group taxa Cernuellopsis Manganelli 8
Giusti, 1988 (type species: С. ghisottii Man-
ganelli & Giusti, 1988), Cernuella Schlúter,
1838 (type species Helix variabilis Drapar-
naud, 1801), and Xerocincta Monterosato,
1892 (type species: Helix neglecta Drapar-
naud, 1805).

Character polarity was determined by out-
group comparison, using the genus Hygromia
Risso, 1826 (type species: Helix cinctella
Draparnaud, 1805) as the outgroup. Hygro-
mia, Cernuella, Cernuellopsis, Xeroamanda
and Xerocincta share many character states
(structure of penial papilla, O + 2 dart-sac
complex, dart-sac complex adhering to va-
gina (not stalked as in Xerosecta), inner sty-
lophore with thick, muscular walls and narrow
lumen, stylophores opening independently of
each other, dart-gun structure) (Figs. 13, 14),
which form robust synapomorphies of the
Hygromia—Cernuella group (Manganelli 8
Giusti, 1988; Giusti et al., 1992). The Cernuella

TABLE 2. Taxa and data matrix.

VIANA gor 160) TOO
Hygromia 0207 2052052 07230520 220720
Cernuellopsis 1 1 0 0 0 0 0 1 1
Xerocincta о ооо
Хегоатапаа о 00
Cernuella IA IE OO

group is supported with respect to Hygromia
by two synapomorphies: penial nerve from
pedal ganglion and retractor of right ommato-
phore free from genitalia.

The data matrix is 100% consistent with
only one possible phylogenetic hypothesis
nine steps long (Fig. 24). The relationships
between the four genera of the Cernuella
group are interpreted as follows: Cernuellop-
sis (based оп 8 (1) and 9 (1)) is a sister group
of Xerocincta-Xeroamanda-Cernuella s. str.
(based on 3(1)); Xerocincta is a sister group of
Xeroamanda-Cernuella, s. str. (based on 4
(1).

Cernuella is therefore taken to be the
monophyletic group supported by the syn-
apomorphy 3(1) and including three subgen-
era: Xerocincta, Xeroamanda and Cernuella
s. str. Cernuella s. str. is a monophyletic
group based on three autoapomorphies:
5(1), 6(1) and 7 (1). The monophyly of Xe-
rocincta and Xeroamanda is not supported
by any autoapomorphies. In the case of Xe-
rocincta, which includes only one species,
there are no problems. On the contrary in the
case of Xeroamanda, which currently т-
cludes two possibly distinct species, para-
phyly cannot be excluded.

360 MANGANELLI,
ACKNOWLEDGMENTS

We thank Mr. L. Gamberucci and Mrs. A.
Daviddi for technical assistence, Mrs. H.
Ampt for linguistic revision, and Ignazio Spar-
acio and Vittorio E. Orlando, both of Palermo,
Italy, for material of С. amanda.

This research was supported by CNR,
MURST 40% and MURST 60% grants.

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Revised Ms. accepted 16 July 1995

MALACOLOGIA, 1996, 37(2): 363-373

SHELL MICROSTRUCTURE OF VESICOMYID CLAMS FROM VARIOUS
HYDROTHERMAL VENT AND COLD SEEP ENVIRONMENTS

Michael J. Kennish,' Antonieto S. Tan,” & Richard A. Lutz'

ABSTRACT

The shell structure of three deep-sea vesicomyid clams (i.e., Calyptogena magnifica, Calyp-
togena phaseoliformis, and Calyptogena c.f. pacifica) is characterized by scanning electron
microscopy (SEM). SEM examination of shell fracture sections of these vesicomyids reveals a
variety of shell microstructures in distinct arrangements. The shell of Calyptogena magnifica
consists of layers of planar spherulitic, fine complex crossed lamellar, cone complex crossed
lamellar, and irregular simple prismatic structure, and that of Calyptogena phaseoliformis 1$
comprised of layers of fine grained homogeneous, planar spherulitic, fine complex crossed
lamellar, irregular spherulitic prismatic, and irregular simple prismatic structure. In Calyptogena
c.f. pacifica, the shell contains layers of planar spherulitic, fine complex crossed lamellar,
vertical non-denticular composite prismatic, and irregular simple prismatic structure. While
cone complex crossed lamellar structure is only observed in Calyptogena magnifica, fine
grained homogeneous and irregular spherulitic prismatic structures only occur in Calyptogena
phaseoliformis. Vertical non-denticular composite prismatic structure is found exclusively in
Calyptogena c.f. pacifica.

Key words: vesicomyid clams, shell microstructure, deep-sea hydrothermal vents, cold-seep

environments.

INTRODUCTION

The genus Calyptogena Dall, 1891, occur-
ring from Eocene to the Recent, is a member
of the archibenthal infauna of the northern
Pacific (Bernard, 1974). Although Dall (1891)
initially placed the genus in the Carditidae,
Woodring (1938) later united the taxon with
the Vesicomyidae (a family instituted by Dall
& Simpson [1901]) based principally on its
shell hinge morphology and associated het-
erodont dentition (Woodring, 1938). In addi-
tion, the microstructure of the aragonitic
shells of Calyptogena differed from that of
the carditids, with the outer layer being
homogeneous in Calyptogena and crossed
lamellar with a system of tubulations in the
Carditidae (Oberling 4 Boss, 1970; Taylor
et al., 1973; Lutz & Rhoads, 1980a; Lutz,
1982). According to Oberling & Boss (1970),
the macroscopic characteristics of Calypto-
gena—chalky, ponderous, smooth, and whit-
ish valves—are also distinctly different from
the carditid traits of strong radial sculpture,
ventral crenulations, polished interior, and
brownish coloration.

Calyptogena spp. appear to be restricted
to deep-sea habitats (Allen, 1983). Living or

recently dead specimens of species within
the family have been collected or photo-
graphed from hydrothermal vents or cold
seeps in the eastern Pacific (Explorer Ridge,
Juan de Fuca Ridge, Gorda Ridge, Santa
Barbara Channel, Monterey [Ascension Fan]
Canyon, Guaymas Basin, Galapagos Rift and
21°N, 13°N, 11°N, 9-10°М, and 17-22°$
along the East Pacific Rise), Sea of Japan,
Gulf of Mexico (Florida Escarpment, Louisi-
ana Slope, and Alaminos Canyon), and Lau-
rentian Fan (Swinbanks, 1985; Turner, 1985).
In soft sediment, sulfide/hydrocarbon loca-
tions, all vesicomyids are infaunal, shallow
burrowers. However, C. magnifica has ac-
quired an epifaunal habit at vent sites on bare
volcanic rock where it lives, nestling in crev-
ices on the hard rock substratum, a habit that
appears to be unique among the vesicomy-
ids (Kennish & Lutz, 1992).

The systematics of the family Vesicomy-
idae (Calyptogena and Vesicomya) is beset
with problems owing to the paucity of sam-
ples thoroughly analyzed, their considerable
variability, insufficient anatomical data, and
the ill-defined boundaries of the numerous
families of heterodont bivalves with which
vesicomyids have been associated (Boss 4

‘Institute of Marine and Coastal Sciences, Rutgers University, New Brunswick, New Jersey 08903, USA
“Department of Biology, Worcester State College, Worcester, Massachusetts 01602, USA

364 KENNISH ЕТ AL.

Turner, 1980). Ambiguities in the classifica-
tion of vesicomyids are evident from past
efforts by various authors to place many gen-
era and species, now included in the Vesi-
comyidae, in other families (i.e., Arcticidae
[Cryprinidae], Carditidae, Kelliellidae, and
Veneridae). Although the dearth of speci-
mens, lack of adequate ontogenetic series,
and occurrence of many species from single
localities at great depths in the ocean have
hindered prior systematic work on the group
(Lamy, 1922; Odhner, 1960; Boss, 1969), col-
lections of relatively large numbers of speci-
mens from a wide range of sulfide-rich hab-
itats (e.g., hydrothermal vents, cold water

sulfide seeps, and the carcasses of decaying
whales) have revealed certain common ana-
tomical features (Kanno, 1971). The shell mi-
crostructures of the vesicomyids reported
here may provide additional information of
value in taxonomic differentiation of mem-
bers of the group.

MATERIALS AND METHODS

Ten Calyptogena magnifica specimens
ranging in length from 4.4 cm to 26.9 cm
were collected from hydrothermal vent fields

SHELL MICROSTRUCTURE OF VESICOMYID CLAMS 365

along the East Pacific Rise (6 specimens
from 21°N and 2 specimens from 9°N) (Alvin
Dives 1218, 1220, 1223, and 1227) and Gal-
apagos Rift (2 specimens) (Alvin Dive 2224)
and prepared for SEM examination by shell
fracturing and sectioning. Subsequent to
fracturing, specimens were sonicated in dis-
tilled water, dehydrated in 95% ethanol, air
dried, and coated with gold/palladium (prep-
aration 1) prior to observation in a Hitachi 450
SEM and Атгау 18301 SEM. Radial (perpen-
dicular to the shell margin), oblique (non-ver-
tical and non-horizontal relative to the plane
of the shell layer), and horizontal (parallel to
the plane of the shell layer) sections (Carter 8
Clark, 1985) were prepared by first embed-
ding specimens in epoxy 815 resin, and sec-
tioning, polishing, and etching them with 6%
HCL (preparation 2). After etching the pol-
ished sections, they were rinsed with distilled
water, dehydrated in 95% ethanol, air dried,
and coated with gold/palladium prior to ob-
servation in the SEM.

The same methods of sample prepara-
tion were employed on specimens of Calyp-
togena phaseoliformis and C. c.f. pacífica.
Eleven C. phaseoliformis specimens ranging
in length from 16.6 cm to 25.4 cm and six

specimens of C. c.f. pacifica ranging in length
from 2.8 cm to 3.7 cm were collected from
sulfide/methane cold seep sites in Monterey
Canyon (Alvin Dives 2286 and 2287) and the
Axial Seamount on the Juan de Fuca Ridge
(Alvin Dive 2426), respectively. Both frac-
tured and sectioned specimens were pre-
pared for SEM observation according to the
aforementioned methods.

Some Calyptogena phaseoliformis speci-
mens were prepared by first sectioning the
shells and then polishing and etching them
with 6% HCL. Subsequent to etching the pol-
ished sections, they were rinsed with distilled
water, dehydrated in 95% ethanol, air dried,
and coated with gold/palladium prior to ob-
servation in the SEM (preparation 3). These
shells were not embedded in epoxy 815 resin
prior to sectioning as in the case of prepara-
tions 1 and 2. Although shells prepared ac-
cording to preparation 3 were examined by
SEM, no photographs of their microstruc-
tures are presented here.

In addition to the methods of shell prepa-
ration outlined above, some shells were frac-
tured but not sonicated prior to observation
in the SEM. Others were treated with sodium
hypochlorite (bleached). However, most of

FIG. 1. Calyptogena magnifica. Radial section, preparation 2, 500x. Umbo towards the left. External shell
surface towards the top. Planar spherulitic structure (PS) grades into fine complex crossed lamellar struc-
ture at the transition (T) layer. Horizontal field width (HFW) = 142 um.

FIG. 2. Calyptogena magnifica. Horizontal section of planar spherulitic structure, preparation 2, 1000x. HF W
= 74 um.

FIG. 3. Calyptogena magnifica. Radial section, preparation 2, 500х. Same orientation as Fig. 1. Irregular
simple prismatic structure (SP) alternates with fine complex crossed lamellar structure (FCCL). HFW =
142 um.

FIG. 4. Calyptogena magnifica. Radial section, preparation 2, 2500x. Same orientation as Fig. 1. Transition
from fine complex crossed lamellar structure to cone complex crossed lamellar structure resembles fine
vertical non-denticular composite prismatic structure. HF = 29 um.

FIG. 5. Calyptogena magnifica. Vertical section, preparation 2, 400x. External shell surface towards the top
left corner. The cone complex crossed lamellar structure increases in width and starts to branch out
towards the inner surface of shell. HF = 178 um.

FIG. 6. Calyptogena magnifica. Radial section of cone complex crossed lamellar structure, preparation 2,
500x. Same orientation as Fig. 1. HFW = 142 um.

FIG. 7. Calyptogena magnifica. Horizontal section of cone complex crossed lamellar structure, preparation
2, 1000ж. HFW = 71 um.

FIG. 8. Calyptogena magnifica. Vertical section, preparation 1, 600х. Umbo towards the top right corner.
The adductor myostracum (AM) consisting of irregular simple prismatic structure is embedded within fine
complex crossed lamellar structure. HFW = 118 um.

FIG. 9. Calyptogena magnifica. Vertical section, preparation 1, 985x. Umbo towards the bottom right
corner. The pallial myostracum (PM) consisting of irregular simple prismatic structure is embedded within
cone complex crossed lamellar structure. HFW = 71 um.

366 KENNISH ET AL.

the shells were prepared according to prep-
arations 1, 2, and 3.

Summary of Methods for Preparing
Shell Samples

Preparation 1. Shell fractured, sonicated in
distilled water, dehydrated in 95% ethanol,
air dried, coated with gold/palladium.

Preparation 2. Shell embedded in resin, sec-
tioned, polished, etched with 6% hydrochlo-
ric acid, rinsed with water, dehydrated in

95% ethanol, air dried, coated with gold/pal-
ladium.

Preparation 3. Shell sectioned, polished,
etched with 6% hydrochloric acid, rinsed
with water, dehydrated in 95% ethanol, air
dried, coated with gold/palladium.

SHELL MICROSTRUCTURE

Calyptogena magnifica Boss & Turner, 1980

Oberling & Boss (1970) reported that the
shell of Са/урюдепа has a granular ec-
tostracum (outer shell layer) and a homoge-

FIGS. 10-18.

SHELL MICROSTRUCTURE OF VESICOMYID CLAMS 367

neous mesendostracum (middle shell layer)
with traces of complex structure. Lutz 4
Rhoads (1980a), Fatton 8 Roux (1981a), Lutz
(1982), and Lutz et al. (1988) identified three
primary microstructural layers plus myostra-
cal regions in the entirely aragonitic shell of
С. magnifica, including (1) an outer granular
layer; (2) a middle fine to irregular complex
crossed lamellar layer; and (3) an inner cone
complex crossed lamellar layer. The shell ex-
terior of larger individuals is often heavily cor-
roded, with the outer granular layer being ab-
sent dorsally due to dissolution (Lutz et al.,
1988). Growth lines present on the shell ex-
terior do not show either continuity on the
whole of the shell or any evidence of regular
periodicity (Fatton & Roux, 1981b). However,
the growth lines are concentrated near the
ventral margin of the shell and appear mor-
phologically similar to those of shallow water
bivalves, with V- or U-shaped indentations in
the shell microstructure.

Detailed observations on Calyptogenamag-

nifica specimens revealed the following shell
structures: (1) planar spherulitic (Figs. 1, 2);
(2) fine complex crossed lamellar (Figs. 3, 4);
and (3) cone complex crossed lamellar (Figs.
4-7). Planar spherulitic structure т С. mag-
nifica (Fig. 2) consists of horizontally flattened
spherulites with second-order subspherical
aggregations of laths which radiate more or
less equally in all directions from a central
point of origin. Individual planar spherulites
are quite variable in shape and measure ap-
proximately 1-5 um in width. They tend to
exhibit highly irregular contacts, yielding a
layer with no particular structural arrange-
ment. When eroded, planar spherulitic struc-
ture appears similar to coarse to fine homog-
enous structures with which it may be easily
confused.

Beneath a layer of planar spherulites, fine
complex crossed lamellar structure is ob-
served. A transition layer of fine microstruc-
ture typically separates the planar spherulitic
and fine complex crossed lamellar layers

FIG. 10. Calyptogena phaseoliformis. Vertical section, preparation 2, 600х. Umbo towards the left. Outer
shell surface towards the top. The outer surface of the planar spherulitic structure (PS) is fine grained
homogeneous structure (FGHS). HFW = 118 um.

FIG. 11. Calyptogena phaseoliformis. Vertical section, preparation 2, 1000x. Same orientation as Fig. 10.
Planar spherulitic structure (PS) overlies irregular spherulitic prismatic structure (ISP). HFW = 71 um.

FIG. 12. Calyptogena phaseoliformis. Horizontal section, preparation 2, 1500x. Umbo towards the left.
Anterior end towards the top. Plane of horizontal section through the broad face of planar spherulitic
structure parallel to the depositional surface. Five to six major rays radiate from the center toward the
periphery of the lenses. HFW = 49 um.

FIG. 13. Calyptogena phaseoliformis. Horizontal section, preparation 2, 4000x. Umbo towards the right.
Anterior end towards the bottom. Plane of horizontal section passing through the transition layer between
р!апаг spherulitic structure and fine complex crossed lamellar structure. The branched ends of the radiating
rays in a lens of planar spherulitic structure interdigitate with irregular laths of the neighboring units of fine
complex crossed lamellar structure. HFW = 18 um.

FIG. 14. Calyptogena phaseoliformis. Radial section, preparation 2, 2000x. Umbo towards the top right
corner. Inner shell surface towards the bottom right corner. The criss-crossing units of the fine complex
crossed lamellar structure can be arranged into tablets. The tablets are staggered irregularly to form a
discontinuous layer. HFW = 37 um.

FIG. 15. Calyptogena phaseoliformis. Vertical fracture, preparation 1, 500x. Same orientation as Fig. 10.
Shell layers from the top of micrograph are: (1) irregular spherulitic prismatic structure; (2) fine complex
crossed lamellar structure. HFW = 142 um.

FIG. 16. Calyptogena phaseoliformis. Horizontal section, preparation 2, 1500x. Umbo towards the right.
Anterior end towards the bottom. Cross section of irregular spherulitic prismatic structure. HFW = 47 um.

FIG. 17. Calyptogena phaseoliformis. Vertical section, preparation 2, 1500x. Same orientation as Fig. 14.
Adductor myostracum consisting of irregular simple prismatic structure is sandwiched between fine com-
plex crossed lamellar layers. HFW = 47um.

FIG. 18. Calyptogena phaseoliformis. Vertical fracture, preparation 1, 1000x. Same orientation as Fig. 14.
Pallial myostracum consisting of irregular simple prismatic structure is sandwiched between fine complex
crossed lamellar layers. HFW = 67 um.

368 KENNISH ЕТ AL.

(Fig. 3). The formation of fine complex
crossed lamellae is often interrupted by the
insertion of pallial or adductor myostracal
layers, and as a consequence, irregular pris-
matic structure is seen alternating with fine
complex crossed lamellar layers (Fig. 3). Pro-
ceeding toward the inner shell surface, one
can follow the development of cone complex
crossed lameller structure (Figs. 4-7).

The adductor and pallial myostraca consist
of irregular simple prismatic structure (Figs.
8, 9). The prism cross sections appear highly
variable in shape along their lengths (Carter,
1990). The adductor and pallial myostraca
are easily distinguished (except when the ad-

ductor myostracum splits) because the ad-
ductor myostracum is substantially thicker
than the pallial myostracum.

Calyptogena phaseoliformis Métivier,
Okutani 8 Ohta, 1986

The thin, elongate shell of Calyptogena
phaseoliformis consists of several character-
istic layers: (1) fine grained homogeneous
(Fig. 10) or planar spherulitic structure (Figs.
10-14); (2) irregular simple prismatic or irreg-
ular spherulitic prismatic structure (Figs. 11,
15, 16, 17, 18); and (3) fine complex crossed
lamellar structure (Figs. 14, 15). One or more

FIGS. 19-27.

SHELL MICROSTRUCTURE OF VESICOMYID CLAMS 369

of these layers may be absent from a given
region of the shell due to growth or dissolu-
tion effects. As in С. magnifica, the adductor
myostracum and pallial myostracum can be
traced down to the inner shell layer, which 15
composed of fine complex crossed lamellar
structure (Figs. 17, 18).

Calyptogena c.f. pacifica Dall, 1891'

Radial sections of polished and acid-
etched specimens of Calyptogena c.f. paci-
fica exhibit four shell layers. Proceeding from
the outer shell surface inwards, these layers
consist of: (1) planar spherulitic structure
(Figs. 19-22); (2) vertical non-denticular com-
posite prismatic structure (Figs. 22-24); and
(3) fine complex crossed lamellar structure
(Fig. 25). The planar spherulites in this spe-
cies measure about 1-3 um in width. As in C.
magnifica and C. phaseoliformis, the adduc-
tor myostracum and pallial myostracum can
be followed down to the fine complex
crossed lamellae comprising the inner shell

‘Morphological characteristics of this species are consis-
tent with assignment to Calyptogena pacifica. In light of
the large geographical separation and differences in water
depth between Axial Seamount (site from which speci-
mens were sampled; depth = 1,540 т) and the type-lo-
cality (Albatross Station 3077, off Dixon Entrance, Alaska,
in 585 m; see Boss 4 Turner, 1980), subsequent genetic
analyses may reveal differences between this species and
С. c.f. pacifica from the type-locality.

layer (Figs. 26, 27). This vesicomyid, having a
much smaller and thinner shell than that of C.
magnifica, is particularly susceptible to dis-
solution effects. Hence, many of the speci-
mens collected from hydrothermal vent fields
of the Axial Seamount (Juan de Fuca Ridge)
have pitted shells, especially in proximity to
the dorsum.

A characteristic feature of Calyptogena
magnifica and С. c.f. pacifica shells 1$ the
presence of stellate formations outside of the
pallial line along the depositional surface of
the shell margin undertucked by the perio-
stracum (Figs. 28-30). These formations ap-
pear as slender and rod-like or triangular
units in the shell microstructure. In C. phase-
oliformis, however, fine grained granules or
hexagonal formations occur on the deposi-
tional surface of the shell margin under-
tucked by the periostracum (Figs. 31-33). In
C. magnifica, with shell structure on the
depositional surface that is well preserved,
smooth surface stellate formations, granu-
lated surface stellate formations, and granu-
lated hexagonal formations can be delin-
eated (Figs. 34-36). These structures occur
sequentially from the shell margin to the pal-
lial line. Hexagonal structures may be formed
by the deposition of particles preferentially
on the angles of the rods that form the stel-
late structures (Figs. 34-36). The hexagonal
formations in C. phaseoliformis may repre-
sent a later stage of formation than the stel-

FIG. 19. Calyptogena c.f. pacifica. Radial section of planar spherulitic structure, preparation 2, 2500x.
External shell surface towards the top. HFW = 29 um.

FIG. 20. Calyptogena c.f. pacifica. Radial section of planar spherulitic structure, preparation 2, 5000x. Same
orientation as Figure 19. HFW = 14 um.

FIG. 21. Calyptogena c.f. pacifica. Horizontal section of planar spherulitic structure, preparation 2, 5000x.
HFW = 14 um.

FIG. 22. Calyptogena c.f. pacifica. Radial section, preparation 2, 2500x. Same orientation as Fig. 19. Planar
spherulitic structure (PS) overlies vertical non-denticular composite prismatic structure (VNDP). HFW =
29 um.

FIG. 23. Calyptogena c.f. pacifica. Radial section of “fine” vertical non-denticular composite prismatic
structure, preparation 2, 2500x. HFW = 29 um.

FIG. 24. Calyptogena c.f. pacifica. Horizontal section of “fine” vertical non-denticular composite prismatic
structure, preparation 2, 2500x. HFW = 29 um.

FIG. 25. Calyptogena c.f. pacifica. Radial section of fine complex crossed lamellar structure, preparation 2,
5000x. HFW = 14 um.

FIG. 26. Calyptogena c.f. pacifica. Radial fracture, preparation 1, 1500x. Umbo towards the right. The
adductor myostracum (AM) is embedded within fine complex crossed lamellar structure. HFW = 47 um.

FIG. 27. Calyptogena c.f. pacifica. Umbo towards the top right corner. Radial fracture, preparation 1, 1500x.
The pallial myostracum (РМ) is embedded within fine complex crossed lamellar structure. HFW = 47 um.

370 KENNISH ЕТ AL.

late structures from which they likely derive.
These stellate- hexagonal formations repre-
sent stages in the formation of planar spher-
ulitic structure.

DISCUSSION

The shell microstructures of Calyptogena
magnifica, С. phaseoliformis, and С. c.f. рас!-
fica share a number of common features, and
they have several differences. The shell of C.
magnifica contains prismatic, planar spheru-
litic, and various crossed structural types.
These structures differ from one region of the

shell to another owing to growth and disso-
lution effects. A complete sequence of shell
structures in this species is most likely en-
countered about midway between the umbo
and pallial line. One or more of these struc-
tures may be absent toward the dorsum
where dissolution effects are most рго-
nounced and toward the ventrum where in-
complete shell formation occurs ventral to
the pallial line. Fatton & Roux (1981b), exam-
ining the shell of a C. magnifica specimen
collected in a hydrothermal vent field of the
East Pacific Rise at 21°N, identified an onto-
genetic process in the clam leading from
granular to various crossed structural types

FIGS. 28-36.

SHELL MICROSTRUCTURE OF VESICOMYID CLAMS 371

which differ from one region of the shell to
another. They suggested that the various
structural types that develop in the shell of
this species seem to be related to growth
rate and environmental fluctuations.
Although fine complex crossed lamellar
structure, cone complex crossed lamellar
structure, and irregular simple prismat-
ic structure (myostracal regions) have been
identified previously in Calyptogena mag-
nifica (Fatton 8 Roux, 1981a; Lutz et al.,
1988), planar spherulitic and (formation of)
cone complex crossed lamellar structures
have never been documented in this species.
In addition, the cause of the repetitive struc-
tural changes first reported in C. magnifica by
Fatton & Roux (1981a) has not been ade-
quately addressed in the literature. Irregular
prismatic structure (myostraca) in C. mag-
nifica may be seen alternating with either fine
complex crossed lamellar or irregular com-
plex crossed lamellar layers (Fig. 3). These
repeated structural changes signify a change
in the physiological environment of shell dep-
osition, with the pallial myostracal layers
marking interruptions in the deposition of a
particular shell structure (Batten, 1984). Lutz
8 Rhoads (1980b) noted that myostracal in-
terruptions in the formation of the complex
crossed lamellar layer in the bivalve Arctica
islandica (Linnaeus, 1767) may develop dur-

ing aperiodic burrowing events when the or-
ganism respires anaerobically. During these
episodes, the energy requirements of the an-
imal are supplied by anaerobic glycolysis,
with the acidic end products of this metabo-
lism being neutralized by dissolution of the
shell (Lutz 4 Rhoads, 1977; Crenshaw, 1980).
This type of metabolic activity may also con-
trol the repetitive shell structures observed in
C. magnífica.

All of the shell structures identified in Ca-
lyptogena c.f. pacifica, except vertical non-
denticular composite prismatic structure,
also occur in C. magnifica. Irregular complex
crossed lamellar structure and cone complex
crossed lamellar structure, both present in
the shell of C. magnifica, have not been ob-
served in C. c.f. pacifica. Planar spherulitic
structure in C. c.f. pacifica compares favor-
ably with that in С. magnifica, although there
is a difference in the maximum width of the
spherulites in these two species. In this
structure, the spherulites are horizontally flat-
tened with concentrically arranged aggrega-
tions of laths radiating outward from a com-
mon center. The spherulites in C. c.f. pacifica
are variable in shape, have irregular contacts,
and range from 1-3 mu in width.

As in the shells of Calyptogena magnifica
and C. c.f. pacifica, the shell of C. phase-
oliformis contains planar spherulitic, fine

FIG. 28. Calyptogena magnifica. Preparation 1, 5000x. Stellate structures on the shell margin undertucked
by periostracum. The stellate structure consists of triangular units. HFW = 14 um.

FIG. 29. Calyptogena c.f. pacifica. Preparation 1, 1000x. Stellate formations on the depositional surface
undertucked by periostracum. HFW = 71 um.

FIG. 30. Calyptogena c.f. pacifica. Preparation 1, 5000x. Stellate formations on the depositional surface
undertucked by periostracum. HFW = 14 um.

FIG. 31. Calyptogena phaseoliformis. Preparation 1, 1000x. Fine grained homogeneous structure on the
depositional surface undertucked by periostracum. HFW = 74 um.

FIG. 32. Calyptogena phaseoliformis. Preparation 1, 1000x. Hexagonal tablets on depositional surface
undertucked by periostracum. HFW = 71 um.

FIG. 33. Calyptogena phaseoliformis. Preparation 1, 5000x. Hexagonal tablets on depositional surface
undertucked by periostracum. HFW = 14 um.

FIG. 34. Calyptogena magnifica. Preparation 1, 5000x. Stellate structures on the shell margin undertucked
by periostracum. Units of the stellate structure are slender and rod-like. Many stellate structures are
superimposed on one another. HFW = 14 um.

FIG. 35. Calyptogena magnifica. Preparation 1, 5000x. Particles begin to mask the stellate structures. HFW
= 14 um.

FIG. 36. Calyptogena magnifica. Preparation 1, 5000x. Between the shell margin and the pallial line,
particles have fully masked the stellate structures. The composite structure appears hexagonal. Compare
with hexagonal structures of Calyptogena phaseoliformis in Figs. 32 and 33. HFW = 14 um.

372 KENNISH ЕТ AL.

complex crossed lamellar, and irregular sim-
ple prismatic structures, although there are
differences in the relative thicknesses of these
structures among the three vesicomyids. Un-
like C. magnifica and C. c.f. pacifica, C. phase-
oliformis also exhibits fine grained homo-
geneous and irregular spherulitic prismatic
structures. The fine grained homogeneous
structure in C. phaseoliformis consists of an
aggregate of aragonitic granules which mea-
sure about 1-4 um т size. The granules have
irregular boundaries and variable shapes,
yielding a layer with no particular structural
arrangement.

The planar spherulitic structure in Calypto-
gena phaseoliformis, as in C. magnifica and
C. c.f. pacifica, is comprised of horizontally
flattened spherulites that appear similar to
coarse to fine grained homogeneous struc-
ture when eroded (Figs. 10-13). In C. phase-
oliformis, the planar spherulitic structure
is wider than that observed in C. magnifica
and C. c.f. pacifica. This structure is underlain
by fine complex crossed lamellar, irregular
spherulitic prismatic, and irregular simple
prismatic structures. The fine complex
crossed lamellar (Figs. 14, 15) and irregular
simple prismatic layers (Figs. 17, 18) in С.
phaseoliformis are structurally identical to
those described above in C. magnifica and C.
c.f. pacifica. Irregular spherulitic prismatic
structure, observed only in C. phaseoliformis
(Figs. 11, 15, 16), is composed of more or less
conical first-order prisms that, in some in-
stances, strongly interdigitate along their mu-
tual boundaries (Carter & Clark, 1985; Carter,
1990). Hence, it is readily differentiated from
planar spherulitic structure.

Stellate formations occur along the depo-
sitional surface of the shell margin of Calyp-
togena magnifica and C. c.f. pacifica (Figs.
28-30). These formations can be distin-
guished from the hexagonal formations ob-
served outside the pallial line along the shell
margin of C. phaseoliformis (Figs. 31-33).
The hexagonal structures may represent a
later stage of formation than the stellate
structures.

SUMMARY AND CONCLUSIONS

The shell structure of the vesicomyids Ca-
lyptogena magnifica, C. phaseoliformis, and
C. c.f. pacifica has been examined with a
scanning electron microscope. It was found
that the shell of C. magnifica consists of pla-

nar spherulitic, fine complex crossed lamel-
lar, cone complex crossed lamellar, and ir-
regular simple prismatic structures. The shell
of C. phaseoliformis is comprised of fine
grained homogeneous, planar spherulitic,
fine complex crossed lamellar, irregular
spherulitic prismatic, and irregular simple
prismatic structures. The shell of C. c.f. paci-
fica contains planar spherulitic, fine complex
crossed lamellar, vertical non-denticular
composite prismatic, and irregular simple
prismatic structures. Only C. magnifica pos-
sesses cone complex crossed lamellar struc-
ture and C. phaseoliformis, fine grained
homogeneous and irregular spherulitic pris-
matic structures. Vertical non-denticular
composite prismatic structure is present ex-
clusively in C. c.f. pacifica. Variation in the
relative thicknesses of the aforementioned
structures in these three species is substan-
tial, accounting for much of the difference
observed in the overall shell thicknesses of
the vesicomyids.

ACKNOWLEDGMENTS

This is New Jersey Agricultural Experiment
Station Publication No. D-32402-15-95 and
Contribution No. 95-17 of the Institute of Ma-
rine and Coastal Sciences, Rutgers Univer-
sity, supported by New Jersey State funds,
the Fisheries and Aquaculture Technology
Extension Center, and NSF Grants OCE 87-
16591, OCE 89-17311, OCE 92-17026 and
OCE 93-02205.

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SHELL MICROSTRUCTURE OF VESICOMYID CLAMS 373

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Revised Ms. accepted 12 July 1995

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MALACOLOGIA, 1996, 37(2): 375-442

THE EVOLUTIONARY RELATIONSHIPS OF CEPHALASPIDEA S.L. (GASTROPODA:

OPISTHOBRANCHIA): A PHYLOGENETIC ANALYSIS

Paula M. Mikkelsen'

ABSTRACT

Cephalaspid opisthobranchs, or ‘‘bubble-shells,’’ comprise a diverse group of snails com-
monly considered “transitional” between prosobranchs and “higher gastropods.” Compara-
tive morphological investigations at gross, light, and scanning electron microscopic levels,
involving 20 taxa of cephalaspids and related shelled opisthobranchs in 16 genera, produced
a data matrix of 47 new and modified-traditional characters. The results present the first
phylogenetic hypothesis for shelled opisthobranchs generated using parsimony-based cladis-
tic methods.

The preferred cladogram (length 117, ci 0.50, ri 0.70) has the following topology: (Outgroup
(Acteon, Gegania) (Hydatina ((Ringicula A, Ringicula B) ((Cylindrobulla (Ascobulla, Volvatella))
((Aplysia, Akera) ((Bulla (Haminoea, Smaragdinella)) (Cylichna (Retusa A, Retusa В) (Acteocina
(Scaphander (Philine A, Philine B)))))))))). Non-homoplastic or at least strong clade-supportive
characters were determined from external anatomy, mantle cavity, and digestive, nervous, and
reproductive systems. From the preferred tree topology, the Anaspidea and Sacoglossa (=
Ascoglossa) were confirmed as monophyletic groups, with Cylindrobulla as an unambiguous
member of the Sacoglossa. Traditional Cephalaspidea was split into two major clades: (a)
Acteon, Ringicula, and Hydatina, removed to the as-yet-unresolved, paraphyletic ‘‘architecti-
branchs” or “lower heterobranchs,” and (b) the remaining cephalaspids as the monophyletic
group Cephalaspidea s.s., in sister-group relationship with Anaspidea. Homoplasy was evident
in 25 characters, and significant in six, confirming the existence of ‘‘rampant parallelism” in
shelled opisthobranchs.

Tree topology suggested several evolutionary scenarios. (a) Formation of the gizzard (most
plesiomorphic in Anaspidea) involved the gizzard plates (many to three) and gizzard spines
(present in Anaspidea and Bulloidea, lost in Philinoidea). (b) The internal sperm-conducting duct
(“vas deferens””) is presumed homologous with the prosobranch external ciliated groove. A
second (novel) external groove, located laterally, developed in shelled opisthobranchs, initially
for egg transport, and co-occurs with the internal duct in Sacoglossa. The internal duct was lost
in Anaspidea and Cephalaspidea s.s., with the external groove assuming the task of sperm
transport. (c) Allosperm storage sacks include a proximal receptaculum seminis and distal
gametolytic gland, the latter probably formed from the prosobranch bursa copulatrix. The
“bursa copulatrix” of sacoglossans is probably secondary. Some of the “lower heterobranchs”’
may share a proximal ‘‘receptaculum apparatus,” with the receptaculum and gametolytic gland
in tandem arrangement. (d) A herbivorous diet is presumed plesiomorphic, with carnivory
evolving independently at least five times, associated with different suites of digestive system
characters.

Key words: Cephalaspidea, Opisthobranchia, Heterobranchia, systematics, cladistics, phy-
logeny, anatomy, characters.

INTRODUCTION

The virtual revolution occurring in gastro-
pod systematics has stemmed from a variety
of causes, including a profusion of new dis-
coveries and development of new techniques
(Haszprunar, 1988; Bieler, 1992). Paramount
among the latter is phylogenetic methodology
(= cladistics), now the almost universally ac-

cepted procedure for reconstructing interre-
lationships among taxa. The resultant insis-
tence upon monophyletic groups defined by
synapomorphies is now overturning tradi-
tional gastropod classification, reducing such
familiar groups as Prosobranchia and Meso-
gastropoda to the level of informally used
common names.

Among the newly recognized higher taxa is

"Harbor Branch Oceanographic Institution, 5600 U. $. 1 North, Ft. Pierce, Florida 34946, and Department of Biological
Sciences, Florida Institute of Technology, 150 W. University Boulevard, Melbourne, Florida 32901 U. S. A.
Present address: Department of Malacology, Delaware Museum of Natural History, P. O. Box 3937, Wilmington, Delaware

19807-0937 Ц. $. А.

376 MIKKELSEN

Heterobranchia (Haszprunar, 1985a; Ponder
8 Lindberg, 1992), now believed to be the
monophyletic sister group to Caenogas-
tropoda (Haszprunar, 1988; Lindberg & Pon-
der, 1991). Heterobranchia includes the
Opisthobranchia and Pulmonata (collectively
Euthyneura, or “higher gastropods’’) plus the
“lower heterobranchs” [= Heterostropha (in
part), Allogastropoda (in part); Haszprunar,
1988; Ponder 8 Warén, 1988; Bieler, 1992],
an enigmatic set of families (e.g., Pyramidel-
lidae, Architectonicidae, Mathildidae) with
mosaic sets of prosobranch and opistho-
branch characters (Robertson, 1974). This
last unresolved assemblage 1$ the subject of
much of the current research in gastropod
systematics.

Within the Heterobranchia, the Order Ceph-
alaspidea [= Bulloidea, Tectibranchiata (in
part), “bubble-shells””] occupies a tradition-
ally “basal” or “transitional” position be-
tween prosobranchs and “higher” opistho-
branchs (Boettger, 1955; Schmekel, 1985),
placing them in close proximity to the con-
troversial lower heterobranchs. Recognizing
that systematics of Cephalaspidea is based
upon anagenetic organizational grades and
phenetic similarities, several authors (Rud-
man, 1972c, d; Gosliner, 1992) have con-
cluded that Cephalaspidea is probably not
monophyletic, yet to date, no parsimony-
based cladistic analysis, presenting a com-
plete data matrix and suggesting an alterna-
tive classification, has appeared in the
literature.

In earlier reviews (Mikkelsen, 1993, 1994),
| surveyed all published phylograms and
the 49 most frequently-used characters for
the 31 traditional families in this order. | noted
a general lack of morphological definition for
the taxon as a whole and that 92% of the
characters traditionally employed are prob-
lematic under modern phylogenetic stan-
dards. | concluded that in order to effectively
apply cladistic methodology to this group,
a thorough re-evaluation of cephalaspid
morphology was necessary to generate an
improved set of taxonomically informative,
homologous characters. This work is the re-
sults of that study and presents the first cla-
distically generated, testable phylogeny for
cephalaspids and closely related shelled
opisthobranchs. Importantly, this provides a
solid morphology-based framework for fu-
ture work involving more refined techniques
at the anatomical, ultrastructural, and/or mo-
lecular levels.

MATERIALS AND METHODS
Taxa

The ingroup comprised 16 genera repre-
senting key families in Cephalaspidea and
other shelled opisthobranch groups with his-
torically close affinity. From present Ceph-
alaspidea, these included Acteon (Acteon-
idae), Ringicula (Ringiculidae), Hydatina
(Aplustridae), Scaphander (Cylichnidae, often
separated as Scaphandridae), Philine (Philin-
idae), Cylichna [Cylichnidae, included also be-
cause of the excellent anatomical work of
Lemche (1956)], Acteocina (Cylichnidae, often
separated as Acteocinidae), Bulla (Bullidae),
Haminoea (Haminoeidae), Зтагадате!а
(Haminoeidae, often separated as Smarag-
dinellidae), and Retusa (Retusidae). Three
taxa were chosen from the shelled members
of Sacoglossa (= Ascoglossa), all formerly in
Cephalaspidea: Volvatella (Volvatellidae), Cy-
lindrobulla (Cylindrobullidae), and Ascobulla
(Ascobullidae, often combined into Volvatel-
lidae). The Anaspidea were represented by
Akera (Akeridae, formerly in Cephalaspidea)
and Aplysia (Aplysiidae, representing tradi-
tional sea-hares).

Exemplar species (explained especially
well by Griswold, 1993) were chosen for the
various genera, selected on the basis of avail-
ability of adequate anatomical material and
literature data. Name-bearing type species
were used whenever possible; this is partic-
ularly pertinent in such studies as this, in
which the higher taxa being considered (i.e.,
families) have not themselves been recently
revised and cannot a priori be considered
monophyletic. Although the name-bearing
type might not exhibit the most plesiomorphic
dataset for its family (as currently defined),
such ап exemplar secures the validity of the
results regardless of subsequent redefinition
of families or genera, or reassignment of other
species. Strict adherence to the exemplar
method was suspended during the creation of
the present dataset in only three cases (Rin-
gicula, Philine, and Retusa) which showed
sufficient anatomical variation at the generic
level to require two ingroup members each,
designated A and В, with A representing the
primary species investigated and В repre-
senting variable character states present in
one or two alternate species. In all other cases
of variability within a genus or other closely
related group (fully explained below in the
Characters and Coding section), coding was

PHYLOGENETICS OF CEPHALASPIDEA 377

assigned according to the exemplar. The total
ingroup thus included 19 taxa.

One of the underlying assumptions in cla-
distics is that the ingroup being analyzed is a
monophyletic group, and this criterion con-
tributed to the choice of the outgroup. Al-
though the primary goal of this study was to
reevaluate Cephalaspidea [the problems of
which with monophyly were reviewed earlier
by Mikkelsen (1993, 1994)], because non-
(but former) cephalaspid taxa were included,
the ingroup under consideration here 1$ rather
the Opisthobranchia, albeitincomplete. Ques-
tions still exist about monophyly of the Opis-
thobranchia, specifically whether it is rather a
paraphyletic grade “leading to the pulmonate
level of organization” (Haszprunar, 1988:
426). However, the monophyly question can
be surmounted here if the ingroup can be
considered as the Euthyneura, which is pres-
ently thought to be monophyletic (Haszpru-
nar, 1988; Bieler, 1992). Thus, although Pul-
monata has been considered by many (e.g.,
Gosliner, 1994) as the sister group to Opistho-
branchia, and might be argued to be the ¡ideal
choice for outgroup selection, it was rejected
primarily because of these considerations.

A representative of the lower hetero-
branchs, unresolved but in presumed sister-
group relationship to the Euthyneura, was
considered the most appropriate outgroup
for this study. This group, unfortunately, pro-
vided few taxa for which adequate character
information was available in the literature or
could be supplemented through original
study. Many are also highly derived in at least
one anatomical respect. For example, Pyra-
midellidae was one possibility because pop-
ulations of several species were readily avail-
able in the area surrounding my laboratory.
However, pyramidellids have a highly spe-
cialized digestive tract (including lack of a
radula) associated with a parasitic lifestyle,
considered a disadvantage because diges-
tive characters were anticipated as important
in this study. In addition, pyramidellids are
derived in lacking a gill and in their euthyneu-
rous nervous system.

Members of other lower heterobranch taxa
are either known primarily from shells without
much anatomical data (e.g., Architectoni-
cidae) or are extremely minute and poorly
understood taxonomically (e.g., Rissoelli-
dae, Valvatidae, Cornirostridae). Although
typically deep-water, one species in the fam-
Ну Mathildidae, Gegania valkyrie Powell,
1940, has been relatively well studied (Climo,

1975; Haszprunar, 1985b; Bieler, 1988), and
the availability of preserved and sectioned
specimens made this the best choice for an
outgroup in this study. But because it was
based on an extant biological species, Gega-
nia was not the ideal outgroup in every char-
acter. Not all of Gegania’s character states
were plesiomorphic according to the ‘‘larger
outgroup” (= caenogastropods) (e.g., shape
of the eye lens coded 2; see below). In addi-
tion, the copulatory organ, which provided
most of the reproductive characters in the
analysis, is absent in G. valkyrie. For these
reasons, a hypothetical plesiomorphic (all-
zero) outgroup was used in a second analysis
that included Gegania. [This hypothetical all-
zero taxon was itself “imperfect”; for in-
stance, in the absence of a gizzard (character
17 = 0), all gizzard characters (characters 18-
21) must be coded as “not applicable.’’]

Taxon abbreviations in the text are as fol-
lows: Ac, Acteon; Ai, Acteocina; Ak, Akera;
Ap, Aplysia; As, Ascobulla; Bu, Bulla; Cb, Cy-
lindrobulla; Cy, Cylichna; Ge, Gegania; Ha,
Haminoea; Hy, Hydatina; Ph, Philine [PhA,
based on P. aperta (Linné, 1767); PhB, based
on P. falklandica Powell, 1954, and P. gibba
Strebel, 1908]; Re, Retusa [ReA, based on R.
obtusa (Montagu, 1803); ReB, based on R.
truncatula (Bruguiere, 1792) and Я. semisul-
cata Philippi, 1836]; Ri, Ringicula [RiA, based
оп Я. nitida Verrill, 1873; РВ, based on R.
buccinea (Brocchi, 1814) and R. conformis
Monterosato, 1875]; Sc, Scaphander; Sm,
Smaragdinella; Vo, Volvatella; 00, all-zero
outgroup. Sources of data (e.g., specimens
and literature consulted for each taxon) and
summarized anatomical descriptions are pre-
sented in Appendix 1.

Cited repositories, institutions, and other
sources of material are: CAS, California Acad-
emy of Sciences, San Francisco; DMNH, Del-
aware Museum of Natural History, Wilm-
ington; FIT, Florida Institute of Technology,
Melbourne; FMNH, Field Museum of Natural
History, Chicago, Illinois; HBOI, Harbor
Branch Oceanographic Institution, Ft. Pierce,
Florida; HBOM, Harbor Branch Oceano-
graphic Museum, HBOI; MNHN, Museum Ма-
tional d’Histoire Naturelle, Paris, France;
NMNZ, National Museum of New Zealand,
Wellington; NNM, Nationaal Naturhistorisch
Museum, Leiden, The Netherlands; PMM, col-
lections by the author; SMSLP, Smithsonian
Marine Station at Link Port, Ft. Pierce, Florida;
USNM, National Museum of Natural History,
Smithsonian Institution, Washington, D. C.;

378 MIKKELSEN

ZMUC, Zoological Museum, Copenhagen,
Denmark.

Morphology and Comparative Anatomy

Comparative methods began with surveys
of the extensive literature on opisthobranch
anatomy to provide suggestions for usable
characters. In most cases, these were veri-
fied and supplemented by original anatomi-
cal investigations involving gross dissection
of fresh and preserved material, light histol-
ogy, scanning electron microscopy, and life
history studies. Character selection placed
emphasis on the search for derived novel
features and on presumably non-homoplas-
tic characters.

Gross dissections employed a Wild-M5
dissecting microscope equipped with draw-
ing tube and ocular micrometer; standard
fine dissecting tools and differential tissue
stains were used. All organ systems, tradi-
tional and non-traditional, were considered
as sources of characters. Radulae, jaws, and
gizzard plates were extracted by dissolving
the surrounding soft tissue in an aqueous so-
lution of potassium or sodium hydroxide, or
sodium hypochlorite (household bleach), at
room temperature, followed by a rinse in dis-
tilled water and storage in 70% ethanol. For
light histology, radulae were permanently
mounted in Turtox CMC-9AF low-viscosity
stain-mountant tinted with acid fuchsin (Mas-
ters Chemical Co., Des Plaines, Illinois). Indi-
vidual organs (e.g., gills) were critical-point
dried for scanning electron microscopy
(SEM). Items thus prepared (plus shells) were
air dried, mounted on stubs either in water-
soluble white household glue or on double-
sided adhesive tape, sputter-coated with
gold-palladium (at НВОМ) or gold (at ЕММН),
and studied using a Zeiss Novascan-30 (at
HBOM) or Amray-1810 (at FMNH) scanning
electron microscope.

Histological study was required for verify-
ing gross dissections, especially in small
individuals, and for discerning tissue differ-
ences. Tissue fixation varied and was often
unknown in the case of museum material.
Originally collected specimens were fixed in
5% formalin or Bouin's fixative (Humason,
1962: 13). Labels with specimens embedded
by Lemche (ZMUC) noted use of formol
sublimate (saturated corrosive sublimate in
10% formalin; Guyer, 1936: 240), Sanfelice's
(Humason, 1962: 21), Bouin's, or Petrun-
kevitsch’s (1933, cupric-paranitrophenol) Яха-

tives. Removal of the shell was accomplished
through physical means and/or chemical de-
calcification utilizing preservation in Bouin’s
or subsequent treatment in a 1% solution of
ethylene diamine tetraacetic acid (EDTA, ad-
justed to pH 7.2, J. Voltzow, pers. comm.)
until decalcified. Standard 5-10-um (in rare
cases, 15-um) serial sections were produced
for entire paraffin-embedded individuals in
anteroposterior and/or lateral orientation.
Larger specimens (e.g., Hydatina) were cut
into several pieces prior to embedding, and
each piece was serially sectioned. Excep-
tionally large specimens (e.g., Aplysia) relied
on gross dissection supplemented by sec-
tions of selected tissues. Instead of routine
staining procedures, such as hematoxylin-
eosin, the more informative stains Alcian-
Blue/Periodic Acid/Schiff's (PAS) or Gomori’s
Green Trichrome (Vacca, 1985: 280), sensi-
tive to different mucosubstances in the re-
productive tract and mucus glands, were
employed (for full protocols, see Mikkelsen,
1994: appendix IID). The specimen of Gega-
nia valkyrie prepared and described by Hasz-
prunar (1985b; NMNZ M.36712) had been
sectioned at 7 um and stained with Azan.

A qualitative mineralogical test using Al-
izarin Red solution (Friedman, 1959) verified
calcification (e.g., the presence of calcite
and/or aragonite) of gizzard plates. Alizarin
Red solution was prepared by dissolving 0.1
g of Alizarin Red S in 100 ml of 0.2% cold
HCl, and the solution stored in a brown glass
bottle. Each gizzard plate sample was first
broken to expose interior layers for testing,
then soaked in the bright yellow test solution
for 4-5 тт. Tests were concluded by stop-
ping the reaction in a distilled water rinse. A
positive reaction stained the cut surface of
the plate bright red.

Phylogenetic Analysis

Characters and character states employed
in the final analysis (with notes on traditional
use and coding assigned to specific taxa) are
outlined in the Results. Characters used in
the dataset were numbered, beginning with
“0” in accordance with the requirements of
the analytical algorithm, Hennig86. Polarity
(plesiomorphic and apomorphic character
states) was assigned on the basis of the con-
dition in “the larger outgroup,” or a general-
ized caenogastropod, not in accordance with
the condition in the outgroup taxon. As men-

PHYLOGENETICS OF CEPHALASPIDEA 379

tioned earlier, this resulted in several cases in
which the outgroup was not coded 0, when
good evidence existed to believe that the
character state in Gegania is derived.

Within the data matrix, a “not applicable”
character was coded as а '“'n”; unknown char-
acter states were coded as “u.” This system
provides the reader with information, а|-
though the two are treated identically by the
algorithm. Autapomorphic characters (i.e.,
those for which the apomorphic state occurs
in only one taxon, e.g., radula absent in Re-
tusa only) were not included in the analysis,
because they are uninformative in this con-
text.

The data matrix was analyzed using Hen-
nig86 (Farris, 1988), version 1.5, on an IBM-
compatible (PC-DOS) 486-class personal
computer. Multistate characters were treated
as unordered. As mentioned previously, two
analyses were conducted, the first with Ge-
gania as outgroup, and the second with an
all-zero outgroup.

Neither dataset allowed the /e algorithm (=
implicit enumeration, guaranteed to find all
trees of minimal length), the most exhaustive
algorithm in Hennig86, to conclude within a
reasonable time frame. Even the most basic
version of this algorithm, ¡e-, guaranteed to
identify one tree of minimal length, did not
find solutions in over ten hours of run-time
per dataset. The most extensive algorithm
that could be successfully executed using ei-
ther dataset was the mhennig*/bb* combina-
tion, which constructs several shortest trees
by single passes through the data and then
applies extended branch-swapping to the re-
sults.

Because neither dataset allowed execu-
tion of the ie algorithm (and thus were not
guaranteed to have produced the shortest
possible trees), successive approximations
weighting (Farris, 1969) was employed as an
additional test. Except for this method, all
analyses utilized equally weighted characters.

The tree-manipulation program Clados ver.
1.1 (Nixon, 1991) was used for interpretation
of character state transitions on the trees.
Component ver. 2.0 (Page, 1993) facilitated
production of the final cladogram in publish-
able format.

From the results of the two mhennig*/bb*
analyses plus the successive weighting runs,
a “preferred tree” was constructed by exam-
ining variation in each section of the tree, and
selecting the topology which was best sup-
ported by character state changes.

RESULTS
Characters and Coding

A complete list of the 47 characters ap-
pears in Table 1. Thirty-six characters are
binary; 11 characters are coded as multi-
state. Within each of the following sections,
characters considered but not used are
first explained, followed by full discussion of
numbered, coded characters. Within each
“coding” list, a ‘‘?”’ indicates that some de-
gree of uncertainty remained, and assigned
coding was provisional.

Shell: The shell in opisthobranchs is a more
difficult structure to use taxonomically than in
most gastropods because it is generally re-
duced in size and thickness. It usually has
little color (except in Acteonidae and Aplus-
tridae) and sculpture (other than spiral
grooves), and is often completely lost. When
present, it is most diagnostic at the species
level.

In the present dataset, adult shells are
present for all taxa, therefore the traditional
character “shell present/absent” was not
pertinent. Taxa with species lacking shells as
adults are present within several opistho-
branch orders under consideration here, for
example, Bursatella (Anaspidea), and Elysia
(Sacoglossa).

The traditional character “shell thick/thin”
is relative to overall body size, which varies
with age or preservation and was therefore
considered too subjective to be reliably
coded. This can also vary with environmental
conditions or within a genus, for example, in
Acteocina with thin- and thick-shelled spe-
cies [A. atrata Mikkelsen & Mikkelsen, 1984,
and A. canaliculata (Say, 1826), respectively;
Mikkelsen & Mikkelsen, 1984].

As in the previous case, the traditional
character “shell reduced” has been linked to
body size and was also rejected as uncod-
able.

Protoconch type (i.e., many tapered whorls
vs. few bulbous whorls), reflecting larval
development (i.e., planktotrophic vs. leci-
thotrophic/direct-developing, respectively)
was not used, because this character com-
monly varies intragenerically among opistho-
branchs (see below, Larval Development).The
same is true of the position о the protoconch,
that is, whether the apex is exposed (e.Q.,
Acteon, Ringicula) or involute (e.g., Bulla,
Scaphander). Involution probably occurred

380 MIKKELSEN

TABLE 1. Characters and coding. M = character modified from traditional use; NEW = new character;
NON = traditionally used character for non-cephalaspids; U = character used unchanged from
traditional use.

. Shell internalization (U). O = external; 1 = internalized.
. Operculum (in adult) (U). O = present; 1 = absent.
. Parapodia (M). O = absent; 1 = present.
. Posterior foot (U). O = absent; 1 = present.
. Mantle cavity opening (M). O = anterior; 1 = lateral.
. Adductor muscle (NEW, NON). 0 = absent; 1 = present.
. Plicatidium (М). 0 = two-sided, suspended in mantle cavity; 1 = one-sided, fully attached to roof of
mantle cavity.
7. Ciliated strips (NEW). O = blunt at mantle edge; 1 = flexed at mantle edge only; 2 = exogyrous,
flexed at both ends to form an arch.
8. Pallial caecum (M). O = absent or short; 1 = long.
9. Jaws (U). O = present; 1 = absent.
10. Oral cuticle (NEW). O = smooth; 1 = with processes; 2 = thickened cuticular ring.
11. Descending limb/ascus (NEW, NON). O = absent; 1 = present.
12. Tooth size (NEW, NON). O = uniform throughout existing ribbon; 1 = increasing within existing
ribbon.
13. Rachidian tooth (М). 0 = rhomboid, with larger median cusp; 1 = rhomboid, bilobed cutting edge,
with median indentation; 2 = dagger-shaped; 3 = еопдаеа plate.
14. Enlarged sickle-shaped lateral teeth (M). O = not present (but other laterals/marginals present); 1 =
present.
15. Lateral/marginal teeth number (M). O = > 1, more than one form; 1 = > 1, identical in form; 2 = 1.
16. Pharyngeal pouches (NEW, NON). O = absent; 1 = present.
17. Esophageal gizzard with gizzard plates (M). O = absent; 1 = present.
18. Gizzard plate calcification (M). O = not calcified; 1 = calcified (as evidenced by positive Alizarin Red
S reaction).
19. Gizzard plate number (Ц). 0 = > 3; 1 = 3.
20. Gizzard plates tuberculate (M). O = not tuberculate; 1 = tuberculate.
21. Gizzard spines (NEW). O = present; 1 = absent.
22. Filter chamber (NEW, NON). 0 = absent; 1 = present.
23. Stomach (NEW). O = with pouch-like chamber; 1 = simple flow-through, without pouch-like
chamber.
24. Caecum extending from stomach (NEW, NON). 0 = absent; 1 = present.
25. Extent of intestinal typhlosole (NEW). 0 = partial; 1 = absent; 2 = entire.
26. Nerve ring location (U). 0 = prepharyngeal; 1 = postpharyngeal.
27. Cerebral/pleural ganglia (M). O = separate; 1 = fused.
28. Relative length of cerebral commissure (NEW). O = long; 1 = short (adjacent).
29. Relative length of pedal commissure (NEW). O = long; 1 = short (adjacent).
30. Position of left pallial ganglion (NEW). 0 = LA, fused or adjacent; 1 = L-A or L—A, separate.
31. Position of subesophageal ganglion (NEW). O = B-V; 1 = B migrated toward nerve ring; 2 = BV,
migrated toward visceral.
32. Position of supraesophageal ganglion (NEW). O = V-P; 1 = P migrated toward nerve ring; 2 = VP.
33. Position of right pallial ganglion (NEW). O = AL; 1 = A-L; 2 = PA, migrated toward supraesophageal.
34. Position of genital ganglion (NEW). O = off visceral ganglion; 1 = on visceral loop between V and P.
35. Eye direction (NEW). O = dorsolateral; 1 = ventrolateral.
36. Eye location (NEW). 0 = close to surface [S/C < 0.2]; 1 = midway between surface and nerve ring
[S/C = 0.2-0.5]; 2 = deeply embedded, near nerve ring [S/C > 0.5].
37. Eye lens (shape) (NEW). 0 = solid spherical or oblong-oval; 1 = solid irregular with hollows/anchors;
2 = hollow irregular.
38. Internal sperm duct (M). O = present; 1 = absent.
39. Lateral external ciliated groove (M). O = absent; 1 = present.
40. Copulatory organ (retractability) (U). O = nonretractile; 1 = retractile.
41. Autosperm storage bulb (U). O = absent; 1 = present.
42. Ejaculatory duct (M). O = absent; 1 = present and continuous with external ciliated groove; 2 =
present and continuous with internal sperm duct.
43. Receptaculum seminis (location) (M). O = proximal; 1 = distal.
44. Gametolytic gland (location) (NEW). O = distal; 1 = proximal.
45. Novel bursa copulatrix (NEW). O = absent; 1 = present.
46. Gonad acini (NEW). O = hermaphroditic acini; 1 = separate male and female acini.

O O1 BR © ND HO

PHYLOGENETICS ОЕ CEPHALASPIDEA 381

several times during opisthobranch evolution
(Gosliner, 1981), as evidenced by several gen-
era or families that contain members with ei-
ther exposed or involute apices (e.g., Hyda-
tina, Acteocina, Retusa).

A plate-like shell is known to occur in many
gastropods (e.g., Naticidae, тре, Crepid-
ula spp., plus Aplysia and Philine here), was
considered certainly homoplastic, and there-
fore was not coded.

Shell microstructure revealed no codable
pattern in the taxa under study. In the re-
duced shells of cephalaspids, shell layers are
uniformly cross-lamellar (Gosliner, 1994;
pers. obs.).

Shell mineralogy (aragonite or mixed ara-
gonite/calcite), included in the preliminary
analysis (Mikkelsen, 1994), was omitted from
the final analysis following discussions which
revealed that the underlying assumptions of
Feigl's Test (for aragonite; Friedman, 1959)
could be imprecise (C. Hedegaard, pers.
comm.).

One character was coded from the shell:

0. Shell Internalization. In most species in
the present dataset, the shell is external, as it
also is in caenogastropods. However, in sev-
eral taxa, it is covered by fused mantle tis-
sues, completely (Philine) or with a small
open foramen (Aplysia).

Coding: O = external (in all except follow-
ing); 1 = internalized (in Ph, Ap).

External Anatomy: External anatomy or gen-
eral body form, especially the cephalic shield,
foot, and parapodia, have been widely used
in cephalaspid systematics since the time of
Fischer (1883b). Because some of these fea-
tures are strongly associated with the bur-
rowing habit, and are present in other bur-
rowing snails (e.g., Oliva, Natica), many are
likely to be homoplastic.

The cephalic shield was once considered a
synapomorphy for the order Cephalaspidea.
This is no longer true since the reclassifica-
tion of such taxa as Akera (to Anaspidea) and
Cylindrobulla (to Sacoglossa). The cephalic
shield could only be defined as a shovel-
shaped head including posterior extensions
(= “processes” or “tentacles””) covering the
anterior shell and/or mantle opening. No spe-
cial innervation patterns could be deter-
mined, and this definition was too subjective
and dependent upon functional rather than
morphological criteria to be reliably used in a
cladistic analysis. Traditional consideration
of form of the cephalic processes referred

either to (a) species-specific shape (rounded,
pointed, cleft) or (b) the unique presence in
Ringicula of an inrolled extension forming a
siphon; neither were appropriate for use in
this dataset.

Hancock's organ is the primary chemosen-
sory organ in cephalaspids, located anteriorly
in the cephalo-pedal groove on either side of
the head. Prior to reclassification of such taxa
as Akera, it served as a well-recognized syn-
apomorphy for Cephalaspidea. It is usually
comprised of a series of vertical or oblique
plicae, with distinct anterior and posterior
portions of separate innervation (Edlinger,
19804).

Based on dissection and histological
studies, Edlinger (1980a, b) categorized Han-
cock’s organs among representative ceph-
alaspids and proposed the following evo-
lutionary transition: (a) irregularly folded
(Acteon), giving rise to (b) regularly folded
(Bulla), in turn giving rise to both (c) bipin-
nate (Haminoea) and (d) weakly folded
(Scaphander). The transition from (b) to (d)
was supported by the observation of a regu-
larly folded organ in juvenile Scaphander
(Edlinger, 1980b). The plesiomorphic condi-
tion of (a) (as well as the character state re-
lationships in general) was supported by well-
separated nerves leading to the lip organs
(N1) and anterior Hancock’s organ (N2), plus
the narrow base of the nerve leading to the
posterior Hancock’s organ (N3), both nerve
conditions as present in Hydrobia ulvae (Pen-
nant, 1777) (Caenogastropoda: Rissooidea:
Hydrobiidae).

Edlinger’s transformation series could not
be used in this dataset for three reasons. (a)
Coding of a strongly folded Hancock’s organ
(as in Bulla) and one that is weakly folded (as
in Philine) was hindered by the ontogenetic
shift seen in Scaphander (Edlinger, 1980b),
changes in external morphology through
preservation, and lack of clear character
state limits. (b) Edlinger’s (1980b) evidence
offered to derive the bipinnate Hancock’s or-
gan from the folded version is weak: Edlinger
(1980b) placed credence in Ev. Marcus & Er.
Marcus’ (1967b: figs. 13, 14) drawings of hor-
izontal pinnae in some Haminoea species
over slanted pinnae in others. This difference
seems more likely a matter of preservation
artifact, species variation, or artistic license,
rather than one of phylogenetic significance.
(c) Edlinger’s (1980b) judgment of Acteon as
most plesiomorphic in innervation of Han-
cock’s organ is also unsettled by the close

382 MIKKELSEN

similarity of that in Haminoea (Edlinger,
1980b: fig. 5).

Hancock's organ is extremely variable
throughout the group under consideration.
According to Edlinger (1980a), that in Acteon
consists of irregular folds only. In Bulla, it is
vertically folded like a single unit of the plica-
tidium-type gill, whereas in Hydatina, each
plica has a radial pattern of folds resembling
a ginkgo leaf. In Scaphander and Philine, the
organ 15 detectable as only a lateral bulge,
sometimes with weak oblique ridges. In Ak-
era, alcohol-preserved specimens showed
hardly any trace of Hancock's organ, but for-
malin-preserved specimens (not transferred
to alcohol) clearly showed a whitish oval
patch with vertical folds amidst an otherwise-
brown integument. This degree of variability,
plus apparent preservation artifacts, made
coding the external structure of the organ dif-
ficult and unreliable. The organ 1$ likewise dif-
ficult to analyze in light-histological material.
Ultrastructural studies seem necessary here
before morphology of the Hancock's organ
can be coded. Edlinger's (1980a, b) conclu-
sions were compared to the final result of the
analysis.

Simple presence/absence of the Han-
cock's organ was considered as a character,
evidenced histologically by the presence of
strong innervation of the cephalo-pedal
groove often coupled with dark-staining sen-
sory cells. If one assumes Bullock (1965) was
correct in regarding Aplysia’s rhinophores to
be homologous with Hancock’s organs (Sup-
ported by similar innervation), the organs are
present in the entire ingroup [unconfirmed in
Acteon, Ringicula, Volvatella, but cited for а!
but Ringicula by Ghiselin (1963), Baba (1966),
and Edlinger (1980a, b)]. This potential char-
acter thus became an autapomorphy of the
ingroup, and was discarded from the dataset
as uninformative.

Lip organs were studied by Edlinger
(1980a, b), but were difficult to code here for
the same reasons cited above for Hancock’s
organ. As in the latter case, Edlinger’s con-
clusions will be compared those of this anal-
ysis.

Three characters were coded from external
anatomy:

1. Operculum (in Adult). In opisthobranchs,
with general reduction of the shell as a pro-
tective mechanism, the operculum is dis-
carded in most species shortly after meta-
morphosis of the larva. Because this is loss
(rather than gain) of a character, it could con-

ceivably occur more than once in a ho-
moplastic pattern.

Shape of the operculum, when present,
does not vary substantially. Adult opercula in
Acteon and Retusa (Luque, 1983) are pau-
cispiral. Larval opercula are also paucispiral
and similar in appearance in all known spe-
cies in the present dataset: Acteon (Thomp-
son, 1976), Acteocina (pers. obs.), Aplysia
(Ostergaard, 1950). The larval operculum is
absent in Philine gibba (fide Seager, 1979).

An adult operculum is absent in Retusa ob-
tusa and most other species, but is present in
three species, R. truncatula (some popula-
tions only; Burn & Bell, 1974; Luque, 1983;
Mikkelsen, 1995), R. operculata Minichev,
1966 (Minichev, 1966), and R. chrysoma
Burn, in Burn & Bell, 1974 (Burn & Bell, 1974).
Retusa A is coded as absent; Retusa B is
coded as present.

Coding: 0 = present (in Ac, ReB, Ge, 00); 1
= absent (in remainder).

2. Parapodia. Although the presence or ab-
sence of parapodia could generally be deter-
mined (although not readily from preserved
specimens alone), the traditional categories
of “weak” and “strong” were considered too
subjective to be useful.

Coding: 0 = absent (in Ac, Cb, As, Vo, Ge,
00); 1 = present (in remainder).

3. Posterior Foot. In many species of ceph-
alaspids, the foot is shortened so that it does
not extend beneath the entire animal. Coding
of the relative length of the foot (= traditional
character “foot long/short’’) was considered
too subjective, too relative to overall body
size, too subject to variation due to age,
preservation, and other factors.

Some species of traditional cephalaspids
(e.g., Acteocina) possess a thickening of the
posterior edge of the mantle (= posterior pal-
lial lobe, infrapallial lobe) that functions as a
posterior foot. Although this is formed espe-
cially in species with a shortened foot, a per-
fect correlation does not exist; it is absent
from the shelled sacoglossans (with short-
ened foot), and present in Smaragdinella in
which the foot extends the full length of the
body. A section of the lobe often extends
dorsally over the posterior apex of the shell
(e.g., Haminoea).

An expansion of the posterior floor of the
mantle cavity is present in some taxa (e.g.,
Acteon, Ringicula, Hydatina), but in these
forms it is not thickened and is not used as an
accessory foot.

Coding: 0 = absent (in Ac, RiA, RiB, Hy, Cb,

PHYLOGENETICS OF CEPHALASPIDEA 383

As, Vo, Ap, Ge, 00); 1 = present (in Sc, PhA,
РВВ, Cy, Ai, Bu, Ha, Sm, ReA, ReB, АК).

Mantle Cavity: The osphradium is a chemo-
sensory organ in the mantle cavity adjacent to
the gill. In caenogastropods, it is a large and
often complex structure, for which fine struc-
ture has been judged taxonomically informa-
tive (Haszprunar, 1986). In cephalaspids, the
osphradium 1$ reduced in size to a small knob,
presumably due to development of a new
chemosensory structure, Hancock's organ
(see above). Edlinger (1980a, b) investigated
the osphradium in six cephalaspid genera and
suggested a pattern of phylogenetic reduc-
tion. However, in the present study, Edlinger’s
(1980b) supposedly taxon-specific morphol-
ogies could not be confirmed in histological
sections, although the osphradium and os-
phradial ganglion could be readily located by
tracing the osphradial nerve from the suprae-
sophageal ganglion. No reliable differences
could be discerned in this small organ. Hasz-
prunar (1986) suggested that external shape
can reflect ecological conditions or diet, and
indicated no codable differences in four taxa
of heterobranchs examined. Effective study
of osphradial structure in this group will
probably require ultrastructural techniques.
Edlinger’s conclusions will be compared to
the results of this analysis.

Five characters were coded from the man-
tle cavity.

4. Mantle Cavity Opening. The position of
the mantle cavity opening, either anterior or
lateral, reflects “detorsion” or posterior mi-
gration of the mantle skirt (Brace, 1977b). It is
usually imprecisely expressed in the pub-
lished literature, for example, “the mantle
cavity is mostly to the right, though some-
times anteriorly directed” (for Scaphan-
dridae; Boss, 1982: 1024). To define this
more exactly, diagrammatic mantle cavity
maps (Figs. 1-17) were drawn for each taxon
and divided into quadrants. Position of the
osphradium, anus, and one or both points at
which the mantle fuses with the head-foot
were indicated on each map. The effective
opening of the mantle cavity is delimited by
the osphradium and anus, often (but not al-
ways) adjacent to the mantle fusion points.
An “anterior” opening was defined as one
with the osphradium and anus in the two an-
terior quadrants (Figs. 1, 2). A “lateral” open-
ing was similarly defined, but with the os-
phradium and anus in the two right quadrants
(Figs. 3-17).

The mantle cavities of Acteon and Ringi-
cula have been described as anterior but
“twisted toward the right” (Fretter & Graham,
1954: 567; Fretter, 1960: 540), nevertheless
each fit the quadrant criteria for “anterior” as
coded here.

Coding: 0 = anterior (in Ac, МА, ВВ, Ge,
00); 1 = lateral (in remainder).

5. Adductor Muscle. Several species of tra-
ditional cephalaspids possess transverse ad-
ductor muscles together with a flexible shell.
These are considered to be relocated slips of
the columellar muscle (Morton, 1972), which
in typical snails acts in retracting the snail
into its shell. In the now-sacoglossan taxa
Cylindrobulla, Ascobulla, and Volvatella, a
single adductor muscle 15 located anteriorly
just ahead of the gill. Akera has both anterior
and posterior adductor muscles.

Coding: O = absent (in all except following);
1 = present (in Cb, As, Vo, Ak).

6. Plicatidium. The typical cephalaspid gill,
or plicatidium, was excellently figured by Per-
rier 8 Fischer (1911: fig. H; pl. 1, figs. 2, 3) and
later by Morton (1972: figs. 6a, b; who coined
the term) in both surface and cross-sectional
views. Although its status relative to the cae-
nogastropod ctenidium has been controver-
sial, Gosliner (1994) summarized the argu-
ments for and against homology of the two
structures; for purposes of the all-zero out-
group, here, the plicatidium is considered ho-
mologous with the (albeit highly modified)
ctenidium. The plicatidium 1$ composed of
two parallel laminae separated by a narrow
blood space (Fig. 22; as described by Perrier
8 Fischer 1911: 30), and thrown into a highly
convoluted, evenly ciliated surface (Figs. 18,
19) between efferent and afferent blood ves-
sels. It is attached by suspensory тет-
branes at both edges to the roof of the man-
tle cavity; the edge is mostly free but 1$ also
attached deeply within the mantle cavity.
Both surfaces of the gill are exposed in the
mantle cavity. In histological cross-section
(Fig. 20), this looks like an extremely long,
convoluted, single membrane, attached to
the mantle by one point. In some taxa (e.Q.,
Bulla, Hydatina), each “fold” or leaflet of the
gill is in turn convoluted, to the extent that
each leaflet looks like a plicatidium itself (Fig.
19); the plicatidial unit has multiplied along
the length of the gill. This “two-sided” plica-
dium is more similar in morphology to the
caenogastropod ctenidium, and this charac-
ter state was considered more plesiomor-
phic.

384 MIKKELSEN

FIGS. 1-17. Diagrammatic mantle cavity maps, showing line of mantle fusion, and position of osphradium
(O), anus (A), and ciliated strips (thick line). 1. Acteon [after Fretter, 1939; Minichev, 1967 (after Pelseneer,
1894)]. 2. Ringicula (after Minichev, 1967). 3. Hydatina (after Rudman, 1972a; pers. obs.). 4. Scaphander
(after Perrier & Fischer, 1911; Brace, 1977b). 5. Philine (after Brown, 1934; pers. obs.). 6. Cylichna (after
Lemche, 1956). 7. Acteocina (after Er. Marcus, 1958). 8. Bulla (after Er. Marcus, 1957). 9. Haminoea (after
Er. Marcus, 1958). 10. Smaradinella (after Rudman, 1972c). 11. Retusa (after Gosliner, 1978; pers. obs.). 12.
Cylindrobulla (after pers. obs.). 13. Ascobulla (after Ev. Marcus & Er. Marcus, 1956). 14. Volvatella (after
Baba, 1966; Haszprunar, 1985а). 15. Akera (after Brace, 1977b). 16. Aplysia (after Eales, 1921; pers. obs.).

17. Gegania (after Haszprunar, 19855).

In Cylindrobulla and the other shelled sa-
coglossans, the gill has the same bilamellar
structure, but is one-sided, attached by one
full side to the roof of the mantle cavity (Fig.
2A, D). This condition was assumed to be
more derived.

The traditional condition “gill reduced”
was avoided as being too subjective. Some
authors have used “д! plicate/non-plicate”
as a character within the Cephalaspidea.
Schmekel (1985) used pectinate to describe
an alternate condition, but did not explain
which possessed this morphology. Boss
(1982) used “pinnate” to describe the gill of
Gastropteron, a philinoidean taxon not in-
cluded here. These distinctions did not apply
to the present dataset.

Although Climo (1975) noted that a caeno-
gastropod-like ctenidium was absent in Ge-
gania, Haszprunar (1985b: fig. 13) found gill
filaments suspended from the roof of the

mantle cavity. Although Haszprunar denied
their homology with the plicatidium, personal
observations suggest the same bilaminate
histological structure. Their location near the
kidney and rectum further supports homol-
ogy with the plicatidium. The gill is one-sided,
with each filament directly attached to the
mantle roof as in Cylindrobulla and the other
shelled sacoglossans.

No gill filaments were detectable in either
Retusa obtusa or R. truncatula.

Coding: 0 = two-sided, suspended in man-
tle cavity (in Ac, МА, RiB, Hy, Sc, PhA, РВВ,
Cy, Ai, Bu, Ha, Sm, Ak, Ap, 00); 1 = one-
sided, fully attached to roof of mantle cavity
(in Cb, As, Vo, Ge); n (absent) in ReA, ReB.

7. Ciliated Strips. The dorsal and ventral
ciliated strips (= tracts, raphes, bands) in the
posterior mantle cavity increase the flow of
water in the vicinity of the gills and anus. Ac-
cording to Brace (1977b), their development

PHYLOGENETICS OF CEPHALASPIDEA 385

FIGS. 18-22. Plicatidium. 18. Full plicatidium; Cylindrobulla n. sp. (Bermuda, 8/1990), SEM. 19. Same as 18,
close-up of plicatidial units. 20. Two-sided plicatidium, Akera bullata (ZMUC), histological cross-section,
PAS stain. 21. One-sided plicatidium, Ascobulla ulla (Ft. Pierce Inlet, 8/1990), histological cross-section,
Gomori stain. 22. Plicatidial structure; same as 20. М = mantle; PL = plicatidium; SH = shell. Scales = 400

um (18), 200 um (20, 21), 100 um (19), 50 шт (22).

correlates with reduction in body size and re-
duction of the gill. The ciliated strips pass just
posterior to the anus, and are present in most
heterobranchs.

In most taxa considered here, the ciliated
strips begin bluntly at the mantle edge and
proceed obliquely posteriorly into the mantle
cavity, ending within the pallial caecum if
present (Fig. 1). In the case of this character,
because caenogastropods have no ciliated
strips (and the all-zero outgroup was coded
‘“n”), polarity was based on the outgroup Ge-
дата and on higher frequency of occurrence
in the ingroup. In other taxa (e.g., Cylichna),
the ciliated strips are flexed posteriorly at the
mantle edge (Но. 6); in a few (e.g., Hami-
noea), both ends of the strips are flexed,
forming an arch (Fig. 9). This latter format
was considered as a type of pallial caecum
(“exogyrous,” see below) by Perrier 8 Fis-
cher (1911).

Coding: O = blunt at mantle edge (in Ac,
RIA, РВ, Ну, PhA, PhB, Ai, Cb, As, Vo, Ak,
Ge); 1 = flexed at mantle edge only (in Sc, Cy,
ReA, ReB); 2 = exogyrous, flexed at both
ends to form an arch (in Bu, Ha, Sm); n (ab-
sent) in Ap, 00.

8. Pallial Caecum. The pallial саесит 1$

a terminal extension of the ciliated strips in
the mantle cavity. Morton (1972: 344) sup-
posed that the presence of a pallial caecum
correlated with inhabiting turbid waters or
burrowing into soft substrata, where it is used
as “a long flushing siphon circulating a clean
water current that keeps the pallial cavity
clear of inborne sediment.” Perrier & Fischer
(1911; summarized by and often credited to
Hoffmann, 1933) proposed three types of pal-
lial caecum: (a) free (in Acteon, Hydatina,
Scaphander), in the form of an elongated tube
enrolled within the shell whorls in parallel to
the visceral mass (Fig. 1); (b) adhering (in Ak-
era, Philine), united with the visceral mass and
enrolled as part of it [Fig. 15; not to be con-
fused with the long, free cloacal tentacle
present in Akera, a sensory structure that dif-
fers histologically (Perrier & Fischer, 1911));
and (с) exogyrous [= “fused” of Brace
(1977a); in Bulla, Haminoea], or “winding out-
side” the shell whorls, in a spiral plane entirely
within the mantle cavity and terminating at the
mantle edge (Fig. 8). Type (c) is more properly
considered a form of ciliated strip and 1$
treated here as such (character 7). Other spe-
cies (e.g., in Philine, Acteocina) have a very
short extension of the ciliated strip into a small

386 MIKKELSEN

triangle from the mantle cavity (Fig. 7). Al-
though a pallial caecum in Gegania was not
discussed by either Climo (1975) or Haszpru-
nar (1985b), the mantle's posteriormost ex-
tension is a small pocket in which the end of
the ciliated strips is situated (Haszprunar,
1985b: fig. 1; Fig. 17). This could qualify here
as a short pallial caecum, however, because
of this and other cases of subjective degree
associated with a short pallial caecum, both
absent and short were coded equally as 0. A
long caecum, either free or adherent, was
coded 1.

Coding: O = absent or short (in МА, ВВ,
PhA, PhB, Cy, Ai, Bu, Ha, Sm, ReA, ReB, Cb,
As, Vo, Ge); 1 = long (in Ac, Hy, Sc, Ak); n
(ciliated strips absent) in Ap, 00.

Digestive System: The digestive system of a
typical shelled opisthobranch includes chiti-
nous jaws, a muscular buccal mass contain-
ing the radula, an esophagus extending from
the buccal mass to the stomach [elaborated,
depending on the taxon, by esophageal di-
verticula, crop, muscular gizzard (containing
gizzard plates and spines), and filter chamber
(also with small plates)]. Food manipulation
and maceration typically occur within the
buccal mass and esophageal structures. The
stomach can be a simple flow-through tube
or can have an extensive posterior pouch or
caecum; the ducts to the digestive gland
open into the stomach to receive digestible
food particles. Following the stomach, a
winding intestine compacts fecal particles,
which are expelled into the mantle cavity.

The “hard parts” of the digestive system
(e.g., radula, jaws, gizzard plates) have
played major roles in traditional cephalaspid
systematics. Details of these features are
particularly useful at the generic level, but
can show remarkable variability at the spe-
cies level. A good example here is the radula
of Haminoea spp., in which overall radular
appearance is exceedingly similar although
the numbers of marginal and lateral teeth
vary from six to over 50 (Thompson, 1976),
with confirmed records of ontogenetic т-
creases (Ev. Marcus, 1976).

The traditional character “radula present/
absent” was not used here because Retusa
is the only taxon here lacking a radula; thus,
this becomes an autapomorphy for Retusa
and is not informative within the analysis.

Gastropod radulae are often categorized
into types (e.g., docoglossate, rhipidoglos-
sate, taenioglossate), reflective of the num-

bers and shapes of the teeth in each row.
Instead of using such terms, which have
never been precisely defined for opistho-
branchs, a number of radular features have
been individually coded (characters 13-15,
below).

Two types of salivary glands were cited for
Aplustridae (= Hydatinidae) by Thiele (1931)
and Boss (1982). The second “pair” is actu-
ally an unpaired, elongate tube called the oral
gland. Its presence in Hydatina alone defines
this character as an autapomorphy, not ap-
propriate for coding in this context.

Outpockets or diverticula of the anterior
esophagus (anterior to gizzard) are present
in a number of groups (e.g., Cylindrobulla,
Bulla, Haminoea), and can be single, paired,
or stalked. In some species, these pouches
are glandular, and in some not (Rudman,
1971b). Many caenogastropods also have
esophageal pouches of various types (Fretter
& Graham, 1994). Ghiselin (1966: 370) sug-
gested that “this diverticulum may be a prim-
itive trait which is retained in some herbivo-
rous forms,” but Gosliner (1994) considered
homology among the various forms un-
proven. Until more convincing evidence sup-
ports homology, this character cannot be
used cladistically.

The presence of a thin-walled crop in the
esophagus of shelled opisthobranchs was
considered plesiomorphic by Gosliner (1981),
who later (1994) suggested that its derived
form was the muscular gizzard (discussed
below). However, species in many genera
(e.g., Scaphander, Philine, Haminoea, Re-
tusa, Aplysia) with a gizzard have a distinct
crop preceding the gizzard, which could re-
quire rethinking of Gosliner's proposed ho-
mology. Because the appearance of the crop
varies with preservation and the amount of
food in the gut, it was difficult to confirm in
dissections and histological sections, and
was not coded here. Examination of series of
live animals are needed to adequately assess
this character.

Seventeen characters were coded from the
digestive system.

9. Jaws. Jaws (= mandibles) are here de-
fined as a pair of discrete cuticularized struc-
tures in the oral cavity composed of rod-like
structures (Fig. 3A, B). This differs from con-
tinuous cuticularized epithelium, which stains
similarly (dark pink in PAS) in histological
sections, but lacks rods. Jaws are further-
more inserted into a generative groove in the
oral cavity (where the rods are formed), which

PHYLOGENETICS OF CEPHALASPIDEA 387

is identifiable in histological cross-sections
(Fig. 25, GG); this was confirmed for all taxa
with rod-like elements except Aplysia (not
sectioned). It is not present for taxa with oral
cuticle only (character 10).

The traditional character “jaws armed”
could refer to two situations: (a) jaws with
rodlets, or (b) rodlets with terminal denticles.
Either case prevents the unmodified use of
this terminology. | have, first of all, redefined
“jaws” here so that the absence of rodlets
negates the presence of jaws; jaws without
rodlets is merely oral cuticle (character 10),
which can also be denticulate, but exists
without a generative groove and 1$ an inde-
pendent feature. Denticulation of the jaw el-
ements can be prominent or barely discern-
ible (Gosliner, 1994: figs. 12, 13). This could
not be coded due to subjectivity and be-
cause it also appears to cross generic lines;
for example, in Gosliner's (1994) excellent
scanning photomicrographs, the jaw ele-
ments of Bulla striata Bruguiere, 1792, and
Haminoea natalensis (Krauss, 1848) are
nearly identical, although those of B. striata
are slightly denticulate, whereas those of H.
solitaria (Say, 1821) are strongly denticulate.

Thompson et al. (1985) described and fig-
ured denticulate jaw elements in Ringicula
conformis. Similar structures were noted by
Bouchet (1975) in R. nitida, however, histo-
logical sections of this species showed that
these are actually processes of the cuticle
lining the oral cavity; rod-like structures and a
generative groove were not observed. Fretter
(1960) agreed with this interpretation, de-
scribing “cuticularized epithelium bearing
rows of sharp denticles” in R. buccinea. Rin-
gicula is therefore coded 1. The same is true
of Acteon, with denticulate oral cuticle (rec-
ognized by Fretter 1939; Fig. 26) that has of-
ten been termed “jaws” (Er. Marcus, 1958;
Ev. Marcus, 1974; Gosliner, 1994).

Coding: 0 = present (in Ну, Cy, Ai, Bu, Ha,
Sm, Ak, Ар, 00); 1 = absent (in Ac, РА, Sc,
PhA, ReA, ReB, Cb, As, Vo, Ge); u (unknown)
in АВ, РВВ.

10. Oral Синае. Cuticle occurs т the oral
cavity of most cephalaspids, including those
with jaws (Fig. 25). In some taxa, the cuticu-
larized epithelium is adorned with denticles
or processes (Fig. 26; as discussed under
character 9), or is noticeably thickened [Fig.
27; e.g., the “sphincter” or “inner labial disk”
of Sacoglossa and other taxa (Salvini-Pla-
wen, 1988: 327), and the “cuticularized ring’
of Umbraculacea (Willan, 1987: 225)].

See comments above (character 9) on the
“jaws” of Ringicula and Acteon. Retusa's
oral tube is lined by ciliated cells only, without
any trace of jaws or cuticle.

Coding: 0 = smooth (in Ну, Sc, PhA, Cy, Ai,
Bu, Ha, Sm, Ak, Ap, Ge, 00); 1 = with pro-
cesses (in Ac, RiA); 2 = thickened cuticular
ring (in Cb, As, Vo); u (unknown) in RiB, PhB;
n (absent) in ReA, ReB.

11. Descending Limb/Ascus. The orienta-
tion of the radula within the buccal mass dif-
fers among the gastropods under study. п
lateral view, the buccal mass of a typical snail
or cephalaspid (Fig. 28) has a wide, exposed
area of teeth in use (RF). Posteriorly, the rad-
ula “bends” where the ribbon enters the pos-
teroventral radular sack (RAS) where new
teeth are formed. Generally as old teeth are
worn, they drop off the anterior edge of the
radula and are shed.

From the same perspective, the sacoglos-
san buccal mass (Fig. 29) has the radular
sack (RS) most dorsal in position. A small
number of teeth (one to several) is in active
use (RF) at the “bend,” which 1$ nearest the
mouth. Instead of being shed, worn teeth are
retained in a ventral pouch called the ascus
(ASC). In this radula, the “elbow” marks the
teeth currently in use; newer, larger teeth
form the so-called ascending limb (Fig. 29:
AL) from the radular sack, and older, smaller
teeth form the descending limb (DL) from the
ascus. The radular sack and ascus are also
clearly distinguishable in histological cross-
section. The presence of a descending limb,
with an ascus, is one of several unique fea-
tures cited for the order Sacoglossa (e.g.,
Boss, 1982).

Coding: 0 = absent (in all but following); 1 =
present (in Cb, As, Vo).

12. Tooth Size. In most gastropods, tooth
size does not differ noticeably from row to
row in the extracted radular ribbon. However,
in sacoglossans, tooth size increases notice-
ably from younger to older portions of the
radula. Following reassignment to Saco-
glossa, Cylindrobulla is now the single ex-
ception to this rule.

Coding: 0 = uniform throughout existing
ribbon (in all except following); 1 = increasing
within existing ribbon (in As, Vo); n in Re (not
applicable; radula absent).

13. Rachidian Tooth. The rachidian, or cen-
tral, tooth exists in this dataset in several
general forms, including rhomboid with a
central larger cusp, bilobed with a minute
central cusp, or dagger-shaped. Bulla has a

388 MIKKELSEN

UT

FIGS. 23-27. Jaws and oral cuticle. 23. Complete jaw; Haminoea antillarum (PMM-933), SEM. 24. Same as
23, close-up of rod-like jaw elements. 25. Jaw and generative groove; Bulla striata (PMM-931), histological
cross-section, PAS stain. 26. Oral cuticle with processes; Acteon tornatilis (ZMUC), histological cross-
section, PAS stain. 27. Thickened oral cuticle; Ascobulla ulla (Ft. Pierce Inlet, 8/1990), histological cross-
section, Gomori stain. CUT = cuticle, GG = generative groove; J = jaw. Scales = 100 um (23, 25), 40 um

(27), 20 ит (26), 10 ит (24).

unique, plate-like rachidian, which was
coded separately. Several taxa (i.e., Hyda-
tina, Scaphander, Philine) have a “vestigial”
rachidian tooth:

The rachidian is absent in Ringicula nitida
(fide Bouchet, 1975; pers. obs.) and R. buc-
cinea (fide Fretter, 1960), the two species

upon which Ringicula A and B are here
based. However, R. conformis has the un-
usual formula 1.1.1, with a bilobed, smooth-
margined rachidian (Thompson et al., 1985).
Similar rachidians were seen in the radular
sack of an unidentified Ringicula from off
eastern Florida; no rachidians were noted in

PHYLOGENETICS OF CEPHALASPIDEA 389

BM RAS

29 RF

ESO

DL ASC

FIGS. 28-29. Diagrammatic buccal masses. Ante-
rior at left; dorsal at top. 28. Cephalaspid. 29. Sa-
coglossan (modified after Gascoigne, 1985: fig. 2).
AL = ascending limb; ESO = esophagus; MO =
mouth; RF = functional portion of radula; RAS =
radular sack.

the buccal mass portion of this radula, sug-
gesting that they are “caducous,” as in
Scaphander (below). Although Ringicula A
was coded “n” to reflect the absent rachid-
ian in R. nitida, Ringicula B was coded 1 to
account for the bilobed form seen in other
species.

In Hydatina physis, Rudman (1972a: 130,
fig. 9C) described the rachidian ‘when
present... [аз]... а small elongate plate with
а raised point at the anterior end.” Ev. Mar-
cus & Er. Marcus (1967b: 17, fig. 9B, as H.
vesicaria Lightfoot, 1786) described this as
“a tiny pointed, somewhat irregular, tooth.”
In H. velum (Gmelin, 1791), Eales (1938: 80)
noted that “оп the naked rachis is a very
minute flat рае.” The Hydatina radula
scanned during this study showed no rachid-
ian. Because of this level of variability, Hyda-
tina was coded “u” for this character.

“The central teeth of Scaphander are
known to be caducous” (Ev. Marcus & Er.
Marcus, 1967a: 602), falling off in the func-
tional radular rows and present only in the
radular sack. Because of this, Rudman (1978:
99) stated that “the central tooth [in
Scaphanaer] is a relic structure of little or no

functional importance.” Although descrip-
tions of the rachidian teeth of Scaphander
species often note the presence of a median
cusp [e.g., $. lignarius (Linné, 1758), fide Pils-
bry, 1885a: pl. 61, figs. 39, 40, and Rudman,
1978: fig. 2G; S. clavus Dall, 1889, fide Ev.
Marcus 8 Er. Marcus, 1967a: 602, fig. 2],
most modern descriptions show irregularly
shaped plates, but definitely lacking cusps
[e.g., S. lignarius, fide Thompson, 1976: fig.
63f; S. punctostriatus (Mighels, 1841), fide
Bouchet, 1975, and Thompson, 1976: fig.
659, h]. The latter suggestion was upheld by
scanned radular teeth of $. watsoni during
this study. The condition in Scaphander 1$
therefore undecided and is here coded “u.”

A rachidian tooth is entirely lacking in the
type species of Philine, P. aperta, therefore
Philine A is coded “n.” However, a nonden-
ticulate, “semi-circular, raised plate” is
present in P. falklandica and in P. gibba (fide
Odhner, 1926; Rudman, 1972b), which ac-
cording to Rudman (1972b: 173) is the plesi-
omorphic condition for the genus. Odhner's
figure (1926: fig. 12) of the radula of P. gibba
shows a rectangular plate, without a median
indentation but evenly denticulate, therefore
also without a distinctly larger median cusp.
Rudman’s drawing (1972b: fig. 2a; repro-
duced in Rudman, 1978: fig. 2C) from the
same species is a semicircular nondenticu-
late plate. Although a median indentation is
clearly not indicated, the absence of any
cusps leaves the condition in this taxon un-
decided, therefore Philine B is coded “u”
here.

Coding: 0 = rhomboid, with larger median
cusp (in Ha, Sm, Cb, Ak, Ap, Ge, 00); 1 =
rhomboid, bilobed cutting edge, with median
indentation (in RiB, Cy, Ai); 2 = dagger-
shaped (in As, Vo); 3 = elongated plate (in
Bu); u in Hy, Sc, PhB (vestigial without defin-
able form); n in Ac, RiA, PhA (rachidian ab-
sent) and in ReA, ReB (radula absent).

14. Enlarged Sickle-shaped Lateral Teeth.
Many taxa in this analysis have hook- or
cusp-shaped lateral and marginal teeth. The
enlarged sickle-shaped tooth is here defined
as a robust hook (often marginally denticu-
late) on a handle-like base (as in Acteocina
spp., see Mikkelsen & Mikkelsen, 1984: fig.
3D). This morphology is frequently associ-
ated with a bilobed, denticulate rachidian.

Both the sickle-shaped teeth and the bi-
lobed rachidian (when present) are smooth in
Ringicula (Bouchet, 1975; Thompson et al.,
1985; pers. obs.).

390 MIKKELSEN

Coding: 0 = not present (but other laterals/
marginals present; in Ac, Hy, Bu, Ha, Sm, Ak,
Ар, Ge, 00); 1 = present (in RiA, РВ, Sc, PhA,
РВВ, Cy, Ai); п (absent) in Cb, As, Vo (laterals
absent) and in ReA, ReB (radula absent).

15. Lateral/Marginal Teeth Number. Lateral
teeth are generally larger and more robust
than marginals, but these labels suffer from
lack of any further identifying criteria. In cer-
tain species, the distinction can be difficult to
resolve. In Akera and Haminoea, for example,
all non-rachidian teeth are hook-shaped,
gradually becoming more slender and less
robust outwardly. The innermost 1-6 teeth
are usually denticulate, and although slight,
this is sufficient to distinguish laterals from
non-denticulate marginals. Because of these
vague definitions, however, a character such
as “marginals present/absent” was not used
in this analysis. Gosliner (1994) concurred,
calling the indistinct outermost teeth of
opisthobranchs “outer laterals.” Character
states O and 1 indicate the presence/ab-
sence of different teeth outside of the rachid-
ian, but are independent of the ambiguous
marginal tooth category. Character state 1,
where all non-rachidian teeth are essentially
identical (and usually minute), is exemplified
by the radulae of Acteon and Hydatina spp.
(Gosliner, 1994: fig. 19C, D).

Although the type species of Philine, P. ap-
erta, possesses only a single lateral tooth
(coded 2), other species (e.g., P. gibba, fide
Odhner, 1926: fig 12) have additional ‘‘mar-
ginal” teeth, usually cusp-shaped but smaller
than the “laterals.” Philine В is therefore
coded 0.

Coding: 0 = > 1, more than one form (in
PhB, Cy, Bu, Ha, Sm, Ak, Ap, Ge, 00); 1=>
1, identical in form (in Ac, Hy); 2 = 1 (in RiA,
RiB, Sc, PhA, Ai); n (not applicable) in Cb, As,
Vo (laterals/marginals absent) and in ReA,
ReB (radula absent).

16. Pharyngeal Pouches. Pharyngeal
pouches are lateral outpockets of the central
cavity of the buccal mass, and are character-
istic of sacoglossans. Their comparable ap-
pearance in histological cross-sections of
Cylindrobulla and Ascobulla spp. is one of
the deciding factors in considering the former
taxon a sacoglossan in this study.

Climo (1975) described a bifurcate “Бисса!
pouch” in Gegania valkyrie, but this is prob-
ably not homologous with the pharyngeal
pouches of sacoglossans. That found in Ge-
gania is a caecum originating ventral to the
junction of the buccal mass and esophagus,

and is therefore more like an esophageal di-
verticulum (see below).

Coding: 0 = absent (in all except following);
1 = present (in Cb, As, Vo).

17. Esophageal Gizzard with Gizzard
Plates. The muscular gizzard, always with a
series of gizzard plates well anchored within
the musculature, is one of the most recog-
nizable features of a typical cephalaspid. The
traditional character “weakly/strongly devel-
oped” was interpreted to refer to corneous
versus calcified gizzard plates, coded in
character 18. “Gizzard plates present/ab-
sent” applies to this character, because a
gizzard 1$ never present without plates, and
vice versa.

Because the all-zero outgroup was coded
here as 0 (gizzard absent), characters 18-21,
which involved gizzard plates and spines,
were coded “n” (not applicable) in this hypo-
thetical taxon.

Coding: 0 = absent (in Ac, RiA, RiB, Hy, Cb,
As, Vo, Ge, 00); 1 = present (in remainder).

18. Gizzard Plate Calcification. Gizzard
plates were tested mineralogically to verify
“calcification,” as reported in the literature
for species of the Philinoidea, for example,
Scaphanaer, Philine, Acteocina, and Cy-
lichna (see generalized descriptions in, e.g.,
Boss, 1982). Scaphander, Philine, and
Acteocina tested positively in Alizarin Red
solution, indicating calcification. The gizzard
plates of Cylichna cylindracea tested nega-
tively, disputing previous reports. One sam-
ple tested partly negatively: the gizzard
plates of Scaphander watsoni tested posi-
tively in the portion embedded within the
gizzard wall, but were “capped” by a layer
of unreactive material (three trials). The latter
could be weddellite, a calcium oxalate hy-
drate, determined (by electron-probe micro-
analysis) in the gizzard plates of Scaphander
cylindrellus Dall, 1908 (Lowenstam, 1968).

Gizzard plates in Philine aperta are calci-
fied according to the mineralogical test,
therefore Philine A was coded 1. However,
some species (e.g., P. gibba, P. falklandica)
have been reported to have uncalcified, cor-
neous gizzard plates (Odhner, 1926; Rud-
man, 1972b), a character state considered
plesiomorphic for the genus by Rudman
(19725). Philine В was provisionally coded 0,
however, in view of the conflicting results of
the Cylichna gizzard plate test versus litera-
ture data, this requires verification.

As a control for the gizzard plate samples,
the “corneous” gizzard plates of Bulla striata

PHYLOGENETICS OF CEPHALASPIDEA 391

were tested with Alizarin Red. Results were
negative. All other reportedly corneous giz-
zard plates (in Akera, Aplysia, Haminoea,
Smaragdinella, and Retusa) tested negatively
in Alizarin Red.

Coding: 0 = not calcified (in PhB?, Cy, Bu,
Ha, Sm, ReA, ReB, Ak, Ap); 1 = calcified (in
Sc, PhA, Ai); n (gizzard absent) in Ac, RiA,
АВ, Ну, Cb, As, Vo, Ge, 00.

19. Gizzard Plate Number. The number of
gizzard plates is three in most species of
cephalaspids. In anaspids (Akera, Aplysia),
10-20 small pyramidal gizzard plates occur.
Runcina (Runcinidae; Runcinoidea), a taxon
not included here, has four gizzard plates, a
feature contributing to its placement in its
own order of opisthobranchs by many au-
thors (Mikkelsen, 1993: fig. 2; 1994: table 2).

Determination of polarity of this character,
in the absence of a gizzard in both Gegania
and the all-zero outgroup, was assigned ac-
cording to Gosliner (1994), who suggested
that the presence of numerous plates was
more plesiomorphic.

Coding: 0 = > 3 (in Ak, Ap); 1 = 3 (in Sc,
PhA, РВВ, Cy, Ai, Bu, Ha, Sm, ReA, ReB); п
(gizzard absent) in Ac, НА, RiB, Hy, Cb, As,
Vo, Ge, 00.

20. Gizzard Plates Tuberculate. Gizzard
plates exist in many sculptural forms, includ-
ing smooth, ridged, laterally pinched, and
tuberculate. Most of these were not treated
in this analysis, but tuberculate gizzard plates
were coded, in part, with the goal of unifying
the two species of Retusa, which in early
trials increased the number of possible
trees dramatically through their independent
movements.

The tubercles on the uncalcified gizzard
plates of Retusa spp. (Figs. 30, 31) are black
in fresh or preserved material, and stain yel-
lowish rather than purplish in PAS. In cross-
section (Fig. 31), these extend from the base
to the surface of the plate. The color suggests
different composition from the rest of the
plate; the yellowish hue is similar to that of the
outer tests of foraminiferan prey seen in the
gut. Other heavily sculptured gizzard plates
(e.g., in Haminoea) did not show these kinds
of staining differences. This character is prob-
ably a synapomorphy for the genus Retusa.

Coding: O = not tuberculate (in Sc, PhA,
PhB, Cy, Ai, Bu, Ha, Sm, Ak, Ap); 1 = tuber-
culate (in ReA, ReB); n (gizzard absent) in Ac,
RIA, РВ, Ну, Cb, As, Vo, Ge, 00.

21. Gizzard Spines. Gizzard spines (Figs.
32-35) are here defined as acicular, cartilag-

FIGS. 30-31. Tuberculate gizzard plates. 30. Re-
tusa obtusa (Fleet, Dorset, U.K., 2/1986), SEM. 31.
R. obtusa (Fleet, Dorset, U.K., 2/1986), histological
cross-section, PAS stain. Scales = 100 um (30), 20
um (31.)

inous bodies affixed to the inner wall of the
esophageal gizzard, always in conjunction
with larger gizzard plates. They vary greatly in
size but are always substantially smaller than
the gizzard plates. The base is roughly circu-
lar and flat to slightly concave below. Lo-
cated at the periphery of the main gizzard
plate “field” (i.e., preceding or following the
gizzard plates), they often alternate with the
larger plates. Histologically and at low mag-
nification during gross dissection, a flattened
inner core of longitudinal striations 1$ visible
and often more darkly pigmented (Fig. 6A-C).
The tip of the spine is also flattened in the
plane of the striations. Gizzard spines control
the flow of food particles in and out of the
gizzard, including (depending on location)
prevention of (a) back-washing into the ante-
rior esophagus during processing, and (b)
exit from the gizzard until particles are small
enough for further digestion in the stomach.

Spines precede the gizzard plates in both
Akera and Aplysia. In fact, in Akera, small py-
ramidal plates showed strong similarity to the

392 MIKKELSEN

is

FIGS. 32-35. Gizzard spines. 32. From anterior gizzard; Aplysia brasiliana (НВОМ 065:00414). 33. Same;
Bulla ampulla (NNM). 34. Same; Haminoea antillarum (PMM-933), histological cross-section, PAS stain. 35.
From filter chamber of Akera bullata (ZMUC), histological cross-section, PAS stain. Scales = 1.00 mm (32,

33), 50 ит (34, 35).

larger conical spines, suggesting that the two
structures are in fact homologous. Removal
of both plates and spines from the gizzard
wall is easy and leaves behind a “scar” or
slightly elevated base to which they were af-
fixed. Conical spines also occur in the thin-
ner-walled filter chamber in Aplysia. The filter

chamber in Akera, although probably itself
homologous, contains different “spines”
(Fig. 35). These are thin cartilaginous caps
over elevated fleshy papillae, effectively hol-
low in histological cross-section, are not re-
movable from the gizzard wall, and do not
show longitudinal striations in the chitin. They

PHYLOGENETICS OF CEPHALASPIDEA 393

are not believed homologous to the gizzard
spines defined here.

Gizzard spines are present both preceding
and following the gizzard plates in Haminoea
and Bulla (Er. Marcus, 1957: fig. 3; Fretter,
1939: fig. 11, teeth”). They are tiny com-
pared with the robust plates, often frayed at
the tip, and (especially when small) do not as
readily detach from the gizzard wall. They are
heavily longitudinally striated and flattened,
morphologically resembling only the inner
core of Akera/Aplysia spines. This resem-
blance and their location are taken as evi-
dence of homology with the latter. They often
also show distinct, regular transverse striae,
presumably growth lines. Fretter (1939)
noted that during ontogeny, three groups of
plate-like projections later fuse to form the
three preceding spines. However, this was an
over-interpretation of Berrill (1931), who quite
clearly referred to three projections that fit
between the three gizzard plates in two-
month-old post-larvae.

In Smaragdinella, Rudman (1972c) was un-
clear т noting the gizzard as “very similar” to
that of Phanerophthalmus, which has gizzard
spines (Rudman, 1972c). Spines were ем-
dent preceding the gizzard plates in histolog-
ical sections and dissections of Smarag-
dinella, but they are thin, V-shaped chitinous
structures without obvious striation, and do
not readily detach. By virtue of their location
and chitinous nature, Smaragdinella is provi-
sionally coded 2.

Brown (1934: fig. 18) showed fleshy
“spines” between the gizzard plates of Phil-
ine aperta, with no comment in the text. In
dissection, these were confirmed as fleshy
ridges running the entire length of the gizzard
between each pair of plates. These are nei-
ther homologous nor analogous in function to
gizzard spines. These structures are also ev-
ident in published drawings of the gizzard of
Acteocina canaliculata (fide Ev. Marcus,
1977b: fig. 43).

Minichev (1971) stated that cuticular
spines were present on the gizzard walls in
Retusa instabilis Minichev, 1971. According
to his figures, these are minute; they proba-
bly correspond to small cuticle-covered pa-
pillae on the gizzard wall seen in histological
sections prepared for this study. Gizzard
spines as defined here are absent in R. ob-
tusa and R. truncatula.

In the absence of literature opinion, polarity
of this character mirrored that of character
19; because numerous gizzard plates (the

plesiomorphic state) in Aplysia and Akera are
accompanied by numerous gizzard spines,
the presence of gizzard spines was also con-
sidered plesiomorphic. This decision is ad-
mittedly arbitrary.

Coding: 0 = present (in Bu, Ha, Sm?, Ak,
Ap); 1 = absent (in Sc, PhA, PhB, Cy, Ai, ReA,
ReB); n (gizzard absent) in Ac, RiA, RiB, Hy,
Cb, As, Vo, Ge, 00.

22. Filter Chamber. A filter chamber, or
secondary gizzard, is present in anaspideans
between the gizzard and the stomach. This
character, plus character 24, were added to
the analysis as unifying data for Akera and
Aplysia.

Coding: 0 = absent (in all except following);
1 = present (in Ak, Ap).

23. Stomach. The stomach in opistho-
branchs was defined here as the (at least
slightly) expanded part of the post-esoph-
ageal digestive tract where the digestive
glands open. Unlike its vertebrate namesake,
where maceration of food occurs, this is the
center for sorting and/or absorption of usable
nutrients; food maceration usually occurs
earlier in the buccal mass or gizzard. [In
Acteon and Ringicula, with weak radulae and
without gizzards, maceration occurs in the
posterior region of the stomach (Fretter,
1939: 641; 1960).]

The stomach exists in two main formats:
those with a large posterior chamber (e.g.,
Acteon), and those without this chamber that
are effectively a simple flow-through tube. In
Ringicula, the large posterior pouch is elab-
orated by thickenings of the wall (Fretter,
1960; Gosliner, 1994) that act to crush large
prey items, analogous (but not homologous)
to gizzard plates.

Coding: 0 = with pouch-like chamber (in
Ac, RiA, RiB, Hy, Ge, 00); 1 = simple flow-
through, without pouch-like chamber (in all
others).

24. Caecum Extending from Stomach. The
anaspid stomach has a distal caecum con-
taining two longitudinal folds or typhlosoles
(Fretter & Ko, 1979). This is a unifying char-
acter for the anaspid clade, comprising Akera
and Aplysia.

Coding: 0 = absent (in all except following);
1 = present (in Ak, Ap).

25. Extent of Intestinal Typhlosole. Unlike
its vertebrate namesake, which serves in ab-
sorption of food nutrients, the function of the
opisthobranch intestine is the elaboration of
waste matter (Fretter, 1939). A typhlosole, or
longitudinal fold, acts to consolidate fecal

394 MIKKELSEN

pellets through secretions (Fretter & Ko,
1979). A partial typhlosole extending only a
short distance into the intestine was consid-
ered the plesiomorphic state, due to that in
the larger caenogastropod outgroup (Fretter
8 Graham, 1994: 220).

Coding: 0 = partial (in Ac, ВТА, Sc, PhA, Cy,
Ha, ReA, ReB?, 00); 1 = absent (in Hy, Bu, As,
Vo, Ak, Ap, Ge); 2 = entire (in Cb); u (un-
known) in RiB, PhB, Ai, Sm.

Nervous System: The presumed plesiomor-
phic nervous system in the Euthyneura (mod-
ified from Guiart, 1901: fig. 49; Russell, 1929:
200, text-fig. 1; Williams, 1975: fig. 7A) in-
cludes: six ganglia in a prepharyngeal esoph-
ageal nerve ring (right and left cerebrals, ped-
als, and pleurals), and five ganglia [a single
visceral (= abdominal), supra- and subesoph-
ageals (= -intestinals), and right and left pal-
lials (= parietals)] on a long streptoneurous
visceral loop emanating from the pleural gan-
glia. Other components of the basic system
include an osphradial ganglion emanating
from the supraesophageal, paired buccal
ganglia from the cerebrals, and a genital gan-
glion from the visceral.

The pallial ganglia are believed to repre-
sent detached portions of the pleural ganglia,
supplying the lateral body wall and parts of
the mantle (Bullock, 1965). Bullock (1965;
supported by Brace, 1977a) considered this
separation to be a phylogenetically recent
event. Their presence was given as the
“most important synapomorphy” for the
Pentaganglionata (= Euthyneura) by Hasz-
prunar (1988: 14; but see results for Gegania,
below).

Williams (1975) considered the relative po-
sitions of the nerve loop ganglia to be impor-
tant comparative criteria. Mere numbers of
ganglia on the loop (as in the traditional char-
acter “visceral loop ganglia 5/4/3”) do not
reflect their relative positions nor which gan-
glia are fused with which others. A visceral
nerve loop formula was devised to cladisti-
cally reflect this information. The most plesi-
omorphic arrangement advocated by Gos-
liner (1994) and earlier authors mentioned
above is L-A-B-V-P-A-L, reading from the left
origin on the circumesophageal nerve ring to
the right, where L = pleural (right or left), A =
pallial (right or left), В = subesophageal, \ =
visceral, and P = supraesophageal. [The
“neutral” positions exhibited here by the two
left (A, B) and two right (P, A) ganglia are
periodically referred to below as upper and

lower left, and upper and lower right posi-
tions, respectively.] The most plesiomorphic
condition (for purposes of assigning polarity)
was modified here to LA- -B-V-P- -AL, dif-
fering in location of the pallial ganglia, which
in this format are not yet separated from
the pleurals. No dashes between ganglia in-
dicate adjacency or fusion of the two ganglia;
no distinction is made between adjacent or
fused because the degree of fusion is often
ambiguous (i.e., is a dumbbell-shaped gan-
glion fused or not?). A single dash indicates
a short connective; two or three dashes indi-
cate longer connectives. As an example,
L-A- -BV- - -PAL indicates adjacent (or fused)
subesophageal/visceral and supraesoph-
ageal/right pallial with the latter adjacent to
the nerve ring, a short connective between
the nerve ring and left pallial, a long connec-
tive between pallial and subesophageal, and
a still longer connective between visceral and
supraesophageal. Because nearly all taxa in-
volved in this study had unique overall pat-
terns (making coding especially difficult), cer-
tain features of this were coded separately,
reflecting the location of key ganglia, using
care to create independent characters. Gan-
gliar fusion and streptoneury are not reflected
in this formula. The position of a single gan-
glion must be expressed with reference to
two other ganglia, e.g., the supraesophageal
in the above example 1$ at the V- -Р/РА po-
sition.

Shortening of the visceral loop was dis-
carded as a potential character, as in previ-
ous traditional characters, because of sub-
jectivity and dependence on overall body
size.

Within gastropods, streptoneury (= chias-
toneury) is plesiomorphic to euthyneury (= or-
thoneury) wherein the nerve loop is un-
crossed. Haszprunar (1988: 14) believed that
“euthyneury . . . is a phenomenon of multiple
convergence.” Williams (1975: fig. 31)
showed it evolving six times within the
opisthobranchs. The process of untwisting
has been attributed to causes other than
strict “detorsion” (Brace, 1977b), including
shortening of the visceral loop, posterior mi-
gration of the mantle skirt, differential growth,
and other changes.

The taxa involved in this study exhibit vary-
ing degrees of streptoneury, from fully
crossed (Acteon), fully uncrossed (Aplysia), to
“slight streptoneury” (Haminoea; terminol-
ogy of Williams, 1975), wherein the nerve
cords are uncrossed, but the right portion of

PHYLOGENETICS OF CEPHALASPIDEA 395

the loop is dorsal to the left portion. Position
of the “crossing” also varies, necessitating
precise definition here. Because Brace
(1977a) expressed euthyneury in part as mi-
gration of the supraesophageal ganglion to-
ward the right, assessment is here made at
the level of the supraesophageal ganglion, ei-
ther left (streptoneurous) or right (euthyneur-
ous) of the midline. Thus, Philine is euthyneu-
rous at the level of its supraesophageal
ganglion, even though its nerve loop 1$
crossed farther posteriorly [e.g., Philine sp. 1
of Williams (1975: fig. 8) with a posterior
“vestige of streptoneury” (Brace, 1977a:
19)]. This was also noted in dissections of
Haminoea antillarum (Orbigny, 1841) during
this study.

“Slight streptoneury” (= incomplete euthy-
neury) was determined if the right arm of the
visceral nerve loop was positioned dorsal to
the esophagus. In Aplysia, wherein all nerve
loop ganglia are crowded about the visceral
ganglion, the only “vestige of streptoneury”’
is found in the dorsal position of the suprae-
sophageal ganglion over the more ventral
viscero-subesophageal ganglion (Guiart,
1901); but because both arms of the visceral
loop are ventral to the gut, Aplysia was con-
sidered fully euthyneurous.

Some degree of variability has been im-
plied in Akera with regard to this character.
Both Guiart (1901, in A. bullata Múller, 1776)
and Ev. Marcus (1970, in A. bayeri Marcus 4
Marcus, 1967) depicted the supraesophageal
ganglion distinctly on the left side, that is, п
streptoneurous position. Brace (1977a: fig.
2B; in A. bullata) drew a clearly uncrossed
visceral nerve cord. Hoffmann (1936: fig. 484)
explained this difference as one related to
isolated versus in situ configurations of the
loop.

Following coding of this character, it was
noted in complete congruence with the posi-
tion of the mantle cavity opening (character
4). Because posterior migration has been
proposed as a possible cause of the change
from streptoneury to euthyneury, these char-
acters were considered not independent,
and this character was not used. Coding 1$
here noted for information purposes only: O =
streptoneurous (crossed visceral loop; in Ac,
Ri, Ge); 1 = slightly streptoneurous (un-
crossed but right arm of nerve loop dorsal to
left; in Hy, Sc, Ph, Cy, Ai, Bu, Ha, Re, Cb, As,
Vo, Ak, Ap); 2 = euthyneurous (uncrossed,
arms of visceral loop level; in Sm, Ru).

Statocysts are roughly spherical, fluid-

filled organs, containing hard bodies called
statoliths (large, single) or statoconia (small,
multiple), which assist the mollusk in deter-
mining its own orientation. Dorsett (1986)
noted that the anatomy of molluscan stato-
cysts is poorly known from a comparative
point of view. These were observed here in
histological sections of all taxa except
Acteocina and Volvatella; Aplysia was not
sectioned, therefore also not observed. In all
but a few exceptions, they were appressed to
the surface of the pedal ganglia (near the
junctions with the cerebro-pedal connec-
tives), and contained multiple statoconia.
This agrees with the general description “in
gastropods” given by Dorsett (1986). Stato-
conia were clearly observed in Hydatina,
Bulla, Haminoea, Smaragdinella, and Gega-
та as minute flattened disks, each with a
central thickening, as discussed and figured
for Aplysia californica Cooper, 1863, by Cog-
geshall (1969). In other species, statoconia
were indistinct flakes and dots (Acteon,
Scaphander, Philine A, Cylichna, Cylindro-
bulla) or were inevident (Ringicula, Retusa A,
Ascobulla, Akera). Hoffman’s (1935) figure of
the statocyst of Aplysia suggested both a
larger statolith and multiple statoconia, al-
though this disagrees with the findings of
Coggeshall (1969). Guiart (1901: 120) also re-
marked on the absence of a large “otolithe.”
Fretter 8 Graham (1994) stated that, although
multiple statoconia appeared more charac-
teristic of “primitive”” gastropods and the sin-
gle statolith of “more advanced” forms, no
clear systematic generalizations were possi-
ble. Haszprunar (1988: 399) also regarded
this dichotomy as “highly ambiguous.” In
some species (e.g., Tritonia hombergi Cuvier,
1802), statoliths and statoconia are both
present, with statoconia added progressively
during development (Thompson, 1962). No
coding was possible on this information.

Gegania showed the only exception in sta-
tocyst location, appressed to the pedal gan-
glion near the junction with the pedal com-
missure. Location of the statocysts, although
possibly useful phylogenetically, differed in
only one species and as such would be an
uninformative character in the analysis. Also,
details of the cellular fine structure of the sta-
tocyst itself, discussed as evolutionary infor-
mative by Barber (1968), were not resolvable
by the light histological techniques employed
here.

Twelve characters were coded from the
nervous system.

396 MIKKELSEN

26. Nerve Ring Location. The six-ganglion
nerve ring around the anterior esophagus
can be located either anterior or posterior
to the buccal mass. Gosliner (1994) deter-
mined that the postpharyngeal position 1$
plesiomorphic for Gastropoda (based on
the condition in veti- and caenogastropods),
but that the prepharyngeal position 1$ a
synapomorphy for the Heterobranchia (sec-
ondarily moved postpharyngeally within the
group, hence determining the polarity here
assigned). The prepharyngeal position is
present only in Cephalaspidea within the
Opisthobranchia (Williams, 1975). The post-
pharyngeal position could be associated with
centralization of the nervous system toward
the midpoint of the body (Williams, 1975).
Williams (1975: fig. 33) perceived this shift to
have occurred at least three times during
opisthobranch evolution.

Members of the genus Retusa have no
muscular “pharynx” or buccal mass, render-
ing it difficult to determine the coding of this
character. However, a reasonable determina-
tion was derived from the location of the
paired buccal ganglia relative to the nerve
ring. In most cephalaspids, the buccal gan-
glia are located at the posterior region of the
buccal mass, adjacent to its junction with the
esophagus. In species with a postpharyngeal
nerve ring (e.g., Akera), the buccal ganglia are
located anterior to or in close proximity to the
nerve ring. Alternatively, in species with a
prepharyngeal nerve ring (e.g., Acteon, Cy-
lichna), the buccal ganglia are located poste-
rior to the nerve ring. Histological sections of
R. obtusa showed the buccal ganglia to lie
significantly behind the nerve ring. This ob-
servation is supported by Hurst (1965: figs.
24, 31), who additionally showed the nerve
ring surrounding the “oral region” anterior to
an expanded, weakly muscular “buccal re-
gion.” The latter carries the buccal ganglia
posteriorly, adjacent to the junction with the
esophagus. Hurst's “buccal region” thus ap-
pears homologous to the functional buccal
mass of other cephalaspids; therefore, the
nerve ring was considered prepharyngeal
and this character was coded O for Retusa.
Boss (1982: 1026) apparently agreed but in-
accurately stated “the nerve collar is in front
of the pharynx” (emphasis mine) in his de-
scription of Retusidae.

The nerve ring of Hydatina physis in histo-
logical sections studied here surrounds a
muscular portion of the pharynx containing
the jaws, i.e., somewhat mid-region of the

pharynx. This, however, was anterior to both
the radular region of the pharynx and the
buccal ganglia, concluding that the nerve ring
must be coded as prepharyngeal (0).

Coding O = prepharyngeal (in Ac, Hy, Sc,
PhA, РВВ, Cy, Ai, Bu, ReA, ReB, Ge, 00); 1 =
postpharyngeal (in RIA, РВ, Ha, Sm, Cb, As,
Vo, Ak, Ap).

27. Cerebral/Pleural Ganglia. Fusion of
these two ganglia is clearly indicated in his-
tological sections by the presence of two
connectives (rather than one) between the
cerebral (actually the cerebropleural) and
pedal ganglia. This is identical to the tradi-
tional characters “nerve ring with 6 or 4 gan-
glia.” This and the following two characters
are expressions of “concentration” of the
nervous system, a phenomenon reportedly
occurring independently in many groups of
“lower” opisthobranchs (Rudman, 1972c).

Cerebral and pleural ganglia are fused in
histological sections of Ringicula nitida, but
are separate but closely apposed in R. buc-
cinea (fide Fretter, 1960). This character was
therefore coded as separate for Ringicula В,
and fused for Ringicula A.

Although the cerebral and pleural ganglia
are fused in the type of Retusa, R. obtusa,
these ganglia are separate in many other
species of Retusa (e.g., R. truncatula). This
character is therefore coded 0 (separate) in
Retusa B and 1 (fused) in Retusa A.

Haszprunar's (1985b) claim that the cere-
bral and pleural ganglia of Gegania valkyrie
are separate on the left and fused on the right
was confirmed by examination of Haszpru-
nar's own histological sections as well as an
additional set, however, the fusion on the
right is incomplete and two ganglia are
clearly distinguishable (Fig. 36). Gegania is
coded here as separate.

Coding: 0 = separate (in RiB, Sc, PhA, РВВ,
Cy, Ai, Bu, Ha, Sm, ReB, Ak, Ap, Ge, 00); 1 =
fused (т Ac, РИА, Ну, ReA, Cb, As, Vo).

28. Relative Length of Cerebral Commis-
sure. Within the nerve ring, cerebral and
pedal ganglia are either closely adjacent (CC
or PP) or connected by a distinct commissure
(C-C or P-P). The latter state is true for both
ganglion pairs in the hypothetical ancestral
condition presented by Russell (1929) and
Gosliner (1978, 1981, 1994). Shortening of
commissures (and connectives) is another
reflection of concentration of the nervous
system.

Coding: 0 = long (C-C, in all but following);
1 = short (CC, adjacent; in Hy, Ak, Ap, Ge).

PHYLOGENETICS OF CEPHALASPIDEA 397

FIG. 36. Circumesophageal nerve ring of Gegania valkyrie (MNHN), histological cross-section, PAS stain.
Dorsal at upper right, right side of animal at left. AG = pallial ganglion (right); СС = cerebral commissure;
CG = cerebral ganglion (right); ESO = esophagus; LG = pleural ganglion (right); М = mantle; PA = right

pallial-supraesophageal connective. Scale = 100 um.

29. Relative Length of Pedal Commissure.
See comments under character 28.

Coding: 0 = long (Р-Р, in all but following);
1 = short (PP, adjacent; in Cb, As, Vo).

30. Position of Left Pallial Ganglion. Bul-
lock (1965: 1362) considered the first indica-
tions of concentration of the visceral loop to
be “the movement of the parietal ganglia for-
ward” seen in many cephalaspids. This as-
sumes that the plesiomorphic state is one in
which the pallials are substantially removed
from the pleurals. Here, because the pallial
ganglia are presumed to have split from the
pleural ganglia (Bullock, 1965; Brace, 1977a),
the state LA (adjacent) is presumed plesio-
morphic to L-A or L- -A, which are here both
considered as character state 1.

The left pallial ganglion is reportedly miss-
ing in Scaphander (Brace, 1977a), Cylichna
(Lemche, 1956), and Асеоста (Gosliner,
1979). Brace (1977a) and Schmekel (1985)

considered the left pallial fused with the left
pleural in Scaphander; however, Brace
(1977a) noted (also in Cylichna) the presence
of vestigial somata at the junction of the left
pallial nerve with the visceral loop, in a upper
left position. lt seems more likely therefore
that the left pallial ganglion has been reduced
in these three taxa, rather than fused with the
left pleural. The position of the left pallial is
therefore interpreted through that of the left
pallial nerve; these three taxa are therefore
coded here as separate.

No left pallial was detected in dissections
of Philine aperta or P. orientalis A. Adams,
1854; Philine A was coded “u.” Philine falk-
landica and P. angasi (Crosse 8 Fischer,
1865) apparently have a left pallial ganglion
(as “accessory” ganglion, Rudman, 1972b)
migrated toward the visceral, and Philine В 1$
coded 1 accordingly.

The position of the left pallial ganglion var-

398 MIKKELSEN

ies in the genus Haminoea. It could not be
discerned as a separate ganglion in dissec-
tions of H. elegans (Gray, 1825) and H. antil-
larum, although in each case nerves were
noted innervating the pallial wall from the
pleural ganglion as well as from the lower left
position of the visceral loop (i.e., in the lower
left position). Er. Marcus (1958) found no in-
dication of a left (or right) pallial in dissections
or sections of H. elegans. Guiart (1901: pl. 5)
showed a pallial nerve and a “ganglion pal-
léal gauche” in the lower left position in H.
navicula (da Costa, 1778). Vayssiere (1879-
1880: pl. 12, fig. 114) figured a small pallial
nerve immediately adjacent to the subesoph-
ageal ganglion (i.e., in L- - -A/AB position) in
H. hydatis (Linné, 1758). [Thompson (1976:
fig. 58e) extrapolated a small ganglion at the
base of this nerve in his redrawing of Vays-
siere's (1879-1880) figure.] Rudman (1971a:
fig. 11) found several left pallial nerves, the
largest at the upper left position, and several
smaller ones nearer the adjacent BV ganglia.
Williams (1975: fig. 14) claimed the left pallial
to be fused to the left pleural in H. virescens
(without any indication of pallial nerves). In
this dataset, Haminoea was provisionally
coded 1 in accordance with its exemplar
species.

Although Williams (1975: 96) considered
the left pallial ganglion in julioid sacoglossans
“fused to some degree within the anterior
nerve ring,” two distinct ganglia were found
on the left side of the visceral loop in Cylin-
drobulla and Ascobulla, identifiable as the left
pallial and subesophageal ganglia. In Volva-
tella, the pallial ganglion was less clear in his-
tological sections but suggested the same
configuration. In other species of Volvatella
[V. vigorouxi (Montrouzier, 1861) and У. ven-
tricosa Jensen 8 Wells, 1990), Baba (1966)
and Jensen & Wells (1990) showed the sube-
sophageal in the upper right position, but this
is likely the left pallial ganglion [with the sube-
sophageal fused with the visceral, as in V.
bermudae Clark, 1982].

Haszprunar's (1985b) interpretation of the
nervous system of Gegania included no men-
tion of pallial ganglia, however, both right and
left were identified here. The right pallial gan-
glion (= Haszprunar's right pleural) is imme-
diately adjacent to the partially fused cere-
bro-pleural ganglion; this was confirmed as
pallial, not pleural, because it lies along the
pleural-supraesophageal connective and
does not receive a connective from the pedal
ganglion (as does the pleural). The left pallial

ganglion appears to be what Haszprunar
(1985b) called “accessory ganglion,” de-
picted as separate from the pleural and in-
nervating the columellar muscle; its presence
was not confirmed in sections examined.

Coding: O = LA, fused or adjacent (in RiA,
RIB, Hy, 00); 1 = L-A or L- -A, separate (in Ac,
Sc, PhB, Cy, Ai, Bu, Ha?, Sm, Cb, As, Vo, Ak,
Ge); u in PhA, ReA, ReB, Ар.

31. Position of Subesophageal Ganglion.
Brace (1977a) stated that the posterior mi-
gration of the subesophageal ganglion to-
ward the visceral (i.e., B-V to BV) was caused
by the posterior migration of the mantle skirt.
However, there is imperfect correlation be-
tween this and character 4, therefore this
character was also used in the analysis.

In Hydatina physis, according to Rudman
(1972a), the subesophageal ganglion is in the
lower left position. However, in my dissec-
tions of H. physis and in H. velum according
to Eales (1938), the subesophageal has
clearly migrated posteriorly to lie adjacent to
the visceral. Because of the conflicting ob-
servations of the type species, H. physis, Hy-
datina is here coded “u.”

The subesophageal was confirmed adja-
cent to the visceral in my dissections of
Volvatella bermudae. lts position adjacent to
the left pleural in V. vigourouxi and V. ventri-
cosa (fide Baba, 1966; Jensen & Wells, 1990)
is probably an error, mistaking the prominent
left pallial for the subintestinal.

Coding: 0 = B-V (in Ac, Cb, As, Ge, 00); 1 =
B migrated toward nerve ring (in RiA, RiB); 2
= BV, migrated toward visceral (in Sc, PhA,
PhB, Cy, Ai, Bu, Ha, Sm, ReA, ReB, Vo, Ak,
Ap); u in Hy.

32. Position of Supraesophageal Ganglion.
In Retusa obtusa and R. truncatula, the su-
praesophageal ganglion is in the lower right
position on the nerve loop (Vayssiere, 1893;
pers. obs.). However, in R. semisulcata, it is
adjacent to the right pleural at the nerve ring
(Huber, 1993). Retusa A is accordingly coded
0, and Retusa B is coded 1.

Williams (1975) assigned Cylindrobulla, As-
cobulla, and Volvatella to the julioid type of
nervous system with visceral loop formula
LA- -B-V-P- -AL. However, in this study, the
supraesophageal and right pallial ganglion
were clearly fused in histological sections of
all three taxa.

The supraesophageal ganglion is in the
lower right position on the nerve loop in
Volvatella vigourouxi (fide Baba, 1966), V.
ventricosa (fide Jensen & Wells, 1990), and in

PHYLOGENETICS OF CEPHALASPIDEA 399

my observations of V. bermudae. However, in
his work on V. bermudae, Clark (1982: fig.
1G) placed it adjacent to the right pleural,
close to the nerve ring. Because of the con-
flicting observations in the exemplar, Volva-
tella is coded “и” here.

Coding: 0 = V-P (in Ac, РИА, Sc, РВВ, Cy,
Ai, Bu, Ha, ReA, Cb, As, Ak, Ge, 00); 1 = P
migrated toward nerve ring (in Hy, PhA, Sm,
ReB); 2 = VP (in Ap); u in Vo.

33. Position of Right Pallial Ganglion. See
general comments about pallial ganglia un-
der character 30.

Williams” (1975) julioid formula (see char-
acter 32) assumed fusion of the right pallial
ganglion with the right pleural. However, in
this study, histological sections of Cylindro-
bulla and Ascobulla suggested fusion of the
right pallial with the supraesophageal. This
was especially clear in Cylindrobulla, in which
the presumed fused ganglion was almost bi-
lobed. In these cases, the right pallial and
right pleural are widely separated. Although
histological sections of Volvatella bermudae
also showed the -PA--L configuration,
closely apposed ganglia (-PAL) were re-
corded in V. bermudae by Clark (1982) and
also in V. ficula Burn, 1966, by Burn (1966).
Volvatella was here coded 2, in accordance
with the condition observed here for the ex-
emplar species.

See remarks on Gegania under character
30.

Coding: 0 = AL (in Hy, Sc, PhA, PhB, Cy,
Ai, Bu, Ha, Sm, ReA, ReB, Ge, 00); 1 = A-L (in
Ac, RiA, RiB); 2 = PA, migrated toward su-
praesophageal (in Cb, As, Vo, Ak, Ap).

34. Position of Genital Ganglion. Brace
(1977a: 19) noted several configurations of
the genital ganglion, either on a nerve off the
visceral ganglion or directly on the visceral
loop between the visceral and supraesoph-
ageal ganglia. Because he assumed that
well-formed plexuses and ganglia would
have prevented “migration” of the genital
ganglion through the visceral, he was “fairly
certain that new cells must have arisen to
form the genital ganglion” in the latter case.

Configurations observed during this study
suggest the reverse, that genital ganglia in
either location are homologous. A distinct
ganglion on a nerve emanating from the vis-
ceral ganglion was found in many species,
including Acteon, Hydatina, Volvatella, and
Aplysia; the nerve was confirmed (but not the
ganglion) in some Philine species and Gega-
nia. Alternatively, among those having the

genital ganglion directly on (or on a separate
nerve off) the visceral loop, Scaphander,
Acteocina, Bulla, and Haminoea (also illus-
trated in Retusa semisulcata by Huber, 1993)
confirmed the genital ganglion on a nerve off
the visceral, but also on the loop by its also
receiving the connective from the suprae-
sophageal ganglion. It thus appears that the
VP connective has been relocated.

Polarity of this character was established
by Russell (1929: text-fig. 1) and Williams
(1975: fig. 1), supported by a similar condi-
tion in the caenogastropod Littorina (genital
nerve only; Fretter & Graham, 1994).

Philine B was coded 0 here because of the
condition of P. falklandica and others (Rud-
man, 1972b; Williams, 1975; Gosliner, 1978);
in other species in the genus, including the
type P. aperta (fide Guiart, 1901; Brace,
1977a), the genital ganglion is part of the vis-
ceral loop, therefore Philine A is coded 1.

The genital ganglion in Akera is on the vis-
ceral loop immediately adjacent to the con-
joined visceral and subesophageal ganglia.
Er. Marcus (1970) called the right-most swell-
ing the visceral ganglion, but Brace’s (1977a)
careful tracing of the nerves associated with
the posterior visceral loop confirms that this
is the genital ganglion. The connective from
the supraesophageal ganglion merges with
the genital ganglion attesting its position on
the loop.

Coding: 0 = off visceral ganglion (in Ac, Hy,
PhB, Cb, As, Vo, Ap, Ge, 00); 1 = on visceral
loop between V and P (in Sc, PhA, Cy, Ai, Bu,
Ha, Sm, ReA, ReB, Ak); u in RiA, RiB.

35. Eye Direction. The only traditional char-
acter regarding the eyes is a simple “ргез-
ence/absence,” the latter condition applying
only to Retusa. However, this study shows
that although eyes are indeed absent т R.
obtusa, they are well formed in R. truncatula.
Retusa A is therefore coded “‘п”; coding for
Retusa B for all eye characters is here based
on R. truncatula.

Direction was taken as direction of the cor-
nea and lens, in other words, the direction of
vision. Curvature of the black pigmented
layer (Figs. 38, 40) can also indicate direction,
that is, concave toward the direction of vi-
sion, convex toward the optic nerve. Ventro-
lateral eyes (Figs. 39, 40) are apparently
“seeing” through the lateral groove between
the cephalic shield and foot, which might be
appropriate for burrowing snails.

From the literature as well as from this
study, eye direction is inconstant within the

400 MIKKELSEN

Sacoglossa. Ventrolaterally directed eyes
were found here in Cylindrobulla, Ascobulla,
and Volvatella. They were likewise figured in
Ercolania lozanoi Ortea, 1981, by Fernandez-
Ovies et al. (1984: fig. 1B). Dorsally directed
eyes were noted in Elysia viridis (Montagu,
1804) (pers. obs.) and in Tamanovalva limax
Kawaguti 8 Baba, 1959 (Kawaguti & Yamasu,
1966).

The eye of Volvatella bermudae, as ob-
served here, points ventrolaterally. However,
the eyes of the type species V. fragilis Pease,
1860, are situated on the dorsal surface of
the head, apparently pointing dorsally
(Evans, 1950). This character is coded 1 for
Volvatella, in accordance with the exemplar.

Coding: O = dorsolateral (т Ac, МА, РВ,
Hy, Bu, Ha, Sm, Ap, Ge, 00); 1 = ventrolateral
(in Sc, PhA, PhB, Cy, Ai, ReB, Cb, As, Vo,
АК); n in ReA.

36. Eye Location. Differences in location of
the eye in opisthobranchs were first noticed
by Willem (1892: 141), who designated four
lettered categories. Those of Aplysia, Hami-
noea, and Bulla were (a) superficial ““comme
chez les Prosobranches.” The eyes of other
taxa not involved here were (5) slightly
deeper, “dans l'epaisseur des téguments”
(within the thickness of the skin), (c) deep
within the integument, and (d) completely
within the body cavity next to the cerebral
ganglia. Within this dataset, the eye of
Scaphander could be placed in Willem's cat-
egory d. However, because of the subjectiv-
ity of Willem's categories, plus the difficulties
of dealing with histological sections not а|-
ways cut in perfect cross-section, the more
objective categories used here were devised.

Distance to the surface (S) was taken to the
surface nearest the lens of the eye, that is,
that surface through which the eye would
“see.” Although distance to the nerve ring
might be a more logical denominator for use
in the above ratio, distance to the center of
the gut (C) was a more reliable measurement
in preserved, potentially contracted speci-
mens. The two distances were usually off-
angled, that is, not measured in a single con-
tinuous line (from surface to center) in a
histological cross section. Location was ex-
pressed as a ratio of distance to surface over
distance to the center (S/C).

As with the eye direction, Fretter (1960:
545) associated withdrawal of the eye from
the surface with the burrowing habit.

As with the previous character, Volvatella is
here coded according to the exemplar spe-

cies, although according to Evans (1950), the
eyes of V. fragilis are superficial.

Coding: 0 = close to surface [S/C < 0.2] (in
Ac, Hy, Bu, Ha, Sm, Ak, Ap, Ge, 00); 1 =
midway between surface and nerve ring [S/C
= 0.2-0.5] (in RiA?, RiB?, Ai?, Vo); 2 = deeply
embedded, near nerve ring [S/C > 0.5] (in Sc,
PhA, РВВ, Cy, ReB, Cb, As); п in ReA.

37. Eye Lens (Shape). The lens of the eye
existed in three distinct forms within this
dataset: (a) spherical, (b) irregular, and (c)
hollow. Solid spherical lenses (Fig. 38) are
found in caenogastropods (Willem, 1892) and
in other lower heterobranchs (Valvata, Hy-
man, 1967: fig. 122b), therefore this was
taken as the most plesiomorphic state for this
character. All spherical lenses in histological
sections stained differently centrally than pe-
ripherally, in most cases orange throughout
most of the interior and dark pink at the mar-
gin. Irregular lenses (Fig. 40) stained uni-
formly one color throughout (dark pink in
PAS, light blue in Gomori's). This implies
non-uniform composition in the spherical
lenses, but this was not determined further.
Akera has an oblong-oval lens that 15 histo-
logically similar to the spherical lenses of
other species; it was therefore also coded
as 0.

The irregular lenses of the three shelled sa-
coglossans have distinct hollows associated
with the presence of connective tissue “an-
chors” to the inner surface of the eyeball (Fig.
40, АМС). A similar irregular lens with an an-
chor was figured in Julia japonica Kuroda &
Habe, 1951, but without comment by Ya-
masu (1968: figs. 9, 10). This is not, however,
a clear-cut sacoglossan character, because
the eye of Elysia viridis sectioned during this
study has a plesiomorphic round lens. One
non-sacoglossan, Philine, has an irregular
lens, staining uniformly pink in sections, and
with distinct surface hollows; although the
presence of anchors in this taxon was un-
clear, it was coded 1 here. The relationship
between these hollows and the enclosed
cavity of the derived hollow eye of Gegania is
unknown. (The latter is ovoid in cross-section
and therefore could effectively function as a
solid lens.) Several spherical lenses (e.g.,
Smaragdinella) showed wide lateral “pegs.”
Staining indicated that these were of lens
material, therefore different than the connec-
tive tissue “anchors” seen in other species.

The sectioned round lenses in Haminoea
antillarum and in one of two specimens of
Retusa truncatula showed shallow surface

PHYLOGENETICS OF CEPHALASPIDEA 401

г". 90
P sa

FIGS. 37-40. Eyes. 37. Dorsally directed eye; Runcina coronata (Quatrefages, 1844) (ZMUC), histological
cross-section, PAS stain. 38. Same, close-up of round lens. 39. Ventrolaterally directed eye; Cylindrobulla
п. sp. (Bermuda, 8/1990), histological cross-section, Gomori stain. 40. Same, close-up of irregular lens with
anchor (ANC). АМС = anchor; CO = copulatory organ; Е = eye; Е = foot; L = lens; OT = oral tube. Scales

= 100 ит (37, 39), 20 шт (38, 40).

hollows, but these were without ‘‘anchors”
and the lenses were bicolored, therefore
these cases were coded as spherical (0). The
extremely small lens of Scaphander lignarius
also had shallow hollows and was slightly ir-
regular in cross-section, but it was slightly

orange-stained at the center and lacked “ап-
chors,” therefore it was coded as spherical
(0). It is possible that some irregular shape
and the shallow hollows are fixation artifacts.

Coding: 0 = solid spherical or oblong-oval
(in Ac, RiA, Hy, Sc, Ai, Bu, Ha, Sm, ReB, Ak,

402 MIKKELSEN

Ap, 00); 1 = solid irregular with hollows/an-
chors (in PhA, Cb, As, Vo); 2 = hollow irreg-
ular (in Ge); n (absent) in Cy, ReA; u in RIB,
PhB.

Reproductive System: Ghiselin (1966: fig.
1C; reproduced by Hadfield & Switzer-Dun-
lap, 1984: fig. 1C) figured the generalized
hermaphroditic reproductive system of a hy-
pothetical opisthobranch ancestor. It is mo-
naulic (having a single pathway for both eggs
and sperm through the female gland mass),
with an external ciliated groove connecting
the common genital (= hermaphroditic) open-
ing with the base of the nonretractile copula-
tory organ with external groove only (without
ejaculatory duct), a pallial prostate gland
[“pallial” as used by Ghiselin (1966), implying
embryonic ectodermal origin, not location in
mantle (= pallial) cavity], a proximal receptac-
ulum seminis, and a distal bursa copulatrix.
[Proximal (near gonad) and distal (near com-
mon genital opening, away from gonad) po-
sitions of allosperm sacks are here as used
by Gosliner (1981) and Haszprunar (1988).]
This agrees completely with Gosliner's (1978,
1981, 1994) Hypothetical Ancestral Opistho-
branch.

Some species (e.g., members of Acteon,
Cylindrobulla) are diaulic, with separate path-
ways for autosperm and eggs. In such spe-
cies, an internal sperm duct splits off from the
hermaphroditic duct after exiting the gonad.
This duct travels internally to the copulatory
organ, and continues into the penis as an in-
ternal ejaculatory duct.

The copulatory organ (“реп!5” of authors)
of shelled opisthobranchs 1$ a varied and of-
ten elaborate structure. In species with re-
tractile copulatory organs, a penial sheath
houses the intromittent organ or what | term
the penis (““cirus” of Ghiselin, 1963; “penial
papilla” of Gosliner, 1990). In monaulic spe-
cies, autosperm travels to the copulatory or-
gan via the external ciliated groove, continu-
ing into the penial sheath via additional
groove(s) (“sulcus seminalis”” of Lemche,
1956), then through the penis either in an ex-
ternal groove or an internal ejaculatory duct.
Terminal glands (''prostate” of authors)
and/or sperm storage areas ('“sperm bulb” of
Gosliner, 1990, and others) communicate
with the base of the penis.

Glandular tissue associated with the cop-
ulatory organ was noted in Ringicula,
Scaphander, Philine, Cylichna, Acteocina (?),
Bulla, Haminoea, Smaragdinella, and Retusa

(?). Because gross morphology and cellular
configuration appeared to be highly variable,
homology could not be presumed with con-
fidence across the taxa involved. This struc-
ture has also been credited with spermato-
phore formation in a number of taxa
(Ghiselin, 1966). This character was therefore
not used in the cladistic matrix.

Glandular tissue surrounding the internal
sperm duct in the vicinity of the female gland
mass is here called “prostate” in convention
with the literature. However, two types of ap-
parently non-homologous glandular “‘pros-
tate” were distinguished histologically: (a)
dense tissue surrounding an expanded cen-
tral ciliated lumen [in Acteon, Hydatina (Rud-
man, 1972a: fig. 6); Fig. 41], and (b) glandular
tissue in a “flower” configuration, that is,
large club-shaped cells surrounding a narrow
central ciliated lumen (also shown by Ev.
Marcus, 1972: fig. 15, for Ascobulla; Reid,
1964: fig. 4A, for Elysia; Sanders-Esser,
1984: fig. 4a, for Ercolania; Fig. 42). The hy-
pothetical opisthobranch ancestors of Had-
field & Switzer-Dunlap (1984) and Gosliner
(1981) were said to have pallial prostates;
however, configurations were not inferred by
the authors. Also because of tremendous va-
riety at the histological level, penial “pros-
tates” (see above) were not here considered
homologous with pallial prostates, as tradi-
tionally used.

Prostates surrounding an internal sperm
duct are absent in all species lacking the in-
ternal duct, requiring that 13 of the 20 taxa in
the data matrix be coded as ‘п.’ This type of
character would add too much uncertainty to
the data matrix, and was therefore omitted
from the analysis.

Morphology of the spermatozoa of the taxa
involved in this analysis is too incomplete to
be effectively used here, but will be dis-
cussed below.

Spermatophores can be structurally com-
plex and taxonomically useful in gastropods
(Mann, 1984). They have been reported in a
number of shelled opisthobranchs [Hami-
noea hydatis and Ademnestia arachis (Quoy
& Gaimard, 1833) (Cylichnidae), fide Perrier &
Fischer, 1914; Volvulella persimilis (Mórch,
1875) (Retusidae), fide Ev. Marcus & Er. Mar-
cus, 1960; Metaruncina setoensis (Baba,
1954) (Runcinidae), fide Ghiselin, 1963;
Runcina ferruginea Kress, 1977 (Runcinidae),
fide Kress, 1985], presumably formed by the
“prostate” part of the copulatory organ.
Spermatophores are also evident in caeno-

PHYLOGENETICS ОЕ CEPHALASPIDEA 403

FIGS. 41-42. Pallial prostate glands. 41. Aceton tornatilis (ZMUC), histological cross-section, PAS stain. 42.
“Flower” configuration; Cylindrobulla п. sp. (Bermuda, 8/1990), histological cross-section, Gomori stain.
Scales = 100 um.

404 MIKKELSEN

gastropods (Robertson, 1989), lower hetero-
branchs (Pyramidellidae, Architectonicidae;
Robertson, 1989), Acochlidiacea, and Nudi-
branchia (last two summarized by Mann,
1984). Unfortunately, occurrence of sper-
matophores has been noted in too few taxa
involved in this study, and more information
seems necessary over mere presence/ab-
sence; data are thus too incomplete to be
coded at the present time.

In most taxa of this dataset, the fertilized
eggs exit the reproductive tract via a duct
that passes directly through the so-called
“female gland mass’ complex. This com-
prises the albumen, capsule (= membrane,
winding), and mucus (= nidamental) glands,
which add the nutritional and protective coat-
ings of the eventual egg mass. While the
smaller albumen and capsule glands in some
cases empty their secretions into the main
duct, eggs always directly traverse at least
the large mucus gland. Lloyd (1952) noted
that in several opisthobranchs, instead of
surrounding the main duct, the female gland
mass is a diverticulum. Passage through the
mass is thus a detour off the main duct,
through a loop with entry and exit in close
proximity. In one such taxon, Philine, the
eggs are retained in this gland mass until the
egg mass is complete to be expelled all at
once (Lloyd, 1952); this is in contrast to oth-
ers, for example, Aplysia, that gradually re-
lease their egg masses as long strings.

This character was not used here for two
reasons: (a) the diverticulum configuration,
present in at least Scaphander, Philine, and
Acteocina (Lloyd, 1952; Gosliner, 1979), is
comparable to the direct configuration ex-
cept that the duct and mucus gland are
folded into a loop, and (b) coding cannot be
made with confidence without detailed anal-
yses of egg and sperm pathways from live
animal studies, for example, those of Robles
(1975) for Bulla and of Thompson 4 Bebbing-
ton (1969) for Aplysia.

Some shelled opisthobranchs and nudi-
branchs have been reported with a “райси-
lar kind of ciliated strip . . . which moves the
eggs around the sperm” in the ampulla (=
hermaphroditic duct) considered to have po-
tential in indicating relationships (Ghiselin,
1966: 345-346). Fretter & Graham (1954) de-
scribed this as an unciliated strip in Acteon,
whereas Rudman (1972b) noted a “wide cil-
iated tract” in Philine. Gosliner (1994) noted
that a ciliated strip has also been reported in
anaspideans. This character was not pur-

sued, in part because the absence of cilia
could not be reliably confirmed without ex-
tensive histological study of each species in-
volved. Hadfield & Switzer-Dunlap (1984:
216) also noted ‘‘considerable variability
among species” in the ampullar wall.

Ghiselin (1966) noted that in some shelled
opisthobranchs, the female gland mass (=
pallial gonoduct) is internally divided. Use of
this character awaits better study of the fe-
male gland complex.

Haszprunar (1985a: 30) used the character
“gonoduct pallial/coelomic,” stating that
“the gonoduct is sunken into the haemocoel
of the body” at a branching point on his tree
just prior to Pentaganglionata (= Euthyneura).
If so, his “architectibranch” taxa (here
Acteon, Ringicula, Hydatina) would be pallial
in this regard. None of the taxa examined
here were pallial in the sense understood in
caenogastropods, wherein the female glands
are a prominent component (sometimes an
open trough) in the mantle cavity. This char-
acter was not considered usable in this anal-
ysis. [This distinction is not the same as pal-
lial and coelomic as used by Ghiselin (1966),
who referred to the embryonic origins of the
various organs, not the ultimate locations.]

Nine characters were coded from the re-
productive system.

38. Internal Sperm Duct. This and the fol-
lowing character represent the traditional
character expressed as “reproductive system
monaulic or diaulic.”” Мопаийс hermaphro-
dites have no separation of eggs and auto-
sperm (except perhaps by separate ciliated
tracts) as the latter exit the reproductive tract
via the elaborate female gland mass. Diaulic
forms have developed either a separate in-
ternal duct for eggs (6odiaulic) or sperm
(androdiaulic). The closed internal sperm duct
discussed previously is indicative of androdi-
auly.

Because the terms monaulic and diaulic
are “conditions” rather than discrete charac-
ters, the two characters that determine these
designations (external ciliated groove, inter-
nal sperm duct) were used instead. They
were not, however, treated as homologous
states of the same character (= the traditional
character “sperm duct open/closed”) based
on the following argument.

Referencing a caenogastropod, such as
Littorina (Fretter & Graham, 1994), the plesi-
omorphic open sperm groove is dorsal in po-
sition. An apparently homologous, superfi-
cial, but closed, sperm duct is found in other

PHYLOGENETICS OF CEPHALASPIDEA 405

caenogastropods [e.g., Cyclostremiscus
beauii (Fischer, 1857) (Vitrinellidae); Bieler &
Mikkelsen, 1988]. Both pass from the gonad
to the base of the copulatory organ, travers-
ing prostatic tissue along the way. The
closed internal duct in opisthobranchs (e.g.,
in Acteon, Volvatella), although more lateral in
position, follows the same course, also
traverses prostatic tissue, and is assumed
here to be homologous with that in caeno-
gastropods. Conversely, this structure can-
not be homologous with the lateral opistho-
branch external ciliated groove (e.g., in
Haminoea, Scaphander), as supposed by the
traditional character, for two reasons: (1) the
latter courses from the lateral common gen-
ital opening (distal to the gonad), forward to
the external opening (rather than the base) of
the retracted copulatory organ, without con-
tact with any prostate-type gland; and (2)
both structures—external groove and inter-
nal duct—simultaneously occur in the shelled
sacoglossans studied here, for example, As-
cobulla ulla (Marcus & Marcus, 1970) (fide Ev.
Marcus, 1972; pers. obs.). It therefore follows
that two non-homologous sperm-conducting
structures are under consideration here: (1)
the dorsal caenogastropod open sperm
groove (or closed duct), homologous with the
more-lateral opisthobranch internal sperm
duct; and (2) the lateral opisthobranch exter-
nal groove. This is in opposition to two sets of
traditional opisthobranch “dogma”: (1) the
traditional character ‘‘sperm duct open or
closed,” which assumed homology of the lat-
eral opisthobranch external groove with the
lateral opisthobranch internal duct, and (2)
the assumption that the external groove of
opisthobranchs is homologous with the ex-
ternal groove in caenogastropods.

In accordance with “the larger outgroup”
(caenogastropods), character 38 is coded as
present plesiomorphically and absent in the
derived condition. In the all-zero outgroup,
this is coded as 0 but no assumption is nec-
essary whether the homologue is in its closed
or open form.

Members of the genus Ringicula have been
noted as either androdiaulic (R. conformis;
fide Pelseneer, 1924) or monaulic (R. buc-
cinea, R. nitida; fide Gosliner, 1981; present
study). Ringicula B is here coded with the
plesiomorphic state (0), and Ringicula A with
the apomorphic state (1).

Coding: 0 = present (in Ac, RiB, Hy, Cb, As,
Vo, Ge, 00); 1 = absent (in РТА, Sc, PhA, PhB,
Cy, Ai, Bu, Ha, Sm, ReA, ReB, Ak, Ap).

39. Lateral External Ciliated Groove. As
with the previous, polarity of this character
was determined by “the larger outgroup”
(caenogastropods); the lateral (not dorsal =
character 38) ciliated groove is absent plesi-
omorphically and present in the derived
state. See comments under character 38,
above, concerning homology and the condi-
tion(s) in Ringicula.

Coding: 0 = absent (in Ac, RiB, Hy, Ge, 00);
1 = present (in RiA, Sc, PhA, РВВ, Cy, Ai, Bu,
Ha, Sm, ReA, ReB, Cb, As, Vo, Ak, Ap).

40. Copulatory Organ (Retractability). The
copulatory organ exists here in two general
conditions, retractile and nonretractile. Be-
cause the outgroup is aphallic, polarity was
determined through comparison with ‘‘the
larger outgroup.” As noted by Gosliner
(1981: 205), “the vast majority of proso-
branch gastropods” have а nonretractile
cephalic copulatory organ.

Coding: 0 = nonretractile (in Ac, Ну, 00); 1
= retractile (in all others except Ge); n (ab-
sent) in Ge.

41. Autosperm Storage Bulb. Confirmed
sperm storage areas as рай of the copulatory
organ were of two different morphologies in
the taxa examined: (a) a terminal bulb (glan-
dular) in Haminoea and Smaragdinella spp.,
and (b) as part of the autosperm intake duct
in members of Philine. Type (a) was the only
one for which homology could be considered
between taxa through positional evidence.
Terminal bulbs were also reported in the lit-
erature for Acteocina (Ghiselin, 1966; as Tor-
natina), Bulla (Er. Marcus, 1957), and Retusa
(Ghiselin, 1966; Gosliner, 1978). A glandular
ciliated terminal bulb was observed in Bulla,
but did not contain sperm; Robles (1975)
called this the terminal “caecum” of the
“prostate” in B. gouldiana Pilsbry, 1895; it
was provisionally coded as present for
autosperm storage. The same was true for
Acteocina.

Coding: O = absent (in Ac, Hy, Sc, Ph, Cy,
Cb, As, Vo, Ak, Ар, 00); 1 = present (in А!?,
Bu?, Ha, Sm, Re?); n in Ge (not applicable,
aphallic); u in Ri.

42. Ejaculatory Duct. An ejaculatory duct 1$
defined as an internal closed duct within the
penis of the copulatory organ. Depending on
species, this can be continuous with the ex-
ternal ciliated groove or with the internal
sperm duct. Ghiselin (1966: 361) “assumel[d]
that the reproductive systems with ejacula-
tory ducts are derived [polyphyletically] from
a form without an ejaculatory duct.” The po-

406 MIKKELSEN

FIGS. 43-45. Allosperm storage sacks. 43. Gametolytic gland (GG); Cylichna cylindracea (ZMUC), histo-
logical cross-section, PAS stain. 44. Receptaculum seminis (RS); Cylichna cylindracea (ZMUC), histological
cross-section, PAS stain. 45. Receptaculum seminis (RS) and bursa copulatrix (BC); Cylindrobulla n. sp.
(Bermuda, 8/1990), histological cross-section, PAS stain. Scales = 100 um.

PHYLOGENETICS OF CEPHALASPIDEA 407

larity of this character established here
agrees with this. Traditional usage, “ejacula-
tory duct open/closed,” was modified to re-
flect the two possible forms of the closed
form of the duct.

Both monaulic and androdiaulic forms are
represented in Ringicula (see above). Al-
though the androdiaulic species are here
coded 2 in Ringicula B, coding for the more
plesiomorphic (monaulic) species Я. nitida
was uncertain because the presence of an
ejaculatory duct could not be confirmed in
histological sections. Ringicula A is coded as
a
Coding: 0 = absent (т Су, Ai, Ha, Sm, ReA,
ReB, Ak, Ap, 00); 1 = present and continuous
with external ciliated groove (in Sc, PhA, PhB,
Bu); 2 = present and continuous with internal
sperm duct (in Ac, RiB, Hy, PhB, Cb, As, Vo);
n in Ge (not applicable, aphallic); u in RiA.

43. Receptaculum Seminis (Location). Two
allosperm storage sacks, one distal (bursa
copulatrix) originating near the common gen-
ital opening, and one proximal (receptaculum
seminis) originating near the gonad and com-
mon hermaphroditic duct, are characteristic
of “pentaganglionate” heterobranchs (Hasz-
prunar, 1988). In caenogastropods (e.g., Nu-
cella lapillus (Linne, 1758); fide Fretter & Gra-
ham, 1994: 49), the bursa copulatrix contains
unoriented allosperm received during copu-
lation, and the receptaculum seminis is used
for long-term storage and nourishment of al-
losperm, which are oriented with their heads
embedded in the walls of the pouch. Instead
of a bursa, most opisthobranchs have a ga-
metolytic gland (= spermatheca, bursa resor-
biens), apparently used for digestion of de-
generating eggs and sperm. Hadfield 8
Switzer-Dunlap (1984) suggested homology
with the caenogastropod bursa, although this
may be questionable. In this study, it is the
receptaculum seminis and the gametolytic
gland that are of interest.

Lemche (1956) noted important histologi-
cal differences in the cellular components of
the two sperm storage pouches in Cylichna
species. The spacious gametolytic gland
(Lemche, 1956: 138-139, fig. 243, as sper-
matheca; Fig. 43) is located in histological
cross-sections at the periphery of the body
near the heart. Its walls are composed of tall
columnar cells with prominent vacuoles, sur-
rounded by thin connective tissue. The
smaller receptaculum seminis (Lemche,
1956: 122-123, figs. 219, 221, as spermato-
cyst; Fig. 44) lies deeper within the coil near

the stomach. It is formed of low cuboidal
cells with flattened oval nuclei, surrounded
by a distinct layer of circular muscle fibers.
Schmekel (1971) verified the wall structure of
these two organs examined ultrastructurally.
These features were confirmed here in histo-
logical sections of C. cylindracea (Pennant,
1777), and were used to determine identity of
the sperm storage pouches irrespective of
their location. The presence/absence of ori-
ented sperm, albeit important, was taken as
less reliable because it can vary depending
on the physiological state of the animal at the
time of preservation.

Johansson (1954) maintained that the sin-
gle proximal allosperm pouch in Acteon tor-
natilis (Linne, 1758) was a bursa copulatrix on
the basis of previous work on ‘‘mesogastro-
pods.” In contrast, Fretter & Graham (1954)
called this a receptaculum seminis in part by
its proximal location. The latter authors ob-
served oriented sperm in the receptacular
duct, but extraneous sperm and yolk gran-
ules in the pouch lumen, and concluded that
this pouch serves the dual function of both
receptaculum and gametolytic gland. Gos-
liner (1978, 1981) considered the receptacu-
lum absent in Acteon because the single
proximal pouch appeared to be gametolytic
in function. In this study, it was determined
through the cellular criteria discussed above
that the expanded duct of the pouch qualifies
histologically as а receptaculum seminis,
with a distinct muscular layer, low cuboidal
cells, and oriented sperm. The pouch itself is
a gametolytic gland with tall columnar cells
surrounded by thin epithelium, filled with un-
oriented sperm plus other materials suggest-
ing gametolytic function. It is concluded that
this pouch is a physical as well as functional
combination of receptaculum plus game-
tolytic gland, together in a proximal location.

The receptaculum seminis in Bulla is prox-
imal but is embedded within the proximal por-
tion of the female gland mass (Robles, 1975;
confirmed here). A basal expansion of the
duct of the distal gametolytic gland was called
the seminal reservoir of the gametolytic gland
(called bursa copulatrix by Robles, 1975), but
is histologically similar to the receptaculum
and also stores closely packed sperm.

The three shelled sacoglossans in this
study (Cylindrobulla, Ascobulla, Volvatella)
have two distal sperm pouches with a com-
mon duct (Fig. 45). Both of these consistently
contained sperm and were thin-walled (of low
cuboidal cells); a Cylichna-type gametolytic

408 MIKKELSEN

gland (with tall columnar cells and amorphous
contents) is absent in all three. The slightly
larger (and deeper) of the two sacks always
contained unoriented sperm, thus functioning
as a bursa copulatrix, but differed in histolog-
ical structure (see character 45). The slightly
smaller (and closer to the common genital
duct) of the two sacks contained oriented
sperm. Its histology of low cuboidal cells sur-
rounded by muscle fibers confirmed that this
is the receptaculum seminis. Sanders-Esser
(1984: 196) concurred with this description of
the walls of sacoglossan allosperm storage
sacks; in the “bursa,” however, she noted
that “sperm and prostate secretion are dis-
solved,” indicating some gametolytic func-
tion.

The outgroup Gegania valkyrie has two
sperm pouches connected by ducts in tan-
dem and then opening by a common duct
into the pallial cavity. The two pouches agree
in histology and contents with the receptac-
ulum seminis and gametolytic gland of Cy/-
ichna, with the receptaculum closest to the
opening. Because the common duct opens in
close proximity to the vesicula seminalis or
sperm-collecting duct and far removed from
the male and female gonopores (Haszprunar,
1985b: fig. 1), the position of both sacks is
interpreted here as proximal.

Although “loss” of the receptaculum was
considered apomorphic for opisthobranchs
by Gosliner (1981), all taxa in the present
dataset had this, rendering the traditional
character “receptaculum seminis present/
absent” unusable.

Coding: O = proximal (in Ac, RIA, ВВ, Hy,
Sc, PhA, PhB, Cy, Ai, Bu, Ha, Sm, ВеА, ReB,
Ak, Ap, Ge, 00); 1 = distal (in Cb, As, Vo).

44. Gametolytic Сапа (Location). See
character 43. Polarity of this character was
established by the condition in “the larger
outgroup” (caenogastropods), making the
outgroup Gegania derived.

Because histological sections of Ringicula
were of insufficient quality to resolve this
character, Ringicula A and В are here coded
according to Gosliner (1978, 1981: fig. 1D),
who noted a distal gametolytic gland (as
bursa copulatrix).

Coding: 0 = distal (in RiA?, RiB?, Hy, Sc,
PhA, РВВ, Cy, Ai, Bu, Ha, Sm, ReA, ReB, Ak,
Ap, 00); 1 = proximal (in Ac, Ge); n (absent) in
Cb, As, Vo.

45. Novel Bursa Copulatrix. See discus-
sion under character 43. Because of different
histology [that in one caenogastropod, Cy-

clostremiscus Беаий, is of ciliated tall colum-
nar cells (Bieler 4 Mikkelsen, 1988; pers.
obs.], the “bursa” present in shelled saco-
glossans might not be homologous with that
in caenogastropods (and thus the all-zero
outgroup). For this reason, and because of
different location, the sacoglossan “bursa”
was considered a novel, derived structure.

Coding: 0 = absent (in Ac, RiA?, RiB?, Ну,
Sc, PhA, РВВ, Cy, Ai, Bu, Ha, Sm, ReA, ReB,
Ak, Ap, Ge, 00); 1 = present (in Cb, As, Vo).

46. Gonad Acini. Although a “true her-
maphroditic gland” has recently been con-
sidered characteristic (plesiomorphic?) of
Pentaganglionata (= Euthyneura; Haszprunar,
1988), Lloyd (1952) noted an evolutionary
trend in the Opisthobranchia toward separa-
tion of developing ova and sperm in the acini
or follicles of the gonad (= hermaphroditic
gland). In histological sections, acini are rec-
ognized by a covering of epithelium and mus-
cle filaments. Many heterobranchs produce
ova and sperm in the same acinus; fewer rep-
resentatives have separate male and female
acini (Hadfield & Switzer-Dunlap, 1984). In
species with separate acini, sperm-producing
acini are segregated in the “medullary zone”
or inner curve of the gonad whorl, whereas
egg-producing acini lie in the “cortical zone”
at the outer surface of the gonad (terminology
of Furrow, 1935b). Jensen (1992: 292) noted
that the gonads of most elysiid sacoglossans
are hermaphroditic and few separate, but be-
lieved that this is ““apparently not related to
phylogeny within the group.”

The acini of Philine aperta were noted as
separate male and female by Brown (1934).
Guiart (1901) earlier recorded both separate
and hermaphroditic acini. Sections examined
during this study verified the existence of
hermaphroditic acini, therefore Philine A was
provisionally coded as 0 (hermaphroditic).

Because most caenogastropods are dioe-
cious, the all-zero outgroup was here coded
un

Coding: 0 = hermaphroditic acini (т Ас,
Hy, Sc, PhA?, Ai, Bu, Ha, Sm, ReA, ReB, As,
Ak, Ap); 1 = separate male and female acini
(in RIA, Cy, Ge); u in RiB, РВВ, Cb, Vo; nin 00.

Developmental Characters: Larval develop-
ment is fully unknown in Ringicula, Sma-
ragdinella, Cylichna, and Scaphander. It is
incompletely studied in Cylindrobulla, Asco-
bulla, Volvatella, and Gegania. Because these
cases added many unknowns to the data
matrix, larval characters were not employed

PHYLOGENETICS OF CEPHALASPIDEA

409

TABLE 2. Data matrix of character states for 21 taxa and 47 characters. n = not applicable; u =

unknown. See text for explanation of character numbers.

4

Taxon 0123456789 0123456789
All-Zero 0000000nn0 00000000nn
Gegania 0000001001 00000000nn
Acteon 0000000011 100n0100nn
Ringicula А 0110000001 100n1200nn
Ringicula B 0110000001 10011200nn
Hydatina 0110100010 000u0100nn
Scaphander 0111100111 000u120111
Philine A 1111100001 000n120111
Philine B 1111100001 000u100101
Cylichna 0111100100 0001100101
Acteocina 0111100000 0001120111
Bulla 0111100200 0003000101
Haminoea 0111100200 0000000101
Smaragdinella 0111100200 0000000101
Retusa А 011110n101 nOnnnn0101
Retusa B 001110n101 nOnnnn0101
Cylindrobulla 0100111001 2100nn10nn
Ascobulla 0100111001 2112nn10nn
Volvatella 0100111001 2112nn10nn
Akera 0111110010 0000000100
Aplysia 1110100nn0 0000000100

2 3 4

0123456789 0123456789 0123456
nn00000000 0000000000 000000n
nn00010010 1004000200 nnn0101
nn00000100 1001000000 0020100
nn00001100 010101011 1440001
nn000u1000 0101u01u00 1420004
nn00010110 0u10000000 0020000
0101000000 1200112011 1010000
0101000000 u210112111 1010000
0101040000 1200012411 101000u
0101000000 1200112n11 1000001
0101040000 1200111011 1100000
0001010000 1200100011 1110000
0001001000 1200100011 1100000
000101000 1210100011 1100000
1101000100 u2001nnn11 1100000
1101000000 u210112011 1100000
nn01021101 1002012101 1021n1u
nn01011101 1002012101 1021n10
nn01011101 12u2011101 1021n1u
0011111010 1202110011 1000000
0011111010 u22u000011 1000000

in the analysis; they were compared against
the final result.

Miscellaneous Characters: Аз previously dis-
cussed (Mikkelsen, 1993, 1994), Ghiselin's
(1966) use of haploid chromosome number
suffers from incomplete data, but will be dis-
cussed below.

Complete retraction into the shell, and diet
are “conditional”” characters of traditional
use. They were not employed here because
they lack discrete homologous characters as
their basis.

Spinella, Cimino, and colleagues have in
recent years isolated a number of polypropi-
onate compounds from opisthobranchs,
many of which function as alarm phero-
mones. Included in their studies so far are
Scaphander lignarius (fide Cimino et al.,
1989), Bulla striata (fide Cimino et al., 1987),
several species of Haminoea (Spinella et al.,
1993b), and Elysia timida (Risso, 1818) (Gav-
agnin et al., 1994). Although some of the
compounds are structurally similar and their
conclusions allude to possible taxonomic ap-
plications, identical compounds extracted
from predators and their prey (Cimino et al.,
1987; Spinella et al., 1993a) will interfere with
the phylogenetic use of such data in the case

of carnivorous species. Assimilation of com-
pounds could also conceivably occur in her-
bivorous species from their algal prey.

Phylogenetic Analysis

The final data matrix appears in Table 2.

Gegania-Outgroup Analysis: Using mhen-
nig*/bb*, the Gegania-outgroup dataset pro-
duced 35 trees with a length of 111 [consis-
tency index (ci) 0.53, retention index (ri) 0.71].
Variation occurred in two areas: (a) the Bulla-
Haminoea-Smaragdinella clade(s), and (b)
the Cylichna-Acteocina-Retusa clade(s). (Re-
tusa A and B formed a consistent clade
throughout all analyses and hereafter will be
referred to merely as Retusa.)

Successive weighting produced the same
basic tree topology, with variation in the
Bulla-Haminoea-Smaragdinella clade(s), but
the Cylichna-Acteocina-Retusa arrangement
stablized. Three trees were generated with a
length of 387 (ci 0.72, ri 0.85). The weighting
process reduced seven characters (8, 9, 15,
27-28, 32, 46) to weights of 1 or 0; 15 other
characters (11-12, 16-17, 19-24, 29, 40, 43-
45) were increased to weights of 10. All other

410 MIKKELSEN

characters received final weights of interme-
diate values.

All-Zero Outgroup Analysis: Using this data-
set, including both Gegania (as an ingroup
member) and an all-zero outgroup, 245 trees
were produced with the mhennig*/bb* algo-
rithm, with a length of 116 (ci 0.50, ri 0.70) but
showing much greater variation than the pre-
vious analysis in all portions of the tree ex-
cept the Cylindrobulla-Ascobulla-Volvatella
and Scaphander-Philine clades. Most of the
variation was experienced by the mid-section
of the tree, involving the anaspids (Akera-
Aplysia) and the bulloid cephalaspids (Bulla-
Haminoea-Smaragdinella), which were also
the most variable in the Gegania-outgroup
analyses. Nevertheless, the relative se-
quence of major groups of taxa was mostly
consistent with the latter results.

Successive weighting of this dataset pro-
duced nine trees with a length of 380 (ci 0.71,
ri 0.85), with variation only in the Bulla-Hami-
noea-Smaragdinella clades. This process re-
duced eight characters to weights of 1 or 0
(8-9, 15, 25, 27-28, 32, 46); 15 other charac-
ters (11-12, 16-17, 19-24, 29, 40, 43-45)
were increased to weights of 10. All other
characters received final weights of interme-
diate values.

The Preferred Tree: Consideration of all trees
generated during this analysis lead to con-
struction of a preferred tree (Fig. 46). Deci-
sions regarding choice of included clades re-
lied on presence or absence of supporting
non-homoplastic or other strong character
state changes. Key nodes are labelled A-I for
purposes of discussion. This tree has a total
length of 117 (ci 0.50, ri 0.70). Fifteen char-
acters (11-12, 16-17, 19-24, 29, 40, 43-45)
had a ci and ri of 1, indicating total congru-
ence with the branching pattern of the tree;
three characters (8, 32, 46) had an ri of 0, and
were least congruent with the preferred tree.

Major Clades. There are seven major
clades on the tree.

(a), (b) and (c). Gegania-Acteon, Hydatina,
and Ringicula A-Ringicula В clades. These
three clades varied somewhat throughout the
analysis in relative position to one another
but always appeared in close proximity to
the outgroup. The Gegania-Acteon clade
(present in 210 of the 245 trees and all suc-
cessive weighting trees in the all-zero analy-
sis) was supported by three synapomorphies
(characters 30 and 33, left and right pallial

ganglia separate, respectfully, from left and
right pleural ganglia; 44, gametolytic gland
proximal) and one weaker character state
change (9, jaws absent). The single non-ho-
moplastic character state change, in which
the gametolytic gland is proximal rather than
distal (character 44), points out the hitherto
unrecognized fact that in both of these
cases, the receptaculum seminis and game-
tolytic gland are in tandem with the former as
a pouch on the stalk of the latter.

The Ringicula and Hydatina clades varied
in relative position, each supported by one
relatively strong character change. With Rin-
gicula adjacent to Gegania-Acteon (89 of 245
trees in all-zero analysis), mantle cavity loca-
tion (character 4) required only one change
from anterior to lateral (in Hydatina and all
following taxa). Although this topology did
not change the overall tree parameters
(length, ci, etc.), it required two additional
cases of parallel apomorphic change: (1) the
copulatory organ (character 40) becomes re-
tractile twice—in Ringicula and in all taxa fol-
lowing Hydatina—and (2) the nerve ring
moves from prepharyngeal to postpharyn-
geal twice (same locations). Just the oppo-
site is true in the preferred topology (86 of
245 trees in all-zero analysis, all successive
weighting trees, and all Gegania-outgroup
trees): the copulatory organ becomes retrac-
tile only once (and the nerve ring moves to a
postpharyngeal position only once), but the
mantle cavity shifts to a lateral position twice.
Because retractability of the copulatory or-
gan likely requires the evolution of complex
retractor muscles (as well as the penial
sheath, and other structure), it was consid-
ered more parsimonious to assume that this
occurred only once, and that the mantle cav-
Ку migrated (through differential growth, with
corresponding loss of streptoneury) multiple
times. Therefore, the preferred topology was
chosen over the alternate one.

The Ringicula А-В clade is supported by
one synapomorphy (character 31, subesoph-
ageal ganglion in B—V postion) and seven
homoplastic character state changes (9-10,
13-15, 33, 36). Ringicula À and B share a
number of character states with the taxa at
the upper end of the tree. These include sev-
eral radular characteristics in common with
all or part of the terminal clade (node F), cre-
ating homoplasies in three characters (13-
15). Two of these are reductions [13—bi-
lobed rachidian as reduction of central cusp
(that is still minutely visible in several taxa);

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PHYLOGENETICS OF CEPHALASPIDEA

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412 MIKKELSEN

15—lateral teeth reduced to single] and thus
not unlikely parallelisms. The sickle-shaped
laterals (character 14) have not been thor-
oughly examined by SEM in Ringicula and,
although superficially similar to those in
Acteocina, could reveal differences to ac-
count for the apparent parallelism. Also at
this position, Ringicula A requires parallel-
isms in two reproductive system characters
(38, internal sperm duct absent; 39, external
ciliated groove present) shared with taxa be-
yond nodes D and C, respectively. Neverthe-
less, Ringicula’s disposition at the base of
the tree is supported by seven plesiomorphic
character states [3, posterior foot absent; 4,
mantle cavity anterior (with dependent strep-
toneury); 17, gizzard absent; 23, stomach
with pouch; 30, left pallial ganglion adjacent
to left pleural; 34, genital ganglion off vis-
ceral; 35, eye direction dorsolateral], some of
which would make relocation of Ringicula to
the terminal clade unacceptable through im-
plied reversals. The reported anatomical vari-
ations within the family Ringiculidae warrant
further study as specimens from this infre-
quently live-collected group become avail-
able.

(5) Cylindrobulla - Ascobulla - Volvatella
clade, representing the Sacoglossa. This is
the most robust clade on the tree, and the only
one which did not vary in composition or rel-
ative position in all analyses. It is supported by
nine character-state changes (characters 2,
5-6, 9, 27, 33, 35-37) plus seven synapomor-
phies [oral cuticle thickened (10); descending
limb/ascus present (11); pharyngeal pouches
present (16); pedal commissure short (29); re-
ceptaculum seminis distal (43); novel bursa
copulatrix present (45); and prostate gland in
“flower” configuration (uncoded)]. The As-
cobulla-Volvatella clade is in turn supported
by two synapomorphies [tooth size increasing
within radula (12) and a dagger-shaped
rachidian (13)] and one other change (25).

(c) Aplysia-Akera clade, representing the
Anaspidea. This branch is supported by two
synapomorphies [filter chamber present
(character 22), stomach caecum present (24)]
and three other character changes (25, 28,
33)]. The larvae of Akera and Aplysia species
also share the presence of a colorless pig-
mented mantle organ (uncoded).

(d) Bulla-Haminoea-Smaragdinella clade,
representing the traditional superfamily Bul-
loidea (in part) of Cephalaspidea. Although
this group experienced much of the variation
in the results, unification into a single clade

(as seen in 71 of the 245 all-zero outgroup
trees, 2 of the 35 Gegania-outgroup trees,
and all of both sets of successive weighting
trees) was chosen on the basis of one syn-
apomorphy, the exogyrous ciliated strips
(character 7). This character was considered
sufficiently complex to hypothesize a single
evolutionary invention rather than two parallel
events. Also, a sperm bulb (character 41) is
present in this clade. Bulla is sufficiently dif-
ferent from Haminoea and Smaragdinella to
warrant union of the last two into a single
clade, with regard to rachidian shape (char-
acter 13, derived in Bulla), extent of intestinal
typhlosole (25, derived in Bulla), position of
the nerve ring (26, derived in Haminoea-Sma-
ragdinella), and presence of an ejaculatory
duct (42, in Bulla). Haminoea and Sma-
ragdinella also share two uncoded characters
(diagnostic of the family Haminoeidae) not
exhibited by Bulla: protoconch resorbed by
subsequent whorls, and the presence of
transversely ridged gizzard plates. In view of
the incongruity displayed by this group dur-
ing the analysis, additional study and deter-
mination of additional characters are war-
ranted.

Sixty-three of the 245 trees generated with
the all-zero outgroup united the Anaspidea
and Bulloidea as a monophyletic group. Sup-
port for this clade was restricted to character
21, gizzard spines present, the hypothesized
plesiomorphic state. Three other observa-
tions also support a combined clade: (a) pur-
ple “inking” behavior in both Aplysia and
Haminoea (Winner & Mikkelsen, 1986), (b)
formation of the anterior cephalic shield into
“funnels” directing water into the cephalo-
pedal groove, and (c) string-shaped egg
masses in Bulla and the anaspids (see be-
low).

(e) The terminal Cylichna-through-Philine
clade (node F), representing the traditional
cephalaspid superfamily Philinoidea, with
the addition of Retusa. The configuration of
this clade was fairly stable throughout the
analysis. It is united on the basis of one sy-
napomorphy [gizzard spines absent (char-
acter 21)] and three other strong character
changes—a bilobed rachidian tooth (13),
sickle-shaped laterals (14), and ventrolateral
eye direction (35). The strength of the radular
characters is weakened by the fact that (a)
Retusa lacks a radula (interpreted with bi-
lobed rachidian by the algorithm), and (b)
Ringicula also possesses both of these fea-
tures (see above). Deep eye location (charac-

PHYLOGENETICS OF CEPHALASPIDEA 413

ter 36, reversed to mid-depth in Acteocina)
also contributes to this clade. Within the
clade, lateral teeth are reduced to single (15,
at node G, reversed in Philine B), gizzard
plates are calcified (18, at node G, also re-
versed in Philine B), jaws are lost (9, node H),
an ejaculatory duct is present continuous
with the external ciliated groove (42, node H),
and the shell is internalized (0, node |).

The Retusa A-Retusa В clade 1$ united by
one synapomorphy (character 20, tubercu-
late gizzard plates) and two other (homoplas-
tic) character changes [9, jaws absent; 41,
sperm bulb present]. lts relationship relative
to Cylichna could not be resolved. A clade
uniting Cylichna and the two Retusa taxa was
present in 111 of the 245 all-zero outgroup
trees (plus 11 of the 35 Gegania-outgroup
trees, and all trees of both successive
weighting analyses), but was unsupported by
any strong character changes. Character
support was likewise not evident for separat-
ing the polytomy, either in the pattern Retusa
A-Retusa B above Cylichna, or Cylichna
above Retusa A-Retusa B (each present in 40
of 245 all-zero outgroup trees, plus 8 of 35
Gegania-outgroup trees).

Overall Tree Topology. The arrangement of
the major clades on the tree (Fig. 46) was
determined by many non-homoplastic (nh)
and other strong (s) character-state changes
at key nodes. At node A, the adult operculum
(character 1, s) is lost and parapodia (2, s) are
formed. At node В, the copulatory organ (40,
nh) becomes retractile, and the nerve ring
(26, s) moves from prepharyngeal to post-
pharyngeal. At node C, the stomach pouch
(23, nh) is lost leading to a flow-through con-
figuration. Here also, the mantle cavity
moves to a lateral position (4, s), the left pal-
lial ganglion separates from the left pleural
(30, s), and the external ciliated groove (39, s)
is formed.

_ At node D, the gizzard (character 17, nh) is
formed [with numerous gizzard plates (19)
and gizzard spines (21)]. The posterior foot
(3, s) is formed, the subesophageal ganglion
moves posteriorly to become associated with
the visceral ganglion (31, s), and the internal
sperm duct (38, s) with its continuous ejacu-
latory duct (42, “reversal”) is lost. At node E,
the number of gizzard plates (19, nh) changes
from many to three. Also, the marginal flexure
of the ciliated strips (7, s) occurs (later re-
versed in several taxa), the nerve ring be-
comes (secondarily) prepharyngeal (26, s, re-
versal), and the genital ganglion (34, s) joins

the visceral loop (later reversed in Philine B).
The changes associated with nodes F, G, H,
and | have been discussed previously regard-
ing the Philinoidea.

DISCUSSION
Traditional Versus Cladistic Characters

Fate of the 49 traditional characters used in
cephalaspid systematics (Mikkelsen, 1993:
table 2) have been re-examined by this study
with the following results (Table 3).

Twenty-seven characters (55%) have been
discarded from use in the present cladistic
dataset; 14 of these are unusable because
they are cladistically uncodable (DU), five are
not pertinent to the present taxa (DN), and
eight are not usable awaiting further investi-
gation (DW). Eight characters (16%) have
been used unchanged or with minor recoding
from traditional usage (Table 3, U). Fourteen
characters (28%) have been used but modi-
fied significantly from traditional usage (M).
Twenty-four cladistic characters (51%) are
new (Table 1, NEW), but six of these charac-
ters are not traditionally applied to ceph-
alaspid opisthobranchs (Table 1, NON).

Consistently strong or weak characters
can be determined from resultant weights in
the two successive weighting runs as well as
from character ci and ri values on the pre-
ferred tree. Common strong characters in all
three trials were: ascus (character 11), tooth
size (12), pharyngeal pouches (16), esoph-
ageal gizzard (17), gizzard plate number (19),
tuberculate gizzard plates (20), gizzard
spines (21), filter chamber (22), stomach (23),
stomach caecum (24), pedal commissure
(29), copulatory organ (40), receptaculum
seminis (43), gametolytic gland (44), and
novel bursa copulatrix (45). Most of these
contributed only to individual clades, for ex-
ample, characters 11-12, 16, 29, 43, and 45
to Sacoglossa. The most important charac-
ters to the overall preferred tree structure
were copulatory organ retractile (character
40), and characters 17, 19, 21, and 23, all of
which pertain to the gizzard and stomach.
Consistently weak characters were similarly
determined: pallial caecum (character 8), su-
praesophageal ganglion (32), and gonad
acini (46), each showed ri values of zero in all
three analyses.

414

Character Evolution

MIKKELSEN

TABLE 3. Fate of traditional cephalaspid characters (Mikkelsen, 1993), including
character numbers from the present data matrix. DN = discarded as not pertinent in
the current dataset (autapomorphies, etc.); DU = discarded as uncodable for use in
cladistic analysis; DW = discarded awaiting further study; М = modified; U =
unchanged.

Character Fate No.
SHELL present/absent DN

SHELL external/internal U 0
SHELL thick/thin DU

SHELL not reduced/reduced DU

SHELL exposed spire/involute/plate-like DU

OPERCULUM present/absent U 1
CEPHALIC PROCESSES (form) DU

PARAPODIA absent/weak/strong M 2
FOOT short/long DU

POSTERIOR PALLIAL LOBE absent/present U 3
HANCOCK’S ORGAN weakly/strongly developed DU

HANCOCK’S ORGAN (form) DW

LIP ORGANS (form) DW

MANTLE CAVITY (position) M 4
GILL present/reduced/absent M 6
GILL nonplicate/plicate DN

OSPHRADIUM not reduced/reduced DW

OSPHRADIUM (form) DW

PALLIAL CAECUM short/long/absent M 8
JAWS present/absent U 9
JAWS armed/unarmed DU

RADULA present/absent DN

RADULA (form) M 14
RADULAR TEETH many/few M 15
RACHIDIAN present/absent M 13
SALIVARY GLANDS 1/2 pair DN

ESOPHAGEAL DIVERTICULUM absent/present DW

GIZZARD weakly/strongly developed M 17
GIZZARD PLATES absent/present M 17
GIZZARD PLATES (number) U 19
GIZZARD PLATES (form) M 18, 20
NERVOUS SYSTEM streptoneurous/euthyneurous DU

NERVE RING prepharyngeal/postpharyngeal U 26
NERVE RING GANGLIA 6/4 M 2
VISCERAL LOOP long/short DU

VISCERAL LOOP GANGLIA 5/4/3 DU

EYES present/absent DN

REPRODUCTIVE SYSTEM monaulic/diaulic M 38, 39
SPERM DUCT open/closed DU

COPULATORY ORGAN nonretractile/retractile U 40
SPERM BULB absent/present U 41
EJACULATORY DUCT open/closed M 42
PROSTATE pallial/penial DW
RECEPTACULUM SEMINIS present/absent M 43
PALLIAL GONODUCT simple/divided DW

GONODUCT pallial/coelomic DU

CHROMOSOME NUMBER 12/17 DW

RETRACTION complete/incomplete DU

DIET carnivorous/herbivorous DU

Several evolutionary scenarios were im-
plied by the trees generated during the anal-
yses and are preserved in the preferred tree.

(a) Formation of the gizzard. The gizzard
(character 17) is a synapomorphy of all taxa
beyond the Sacoglossa clade (Fig. 46, node
D). In its first appearance in the herbivorous

Г) x 1996

PHYLOGENETICS OF CEPHALASPIDEA _ 415

anaspids, it has numerous plates and gizzard
spines (as earlier proposed by Gosliner,
1994, upon which coding of these characters
was based) and is followed by a filter cham-
ber preceding the stomach. Beyond the
Anaspidea (node E), the gizzard plates con-
solidate into three large plates (character 19).
In the primarily herbivorous Bulloidea clade,
the gizzard spines are retained. These spines
are lost at node F (character 21). Beyond
node F, innovations in the gizzard occur: giz-
zard plate tubercles (in Retusa), and calcifi-
cation (п Acteocina, Scaphander, Philine А)
with the corresponding switch to obligate
carnivory (see discussion under Diet, below).

(b) Internal sperm duct versus external cil-
iated groove. The preferred tree reitterates
assumptions made about homologies of
these two structures in the Characters and
Coding section (character 38, above). It was
assumed that the external ciliated groove
present in caenogastropods is homologous
with the closed internal sperm duct of heter-
obranchs (in Gegania, Acteon, Hydatina, Rin-
gicula B, and the sacoglossans). The saco-
glossans show the first appearance of the
opisthobranch-type external groove (charac-
ter 39 at поае С) in conjunction with a still-
present internal duct. Beyond the sacoglos-
sans (node D), the internal sperm duct
(character 38) is lost, and the external ciliated
groove takes over as the sperm conduit.
Observations by Hadfield & Switzer-Dunlap
(1984) support this hypothetical transition,
wherein the ciliated groove first functions to
transport eggs in the Sacoglossa, both eggs
and autosperm in Anaspidea, then autosperm
only in Cephalaspidea. The evolutionary hy-
pothesis is, then, that the internal sperm duct
ishomologous with the caenogastropod open
ciliated groove, but that the ciliated groove
of opisthobranchs is a novel structure. Ho-
moplasy in these characters occurs in Ring-
icula A, based on Я. nitida, which is monaulic
with a confirmed external ciliated groove.

(c) Allosperm storage sacks (receptaculum
seminis, gametolytic gland, novel bursa cop-
ulatrix). As mentioned previously, the pre-
sumed plesiomorphic condition for opistho-
branchs is a proximal receptaculum seminis
and a distal bursa copulatrix. The distal ga-
metolytic gland in most taxa examined here
may be formed from (and thus homologous
with) the bursa (see discussion of character
43). According to the preferred tree, the re-
ceptaculum (character 43) remains proximal
in all taxa except the sacoglossans, in which

it moves to a distal position: The gametolytic
gland (character 44) remains distal in all taxa
except Acteon and Gegania, in which it has
moved proximally to unite with the receptac-
ulum in tandem; in both taxa the receptacu-
lum is a sack along the stalk of the game-
tolytic gland. The plesiomorphic bursa is not
coded here; the “bursa” of the sacoglossans
is unique histologically and appears to be a
novel structure.

(d) The visceral nerve loop formula devel-
oped here contradicted the previously held
view in placement of the pallial ganglia in
their most plesiomorphic form: LA- -B-V-P-
-AL (rather than the classic L-A-B-V-P-A-L).
The most critical problem here 1$ the lack of
incontrovertible evidence for position of the
pallial ganglia through innervation of specific
organs; pallial ganglia innervate the wall of
the mantle cavity, as do most other ganglia.
Nevertheless, most taxa could be reliably
coded based on the presence and position of
other ganglia.

The preferred tree supports the proposed
fused/adjacent position of the pallials and
pleurals as plesiomorphic. The left pallial
(character 30) is fused/adjacent in the out-
group, Ringicula, and Hydatina. It becomes
well separated twice, once in Acteon and Ge-
дата, and once at node С. The right pallial
(33) is fused/adjacent throughout most of the
tree, with two changes to separate (A-L con-
figuration) in the Acteon-Gegania and Ring-
icula clades, and two changes to the PA con-
figuration in the sacoglossans and the
anaspids (with Aplysia assigned by the algo-
rithm).

Homoplasy: Not surprisingly, but now with
supporting evidence, parallelism is indeed
“rampant” in the opisthobranchs, at least in
those investigated by this study. Twenty-five
characters undergo at least one parallel apo-
morphic change on the preferred tree. Signif-
icant parallelism (three or more identical
changes) occurred in six characters: elonga-
tion of a pallial caecum (character 8; four
times), loss of jaws (9; five times), loss of in-
testinal typhlosole (25; five times), fusion of
the cerebropleural ganglia (27; five times),
shortening of the cerebral commissure (28;
three times), and supraesophageal ganglion
migration to V- -P position (32; four times).
Reversals were also common, occurring
nine times in eight characters. Some of these
are logical from our knowledge of the biology
of these animals. For example, the adult

416

operculum (character 1) is “reinvented” т
Retusa B; this is not unexpected because
most opisthobranchs still possess larval
opercula and the “reversal” in the tree is
merely ontogenetic retention of a structure
never completely lost. Also, reversal to ab-
sence of the posterior foot (character 3) in
Aplysia could be a consequence of great en-
largement in body size. Others can be inter-
preted as true reversals, for example, a sec-
ondarily prepharyngeal location of the nerve
ring (character 26).

Implications for Cephalaspid Classification

It is a natural response following this type
of analysis to extend the conclusions to a
revision of taxonomic classification. The fol-
lowing arrangement is supported by the pre-
ferred tree:

? “Lower

Acteon
Class Opisthobranchia:
2 “Architectibranchia’’: Hydatina,
Ringicula
Order Sacoglossa: Cylindrobulla,
Ascobulla, Volvatella
Order Anaspidea: Akera, Aplysia
Order Cephalaspidea:
Superfamily Bulloidea: Bulla,
Smaragdinella, Haminoea
Superfamily Philinoidea: Cylichna,
Retusa, Acteocina, Scaphander,
Philine

Several qualifications must be mentioned.
First, Acteon, Hydatina, and Ringicula have
been here removed from Cephalaspidea by
this investigation. Haszprunar (1985a) placed
all three in the paraphyletic pentaganglionate
(= opisthobranch) group “Architectibran-
chia.” This analysis placed one of the three
taxa, Acteon, clearly in a clade with the lower
heterobranch (= non-opisthobranch) Gega-
nía. Although both of these larger groups
(“Architectibranchia” and “Lower Hetero-
branchia”) are at present unresolved, | have
provisionally used them here to avoid naming
new higher groups based on these prelimi-
nary results. Unfortunately, several presum-
ably important sources of characters (e.g.,
Hancock's organ, osphradium, female gland
mass, sperm structure) had to be excluded
from this analysis for lack of cladistically
sound data; their eventual inclusion (clearly
objectives for future work) could define the
limits of Opisthobranchia and resolve the in-
teresting basal groups.

Heterobranchia”: Gegania,

MIKKELSEN

It is interesting to note the single synapo-
morphy uniting Geganía and Acteon — char-
acter 44, gametolytic gland in proximal posi-
tion, in tandem with the receptaculum seminis
as a sack on its stalk. Haszprunar (1985b, c)
noted the same “receptaculum apparatus” in
two other lower heterobranchs: the mathildid
Opimilda maoria Powell, 1940, and the archi-
tectonicid Heliacus variegatus (Gmelin, 1791).
This character could prove with further study
to be important in the phylogeny of lower het-
erobranchs.

Relative to previous classifications, the
Sacoglossa and Anaspidea, both well-recog-
nized, non-cephalaspid clades, are here each
confirmed as monophyletic with synapomor-
phies as outlined earlier. The long list of strong
characters supporting the sacoglossan clade
correspond well to previous lists of sacoglos-
san traits [e.g., suctorial feeding apparatus,
sacoglossan radula (Kay 1968: 20)]. These
findings confirm the placement of Cylindro-
bulla in the Sacoglossa [as advocated by
Thompson (1976), Gosliner (1994), and others
(see Mikkelsen, 1993, 1994)], based on the
type species, С. Беаий, and a new species
from Florida. Further discussion will be pre-
sented in a manuscript in preparation. These
results require redefinition of the order Saco-
glossa as distinguished by the above synapo-
morphies for this clade. Most notably, dag-
ger-shaped radular teeth are no longer
prerequisite for membership in the order.

More germane to the present problem, the
demise of traditional Cephalaspidea has here
been confirmed by this analysis. With
Acteon, Ringicula, and Hydatina now ге-
moved to more basal groups, the remaining
Cephalaspidea survived the analysis and can
be considered monophyletic. Synapomor-
phies for this group (at node E) are: ciliated
strips flexed at mantle margin (character 7),
three gizzard plates (19, non-homoplastic),
the nerve ring secondarily prepharyngeal
(26), and the genital ganglion on the visceral
loop (34). Characters 7, 26, and 34 experi-
ence reversals within the clade.

Within Cephalaspidea, there are two
monophyletic clades, the Bulloidea and the
modified Philinoidea. Bulloidea is supported
by one non-homoplastic synapomorphy
(character 7, exogyrous ciliated strip) and the
weaker character, presence of a sperm bulb
(41). Philinoidea is based on one non-ho-
moplastic character—gizzard without gizzard
spines (2)—and four homoplastic characters:
bilobed rachidian teeth (13), sickle-shaped

PHYLOGENETICS OF CEPHALASPIDEA 417

lateral teeth (14), ventrolaterally directed eyes
(35), and deeply located eyes (36).

The position of Retusa warrants special
comment. Superficially, the living animals of
Retusa spp. closely resemble those of
Acteocina or Cylichna, but Retusidae has
been most often grouped in Bulloidea or at
least near Bullidae/Haminoeidae (Mikkelsen,
1993, 1994). The basis for this is difficult to
determine because Bulloidea, like most
cephalaspid superfamilies, has rarely been
defined. Among the few examples, Steinberg
(1963) and Keen (1971) provided: shell exter-
nal, aperture as long as the shell, and para-
podia absent. The first of these 15 clearly ple-
siomorphic, the second also pertains to
forms such as Philine and Hydatina, and the
last is in error. Ghiselin (1966) included Re-
tusidae in Bulloidea by the presence of a
sperm bulb (character 41), a character provi-
sionally assigned but not confirmed here.
Therefore, traditional placement in Bulloidea
(which never occurred during this analysis) 1$
weaker than the evidence provided here for
placing Retusa in Philinoidea. Unknown char-
acter states assigned to Retusa by the algo-
rithm through its membership in Philinoidea
involve location of the left pallial ganglion (30,
which could not be determined), configura-
tion of the eye (35-37, absent in Retusa A),
and the appearance of the radula (11-15, as-
signed by the algorithm with bilobed rachid-
ian and sickle-shaped laterals prior to its pre-
sumed loss). Gosliner (1978) also advocated
philinoidean membership for Retusidae
based on carnivory and configuration of its
reproductive system.

Outgroup Choice: Gegania Versus
Hypothetical Ancestors

Although the validity of ‘“‘hypothetical an-
cestors’”’ can be debated, the hypothetical
all-zero outgroup solved problems in this
study not resolved by the use of a bona fide
outgroup. Because Gegania valkyrie is a real
animal, it has derived features of its own and
is even missing suites of characters impor-
tant in this analysis (e.g., a copulatory organ).
This will always be especially true within the
Heterobranchia, in which evolutionary re-
placement of classic gastropod characters
with experimental novelties seems ubiqui-
tous. One might question whether Gegania,
or in fact any of the lower heterobranchs, was
an appropriate choice for an outgroup. (The
arguments against using a pulmonate for this

analysis were discussed earlier.) The lower
heterobranchs have been considered by
most authors (Haszprunar, 1988; Bieler,
1992; Healy, 1993) to be a paraphyletic
grade, yet it seems likely that at least some of
the included families lie in sister-group rela-
tionship to the Opisthobranchia. Whether
they comprise one monophyletic group or a
dozen is immaterial here; the best choice for
an outgroup in opisthobranch studies still
rests within the lower heterobranchs. Which
taxon is chosen depends largely on available
material and the characters being tabulated.
Based on sperm ultrastructure, Healy (1993)
concluded that several alternate superfami-
lies (Pyramidelloidea, Rissoelloidea, Omalo-
gyroidea) were closer to the Opisthobranchia
than the Architectonicoidea (containing Ge-
gania). So, for sperm studies, a pyramidellid
might be the best choice. However, for rea-
sons discussed earlier, a pyramidellid was
not chosen for this anatomy-based work.
One unanticipated result of the all-zero
outgroup analysis was that Gegania joined
the ingroup, forming a clade with Acteon.
Some might interpret this as evidence that
Gegania was a poor choice for an outgroup.
| prefer to suggest that this result showed
that Acteon quite possibly does not belong to
the Opisthobranchia, and that the ingroup
should have been considered the Hetero-
branchia. In this case, a caenogastropod (i.e.,
that upon which the all-zero outgroup was
based) was a better outgroup choice. In this
interpretation, a pulmonate would have again
been eliminated as a possible choice.
Gosliner (1978, 1981, 1994) proposed a
Hypothetical Ancestral Opisthobranch
(HAO), the features of which can now be
compared against the results of this study.
The HAO agrees with the all-zero outgroup
used here (in those characters discussed)
with one major exception. Although Gosliner
assumed а nonretractile copulatory organ
(although he specified a ‘‘non-protrusible pe-
nis,” his figures show а nonretractile organ),
the plesiomorphic state for opisthobranchs
as proposed here, no internal sperm duct 1$
indicated. Coupled with this, the HAO has an
external ciliated groove, which although most
common in the ingroup, has been suggested
here as derived. In addition, ‘‘prostatic secre-
tions are added from glandular epithelial cells
lining the [open, ciliated] sperm groove”
(Gosliner, 1981: 222), a condition reminiscent
of that in caenogastropods, such as Littorina
(Fretter 8 Graham, 1994), but one not seen or

418 MIKKELSEN

proposed here. Gosliner apparently per-
ceived the open seminal groove of caeno-
gastropods homologous with that in opistho-
branchs—dogma not supported by these
results. In nervous system configuration, the
HAO has a visceral nerve loop formula (L- -
B-V-P--L) compatible with this study,
wherein the right and left pallial ganglia were
not indicated by description nor by figures;
although Gosliner did not discuss this issue,
we can presume that the pallials are, as pro-
posed here, still fused with the pleurals.

Additional Character Mapping

Some available data, although too incom-
plete for inclusion in the analysis, were suffi-
cient for comparison against the preferred
tree.

Chromosome Number: Haploid chromo-
some number has been claimed as conser-
vative in the Euthyneura (Burch & Natarajan,
1967; Patterson, 1969; Schmekel, 1985).
Ghiselin (1966) advocated a clade containing
Anaspidea, Sacoglossa, and Cephalaspidea
(in part), supported only by weak evidence of
a haploid chromosome number of 17 or 18.
Haszprunar (1985a) perceived this as the ple-
siomorphic state (16-18 in Gymnomorpha
and Archaeopulmonata; 17 in lower hetero-
branchs and Cephalaspidea), which is re-
duced to 12-13 in the Eleutherobranchia
(= Notaspidea + Nudibranchia). In his non-
cladistic phylogenetic hypothesis of Euthy-
neura, Salvini-Plawen (1991a: fig. 15; 1991b)
considered 16 as the most plesiomorphic
state for the Euthyneura, 17 for Opisthobran-
chia, and a reduction to 13 for three groups
of traditional Nudibranchia.

Considerable progress has been made in
this field since Ghiselin’s 1966 paper. Within
the group under study here, 16-18 haploid
chromosomes are characteristic of all taxa
above node С in the preferred tree. This in-
cludes: no records for the included shelled
sacoglossans, but many unshelled forms (17;
Inaba, 1959; Patterson, 1969); numerous
records for Aplysia and other Aplysiidae (16-
17; Inaba, 1959; Burch & Natarajan, 1967;
Patterson, 1969; Natarajan, 1970; Vitturi et
al., 1985); one species of Bulla (17; Vitturi et
al., 1985); six species of Haminoea and the
related Cylichnatys (16-18; Inaba, 1959;
Burch & Natarajan, 1967; Patterson, 1969;
Natarajan, 1970; Vitturi et al., 1985); one spe-
cies of Smaragdinella (18; Burch & Natarajan,

1967); one species of Scaphander (18; Vitturi
et al., 1985); and three species of Philine (17-
18; Inaba, 1959; Patterson, 1969; Vitturi et
al., 1985). Node C is the equivalent to that
based on chromosome data on Ghiselin's
(1966: fig. 7, node 5) tree. Below this node,
only one species, Hydatina velum, has been
studied and found with 15 chromosomes
(Natarajan, 1970). Although Natarajan (1970)
considered this state “primitive,” Butot & Ki-
auta (1969) hypothesized that reduction in
chromosome number in stylommatophoran
land snails was derived [as also occurs in
Nudibranchia (13) and Notaspidea (12-13);
Vitturi et al., 1985]. Philinoglossa praelonga
Salvini-Plawen, 1973, a sand-nudibranch
grouped with Retusa by Ghiselin (1966),
showed only 13 pairs (Curini-Galletti, 1985).

Within the lower heterobranch outgroup,
Valvatidae show 9-10 and Pyramidellidae 17
(Furrow, 1935a; Patterson, 1969), but only
two species have been examined so far.

When mapped onto the preferred tree, the
pattern of these data supports Ghiselin's
node 5 character change, however, chromo-
some number does little to support or refute
the component clades. А chromosome пит-
ber of 18 appears on the tree only past node
E, perhaps establishing this number for
Cephalaspidea $.5.

Chemosensory Organs: Edlinger (19805)
constructed a phylogenetic hypothesis of
shelled opisthobranchs based on Hancock's
organ, lip organs, and the osphradium. These
conclusions can now be compared to the
preferred tree from this study. In his single-
organ schemes, Haminoea was considered
most plesiomorphic for the osphradium and
lip organs, and Acteon for Hancock's organ.
Scaphander and Philine comprised a derived
group in all characters, consistent with their
terminal position on this preferred tree. Bulla
was consistently close to Haminoea, but var-
ied in relationship: identically plesiomorphic
in lip organs, derived in osphradium, more
plesiomorphic in Hancock's organ. Acteon
was considered derived in osphradium and
lip organs. Retusa was derived in lip organs,
but was only superficially considered for os-
phradium (“reduziert’’) and Hancock's organ
[“eine Ähnlichkeit mit Bullaria” (= Bulla)].
Edlinger’s (1980b: fig. 9) composite “Еуо-
lutionsschema” was based on his morpho-
logical data (Edlinger, 1980a) and those of
earlier authors, especially Boettger (1955)
and Salvini-Plawen (1970). It agrees with the

PHYLOGENETICS OF CEPHALASPIDEA 419

preferred tree here in four points: (a) Acteon
is nearest the base of the tree, (b) the Saco-
glossa follow, (c) Scaphander and Philine are
in one clade, and (d) Bulla and Haminoea lie
in close proximity. It differs from the preferred
tree in placing Retusa with Bulla, and the
Anaspidea in a clade with the Philinoidea,
rendering the latter paraphyletic. Although
these results lend support to some aspects
of the preferred tree, and refute others,
Edlinger's conclusions were not cladistically
generated, and his morphological data are
not readily translated into codable informa-
tion.

Larval Development: Larval development
type (i.e., planktotrophic, lecithotrophic, di-
rect-developing) was not considered here,
because evidence suggests a high incidence
of homoplasy throughout the Gastropoda.
Planktotrophic development is considered
more plesiomorphic (Strathmann, 1993), evi-
denced by (a) its omnipresence throughout
the class (most common generally indicating
most plesiomorphic), and (b) the presence of
vestigial veliger structures (e.g., velar lobes,
used in locomotion and feeding in swimming
larvae), even in encapsulated embryos later
hatching as benthic, crawl-away juveniles.
Nevertheless, non-feeding larvae (lecitho-
trophic and direct-developing, evidenced
also by bulbous protoconch whorls) are
known throughout the class, even in “primi-
tive” groups (Jablonski, 1986). Direct devel-
opment occurs in members of four genera
involved here [Acteocina (Mikkelsen & Mik-
kelsen, 1984), Retusa (Smith, 1967), Cylin-
drobulla (D. De Freese, pers. comm.), and
Ascobulla (D. De Freese, pers. comm.)], but
in none are all known members of the genus
direct-developing.

Two alternate sources of data related to
larval development were considered here:
form of the egg mass and larval morphology.

Egg Mass Form. Most opisthobranchs de-
posit gelatinous egg masses attached to the
substratum by some means. Study is facili-
tated by the fact that most species readily
(and conveniently) shed an egg mass (fertil-
ized or not) as a stress reaction to the labo-
ratory environment. Although some egg
mass characters were studied here, relative
number of eggs per mass was not used, due
to high variability within genera dependent
upon size and condition of the adult, larval
development type, and other factors.

Form of the egg mass has been catego-

rized by Soliman (1987) and by Hadfield &
Switzer-Dunlap (1984). Three different forms
were evident in the taxa under consideration:
(a) elongated string (round or oval in cross-
section) in Bulla (pers. obs.), Cylindrobulla (D.
De Freese, pers. comm.), Volvatella (K. B.
Clark, pers. comm.), Akera (pers. obs.), and
Aplysia (pers. obs.); (b) flattened ribbon in Hy-
datina (pers. obs.); and (c) gelatinous ball
(ovoid to globular) in Acteon (Fretter & Gra-
ham, 1954), Philine (Tchang Si, 1931; Lloyd,
1952), Acteocina (Mikkelsen 8 Mikkelsen,
1984), Retusa (Smith, 1967; Berry, 1989), As-
cobulla (K. B. Clark, pers. comm.), and Ge-
дата (Climo, 1975).

Egg mass variability 1$ evident within Hami-
noea. Haminoea elegans and H. antillarum
deposit unanchored coiled ribbons attached
to the substratum along one long side; H.
solitaria (fide Smallwood, 1903; Harrigan 4
Alkon, 1978) and H. succinea (Conrad, 1846)
(pers. obs.) produce anchored gelatinous
balls, sometimes elongated into cylinder-
shaped structures; H. zelandiae Gray, 1843,
forms unanchored sausage-shaped masses
attached to vegetation (Rudman, 1971a).

Many opisthobranchs anchor their egg
masses to the substratum via a long mucous
string. This occurs here mainly in gelatinous
ball masses [in Acteon (Fretter & Graham,
1954), Philine (Tchang Si, 1931; Lloyd, 1952),
Acteocina (Mikkelsen & Mikkelsen, 1984),
and some Haminoea (see above)], but is also
true of the egg ribbon in Hydatina (pers.
obs.).

The species in this analysis all deposit
eggs encapsulated within gelatinous egg
masses. (The egg capsule is here the egg
membrane surrounding each embryo and its
albumen supply; it is not homologous with
the resistant, chitinous, external capsules
produced by neogastropods, e.g., Busycon.)
In most cases, each capsule contains one
egg, however several taxa are known to nor-
mally encase more than one egg per capsule
[Hydatina (20-30 eggs/capsule, pers. obs.),
Aplysia (3-12; Tchang-Si, 1931; pers. obs.)].
Because these two are among the largest of
opisthobranchs, this could be an adaptation
to increased body size.

Although Hadfield 4 Switzer-Dunlap (1984:
278) claimed that the number of eggs per
capsule “probably has little significance as a
taxonomic tool,” this reference was to the
actual number among multiple-egg species,
known to vary with age of the adult or posi-
tion within the mass. The latter type of vari-

420 MIKKELSEN

ability has been noted in several taxa. Tchang
Si (1931) considered the normal condition in
Philine aperta to be one egg/capsule, but
noted that 2-3/capsule were found in en-
larged capsules near the end of the egg
string. In P. gibba, Seager (1979) noted mul-
tiple embryos as rare and usually associated
with capsule abnormalities. Haminoea antil-
larum (and others, e.g. H. solitaria, fide Har-
rigan 8 Alkon, 1978) produces encapsulated
single eggs, whereas two eggs per capsule
were reported by Berrill (1931) for H. hydatis,
and Bandel (1976: 94) noted 1-3 eggs/cap-
sule in H. antillarum, although single was
given as “the usual” condition. Bulla striata
examined for this study were consistently
one egg/capsule (with a rare two/capsule in
normally single masses), but as many as 25/
capsule were recorded in B. gouldiana by
Robles (1975).

In spite of the abundance of egg mass
data, few patterns are observable when
mapped on the preferred tree. Gelatinous ball
masses, anchored or unanchored, range the
entire length of the tree, so this (= the most
common) could be the plesiomorphic state
(agreed by Gosliner, 1994). Egg strings are
concentrated mid-tree, between nodes C
and F, however the only clade for which this
is exclusive is the Anaspidea.

Larval Morphology. The premetamorphic
larvae of the taxa under consideration are
remarkably well studied (Thorson, 1946;
Thompson, 1976; and others) and extremely
uniform in morphology. Only one character
was considered appropriate for possible use
here, the pigmented mantle organ (= “larval
kidney”” of authors).

The pigmented mantle organ (PMO; = lar-
val kidney, secondary kidney, anal glana), lo-
cated in the mantle near the anus on the right
side of the veliger, is an important taxonomic
feature of all heterobranchs. Its relation to ex-
cretion, however, is doubtful (Robertson,
1985). Bonar & Hadfield (1974) suggested in-
volvement in accumulation of metabolic
wastes, if not in excretion. Bickell & Chia
(1979) described PMO cells appearing secre-
tory in nature. The PMO has been suggested
as repugnatory in Pyramidellidae (J. B. Wise,
pers. comm.). Although the PMO is usually
“lost” at metamorphosis, it persists in mem-
bers of the genus Philine for at least a short
period after larval life (Horikoshi, 1967), and
throughout adult life in other heterobranchs
(e.g., Pyramidellidae; J. B. Wise, pers.
comm.).

Robertson (1985) noted that the PMO color
(perhaps reflecting biochemical makeup) is
often genus- or species-specific, and thus of
taxonomic use. The color of the PMO was
often recorded in published larval develop-
ment accounts, facilitating its use here. Rob-
ertson (1985: 6) noted that PMO color could
be extremely variable in such families as
Pyramidellidae, and that “there is no basis
for a claim that the color diversity reflects
non-homology.” Likewise one might also
postulate that similar color does not neces-
sarily imply homology, although Edlinger
(1980b) suggested that similar PMO color in
Philinidae, Philinoglossidae, and Haminoei-
dae implied close phylogenetic relationship.
A change in color during ontogeny occurs in
Philine (Thorson, 1946) and Haminoea (pers.
obs., H. succinea). For these reasons, and
because PMO color was often ascertained
from published black-and-white illustrations,
dark and light were used here instead of ac-
tual hue. Dark PMOs (black, red, or orange)
were found in Hydatina (black, pers. obs.),
Philine (red to jet black; Thorson, 1946; Hori-
koshi, 1967), Bulla (dark red, pers. obs.),
Haminoea (red to black, pers. obs.), and Cy-
lindrobulla (orange, pers. obs.). Clear PMOs
have been recorded in Acteon (figured only
as lightly pigmented by Thompson, 1976),
Acteocina (pers. obs.), Akera (figured as col-
orless by Thorson, 1946: fig. 148B, C), and
Aplysia (Thompson, 1976; Kriegstein, 1977;
Switzer-Dunlap & Hadfield, 1977). Retusa
was the only taxon known to exhibit both
character states: dark (R. obtusa; fide Smith,
1967) and light (R. truncatula; fide Rasmus-
sen, 1944). No correlation exists between
type of larval development and PMO color.

Hurst (1967: 279) suggested that a darkly
pigmented PMO is a feature of “the more
primitive opisthobranchs,” suggesting polar-
ity. This is corroborated by colorless РМО$ in
the nudibranchs Phestilla sibogae Bergh,
1905 (Bonar & Hadfield, 1974) and Doridella
steinbergae (Lance, 1962) (Bickell & Chia,
1979). When mapped on the preferred tree,
however, no pattern emerges; clear and dark
PMOs are equally and evenly distributed. As
with egg mass type, the only clade to be sup-
ported is the Anaspidea, with clear PMOs.

Sperm Morphology: Ghiselin (1966) summa-
rized light microscopical observations on
opisthobranch spermatozoa up to time of his
publication, noting variation in the length of
the sperm head (= nucleus; short or long),

PHYLOGENETICS OF CEPHALASPIDEA 421

with short assumed to be the plesiomorphic
state. Healy 8 Willan (1984) explained that
the “long” form was actually a cord-shaped
nucleus helically wrapped around the ax-
oneme. When Ghiselin's (1966) compiled nu-
clear length data, plus additions from subse-
quent literature (Thompson & Bebbington,
1969; Thompson, 1973; Healy, 1981), are
mapped on the preferred tree, little or no pat-
tern emerges. Short nuclei are characteristic
not only of Acteon and lower heterobranchs
(here, Pyramidellidae), but also of Philine and
others. Short and long forms have also both
been documented in Nudibranchia and No-
taspidea (Healy & Willan, 1984, 1991). It 1$
apparent that more sophisticated techniques
will be needed if sperm data are to be useful
here.

For the past decade, Healy (1981, 1982,
1987, 1988a, b, 1993; also Healy & Willan,
1984, 1991) has been amassing ultrastruc-
tural data on sperm and spermiogenesis of
higher gastropods for eventual application to
phylogenetic studies. Healy (1982, 1988a, b)
determined that sperm morphology (at the ul-
trastructural level) of Architectonicidae and
Pyramidellidae corroborated their relation-
ship with opisthobranchs and their position in
Heterobranchia. Healy 8 Willan (1991) exam-
ined sperm from all four superfamilies of
nudibranchs; although no autapomorphies
for Nudibranchia were identified, sperm char-
acters proved useful in indicating familial re-
lationships. In these and other studies, the
following characters have been suggested as
worth investigating for phylogenetic pur-
poses: midpiece membranes (Healy & Willan,
1991), acrosomal morphology (Healy & Wil-
lan, 1991), presence/absence of glycogen
piece (Healy & Willan, 1984), and number of
midpiece helices (Healy, 1988b). Similar in-
formation for Cephalaspidea is not yet avail-
able.

Fossil Record: Because of their calcareous
shells, mollusks have one of the best fossil
records of any invertebrate group. Given that
the evolutionarily oldest taxa should be the
first to branch off a cladogram, comparison
of the stratigraphic distributions of organisms
at, for instance, the family level can provide
an independent test of a morphologically
based phylogeny. This of course assumes
that a relatively complete fossil record exists,
at least representative of actual history.
Because of their reduced, thin-walled
shells, shelled opisthobranchs do not have

as rich a fossil record as most groups of mol-
lusks. Therefore, although occurrence
records for shelled opisthobranchs can be
taken at face value, distributional gaps in the
fossil record are not guaranteed indications
of absences of taxa.

For our purposes here, we are searching
for the oldest fossil record for each taxo-
nomic unit to compare against the preferred
tree [although Schoch (1989: 211 ff.) summa-
rized the arguments against such an exer-
cise, e.g., first occurrences of taxa can only
be objectively compared at a single location].
Tracey et al. (1993) provided a convenient
and comprehensive summation of oldest
records of gastropods at the family level.
They reported that the outgroup family,
Mathildidae, indeed showed the longest fos-
sil record of taxa on this tree, extending into
the Permian [Guadalupian stage; 270 MYBP
(= million years before present)]. Acteonidae
and Aplustridae were the oldest recorded
shelled opisthobranchs under consideration
here, with records in the Jurassic (Sinemurian
stage; 180 MYBP). Other Jurassic taxa were
Bullidae (Pliensbachian stage), Retusidae
(Bathonian), Akeridae (Callovian), and Acteo-
cinidae (Kimmeridgian). Moving forward in
time to the Cretaceous (135 MYBP), the old-
est was Ringiculidae (Neocomian series), fol-
lowed by Cylichnidae (Santonian) and Hami-
noeidae (Maastrichtian). The remaining taxa
with fossil records did not extend past the
Tertiary: Philinidae and Sacoglossa (oldest
Juliidae) (both Eocene series, 60 MYBP), and
Aplysiidae (Miocene series, 25 MYBP).

Comparing these data against the tree pro-
vides little information. While the Permian is
unique to the lower heterobranchs, the next
oldest genera (Acteon, Hydatina, Akera,
Bulla, Acteocina, Retusa; all Jurassic) are dis-
tributed throughout the tree. As previously
noted, the relatively short fossil record for
groups such as the sacoglossans or Aplysia
could be equally (perhaps more) a matter of
poor fossilization.

Diet: One of the original goals of this work
was to evaluate diet and its associated ana-
tomical modifications in relation to ceph-
alaspid phylogeny (Mikkelsen, 1993, 1994).
Diet was presented as one of the potential
sources of homoplasy, and digestive charac-
ters were coded with this in mind. Because
diet is likely the result of a complex interplay
of phylogeny, prey structure, and habitat, it
was not itself coded as a character (as done

422 MIKKELSEN

by Gosliner, 1978) and thus did not partici-
pate in tree construction. But it is important
now to map diet on the preferred tree, to ob-
serve how morphology could have contrib-
uted to feeding biology in the various clades.

From numerous literature sources (summa-
rized by Mikkelsen, 1990), diet has been
mapped on the preferred tree in Figure 47.
The most parsimonious explanation of the
pattern is that herbivory is most plesiomor-
phic in agreement with Purchon (1977; but
contrary to Haszprunar, 1985a). Herbivores,
that is, anaspids and bulloids, also possess
the most plesiomorphic radular forms among
the taxa in this analysis. Within the herbivores,
the sacoglossans specialize as suctorial feed-
ers with uniserial radulae, and the anaspids
and haminoeids handle filamentous algae
with complex gizzard plate configurations.

According to the preferred tree, carnivory
has evolved at least five times, but associated
with different suites of digestive system char-
acters. The philinoideans are the best-recog-
nized cephalaspid carnivores. Most of these
taxa have sickle-shaped laterals capable (at
least in some Philine; Hurst, 1965) of pulling
large, hard-shelled prey (clams, snails, fora-
miniferans) into the gut to be crushed by the
gizzard plates. Retusa, in the midst of this,
feeds (sometimes selectively) on similar prey
but instead of crushing, the gizzard plates
apparently hold an item in the gizzard while
digestive enzymes dissolve its tissues (Burn &
Bell, 1974). Bulla is an omnivore (Tchang Si,
1931; В. Winner, pers. comm.), the only one
on the tree; its radular and gizzard plate mor-
phologies are correspondingly unique among
shelled opisthobranchs.

Acteon and Hydatina are specialists on
polychaete worms [which contributed to Gos-
liner's (1978) clade], correlated with radulae
having numerous, minute, subequal teeth
(Rudman, 1972a; Yonow, 1992; Gosliner,
1994). Gegania valkyrie was found by Climo
(1975) on antipatharian corals, and was as-
sumed to be an ectoparasite. Ringicula spp.,
according to published sources, are general-
ists, feeding upon copepods and foraminifer-
ans (Pelseneer, 1924; Fretter, 1960; Bouchet,
1975); recognizable clams and foraminiferans
were seen in the gut of sectioned specimens
in this study. Fretter (1960) observed Ringi-
cula using its sickle-shaped radular teeth to
pull in large prey items, to be crushed not by
а gizzard (absent in Ringicula), but т the pos-
terior chamber of the stomach that has been
muscularized into “crushing plates.” The tree
supports parallel evolution of the bilobed-

rachidian/sickle-shaped-lateral type of radula
in Ringicula as well as the philinoideans. Ат-
gicula is able to process similar hard-shelled
prey, not in a gizzard, but analogously in a
modified stomach.

One additional character change, loss of
jaws (character 9), seems strongly associ-
ated with dietary modifications. Jaws are lost
five times on the tree: (a) in Acteon-Gegania,
(b) in Ringicula, (с) in the sacoglossans, (d) in
Retusa, and (e) in Scaphander-Philine. This is
affiliated with, respectively, (a) specialized
vermivory or ectoparasitism, (b) crushing
stomach plates, (c) suctorial feeding, (d) loss
of the entire buccal mass, and (e) obligate
carnivory with crushing gizzard plates.

It is thus evident that although diet might
not necessarily be the “driving force” behind
evolution of the shelled opisthobranchs [as
advocated by Thompson (1976) and Salvini-
Plawen (1988)], evolutionary novelties in di-
gestive morphology almost certainly have
“allowed” a wide variety of dietary niches to
be occupied.

CONCLUSIONS

1. The first phylogeny of shelled opistho-
branchs, based on morphology and employ-
ing parsimony-based cladistic methods, has
been generated. Non-homoplastic or at least
strong clade-supportive characters were de-
termined from external anatomy, mantle cav-
ity, digestive, nervous, and reproductive sys-
tems. This work has created a testable
hypothesis, now available for refinement and
comparison with future data from ultrastruc-
tural, biochemical, and/or molecular studies.

2. Monophyletic clades were confirmed for
Anaspidea and Sacoglossa, with the con-
firmed inclusion of Cylindrobulla in the latter
group.

3. Traditional Cephalaspidea has been split
into two groups. (a) Ringicula and Hydatina
were removed to the paraphyletic “archi-
tectibranchs,” and Acteon showed strong af-
finity for the “lower heterobranchs.” These
will require additional data for resolution of
monophyletic groups. Ringicula was the
most enigmatic member of this assemblage,
with suites of non-congruent characters in
need of confirmation and further study. (b)
Cephalaspidea s.s., as a sister group of
Anaspidea, was supported by four synapo-
morphies: flexed ciliated strips, three gizzard
plates, secondarily prepharyngeal nerve ring,
and genital ganglion on the visceral loop.

423

PHYLOGENETICS OF CEPHALASPIDEA

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424 MIKKELSEN

This clade includes two monophyletic sub-
clades: Bulloidea (Bulla, Haminoea, Sma-
гадате!а) supported by an exogyrous cili-
ated strip, and Philinoidea (Cylichna, Retusa,
Acteocina, Scaphander, Philine), based оп
digestive and nervous system configurations.

4. “Rampant parallelism” is upheld as a
evolutionary process within the shelled
opisthobranchs, with 25 of the 47 characters
showing some, and six characters signifi-
cant, levels of homoplasy on the preferred
cladogram.

5. Tree topology suggested a number of
evolutionary scenarios. (a) Formation of the
gizzard: the most plesiomorphic gizzard (in
Anaspidea) has numerous plates and gizzard
spines, consolidating into three plates in
Cephalaspidea s.s.; gizzard spines were re-
tained in Bulloidea, lost in Philinoidea. (b)
Sperm-conducting ducts: the caenogastro-
pod external ciliated groove is homologous
with the internal duct present in some shelled
opisthobranchs. A second (novel) external
groove, located laterally, developed in
shelled opisthobranchs probably initially for
egg transport. Both internal duct and external
groove co-occur in Sacoglossa; the internal
duct was lost in Anaspidea and Cephalaspi-
dea s.s., with the external groove assuming
the task of sperm transport. (c) Allosperm
storage sacks: most shelled opisthobranchs
have a proximal receptaculum seminis and
distal gametolytic gland, the latter probably
formed from the caenogastropod bursa cop-
ulatrix. Sacoglossans have no opistho-
branch-type gametolytic gland; their “bursa
copulatrix”* is probably a novel structure.
Some of the “lower heterobranchs” may
share the synapomorphy of a proximal “re-
ceptaculum apparatus,” consisting of a re-
ceptaculum seminis and gametolytic gland in
tandem arrangement. (d) Diet: herbivory is
presumed plesiomorphic, with carnivory
evolving independently at least five times, as-
sociated with different suites of digestive
system characters.

ACKNOWLEDGMENTS

This work was submitted in partial fulfill-
ment of requirements for a Ph.D. degree in
Biological Sciences at Florida Institute of
Technology (FIT), Melbourne, Florida. | grate-
fully acknowledge my committee chair, Dr.
Kerry B. Clark (FIT), for his advice and inter-
est during the entire course of my graduate
study. | also thank the remaining members of

my committee for their willing involvement:
Drs. Richard L. Turner (FIT), R. Grant Gilmore
(FIT and HBOI), Walter G. Nelson (FIT), and
Gary N. Wells (FIT). Two informal committee
members, Drs. Rudiger Bieler (FMNH) and
Terrence M. Gosliner (CAS), provided coun-
sel on phylogenetics, gastropod morphology,
and opisthobranch biology. Drs. Petra Sier-
wald (FMNH), John B. Wise (George Wash-
ington University, Washington, D.C.), Silvard
Р. Kool (Boston, Massachusetts), Ya.l. Star-
obogatov (Zoological Institute, Russian
Academy of Sciences, St. Petersburg, Rus-
sia), and C. Hedegaard (University of Califor-
nia, Berkeley) also contributed greatly
through discussion of various aspects of this
work. The manuscript also benefitted greatly
from the comments of two anonymous re-
viewers.

This work would not have been possible
without the assistance of colleagues who
provided specimens, and museum curators
or collection managers who allowed me ac-
cess and use of their cataloged material: Dr.
Kathe Jensen and Mr. Tom Schiotte (ZMUC),
Dr. Terrence M. Gosliner (CAS), Dr. Albert F.
Chadwick and Mr. Russell H. Jensen
(DMNH), the late Dr. T. Е. Thompson (Univer-
sity of Bristol, U.K.), Dr. Nathalie Yonow (Uni-
versity College of Swansea, U.K.), Dr. Anders
Warén (Naturhistoriska Riksmuseet, Stock-
holm, Sweden), Mr. Joseph W. Goy (Texas A
& M University), Mr. Leonard C. Hill (Miami,
Florida), Mrs. Jan M. Light (Godalming, Sur-
rey, U.K.), Dr. Kerry B. Clark (FIT), Dr. Duane
E. De Freese (formerly FIT; now Brevard
County EEL Program, Melbourne, Florida),
Dr. Robert W. Virnstein (formerly HBOI; now
St. Johns River Water Management District,
Palatka, Florida), Dr. Rüdiger Bieler (ЕММН),
Prof. Dr. Е. Gittenberger (МММ), Mr. H. L.
Strack (Rumphius Biohistorical Expedition,
Foundation for the Advancement of Biohis-
torical Research, Rotterdam, The Nether-
lands), Dr. Robert Ernest (Applied Biology,
Inc., Jensen Beach, Florida), and Drs. Alan R.
Kabat, José H. Leal, M. G. Harasewych, and
the late Richard S. Houbrick (USNM). A col-
lection of opisthobranchs prepared and em-
bedded by the late Dr. Hennig Lemche, gen-
erously made available by the Zoological
Museum, Copenhagen, provided both an in-
valuable resource and an historically interest-
ing vein to this research. Dr. Rudiger Bieler
also provided access to critical specimens of
Gegania valkyrie: (a) histological slides origi-
nally prepared by Dr. Gerhard Haszprunar

PHYLOGENETICS OF CEPHALASPIDEA 425

(Institut fúr Zoologie der Universitát Wien,
Austria) on loan from National Museum of
New Zealand (Wellington) and upon which
Haszprunar's (1985a) paper was based; and
(b) additional material on loan from MNHN for
joint histological sectioning and SEM. Access
to the scanning electron microscope and in-
vertebrate histology laboratory at ЕММН 1$
also acknowledged (the latter partially sup-
ported by National Science Foundation grant
DEB-9318231 to R. Bieler).

Literature and collecting assistance was
provided by Drs. John B. Wise and Rúdiger
Bieler. Advice and discussion on histological
procedures were provided by Dr. Rúdiger
Bieler, Mr. Woody Lee (SMSLP), Dr. Janice
Voltzow (University of Puerto Rico, Río Pie-
dras), and Mr. Oliver Putz (НВО! and Univer-
sitát Berlin, Germany). Translations of foreign
language publications were performed by Dr.
André Wybou (Vero Beach, Florida), Dr.
Cindy Lee Van Dover (formerly SMSLP), and
Mrs. Irene Guy (Melbourne, Florida).

| gratefully acknowledge financial support
and use of the laboratories, equipment, and
facilities of Harbor Branch Oceanographic In-
stitution, Ft. Pierce, Florida. Librarian Kristen
Metzger and Staff Photographer Tom
Smoyer assisted greatly throughout the
course of this research. Paul S. Mikkelsen
(formerly HBOI; now Palm Beach County En-
vironmental Resources Management, West
Palm Beach, Florida) assisted with collecting
and extracted some of the radulae. Addi-
tional financial assistance was provided by
the Hawaiian Malacological Society (1993
Scholarship), National Museum of Natural
History, Smithsonian Institution (1993 Rose-
water Fellowship), Field Museum of Natural
History (visiting research scholarship from
the Thomas J. Dee Fellowship Fund, 1993),
and Delaware Museum of Natural History.

This 15 Harbor Branch Contribution no.
1104.

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Revised Ms. accepted 30 Aug. 1995

APPENDIX |
SUMMARIZED ANATOMICAL
DESCRIPTIONS

The following are summarized descriptions
of the taxa involved in this study, based on
original dissections and histology, supple-
mented by literature data. References are
cited within the descriptions only for data un-
confirmed by original observations. These
are not complete anatomical descriptions;
rather, they are limited to those characters
used in the phylogenetic analysis. The reader
is referred to the literature examined for each
taxon for additional information, and to the
Characters and Coding section for more

432 MIKKELSEN

complete explanation of characters. Preced-
ing each description 1$ a list of зреситеп ma-
terial and literature examined in generating
the data matrix. Within material examined,
the following abbreviations explain the use of
specimens: D, dissection; H, histology; L, life
history study; M, mineralogy; S, SEM. The
exemplar species studied herein is marked
with an asterisk (*).

Acteon

*A. tornatilis (Linné, 1758) (type species)—
Rhossili, Gonor Peninsula, South Wales,
United Kingdom, no date, ex N. Yonow
[0,$]; МОС, Lemche material (embed-
ded), Frederikshavn, Denmark, 7/1948
[H]; Perrier & Fischer, 1911; Fretter,
1939; Fretter & Graham, 1954; Hurst,
1965; Thompson, 1976; Brace, 1977a.

A. candens Rehder, 1939—Ev. Marcus,
1974.

A. finlayi McGinty, 1955—Carriacou, no date,
ex J. Hamann [$].

A. pelecais Marcus, 1972—Ег. Marcus, 1958,
1972.

А. traskii Stearns, 1897 —Ev. Marcus, 1972.

Shell external. Foot simple. Operculum
present, paucispiral. Parapodia absent. Man-
tle cavity directed anteriorly. Plicatidium-type
gill two-sided, attached to mantle by suspen-
sory membranes. Ciliated strips beginning
bluntly at mantle edge, extending into long,
free pallial caecum, winding parallel to vis-
ceral mass within shell whorls. Jaws absent,
but cuticularized epithelium present adorned
with strong denticles. Radula with formula
100+.0.100+. Rachidian tooth absent. Lateral
teeth minute, numerous, identical in form,
denticulate. Distinct marginal teeth absent.
Tooth size uniform throughout ribbon.
Esophageal gizzard with gizzard plates ab-
sent. Stomach with pouch-like chamber. In-
testinal typhlosole short, not extending full
length from stomach to rectum. Nervous
system streptoneurous. Circumesophageal
nerve ring prepharyngeal. Cerebral/pleural
ganglia fused. Cerebral commissure long;
pedal commissure long. Visceral nerve loop
formula: L-A-B-V-P-A-L (Brace, 1977a, т
part). Genital ganglion on nerve emanating
from visceral ganglion (Brace, 1977a, in part);
genito-rectal nerve present. Eye with solid,
spherical lens, close to surface (S/C not
available), directed dorsolaterally. Reproduc-
tive system androdiaulic. Gonad with her-

maphroditic acini. Internal sperm duct
present; external ciliated groove absent.
Copulatory organ non-retractile, with ejacu-
latory duct continuous with internal sperm
duct; no penial gland or sperm storage area.
Prostate pallial, dense glandular tissue along
ciliated lumen. Single proximal allosperm
storage sack, comprised of a united recep-
tacular duct (with oriented sperm) and termi-
nal gametolytic gland. Egg mass ball-
shaped, anchored with a mucus string, with
single egg/capsule (Fretter & Graham, 1954).
Veliger PMO color not noted, but figured as
lightly pigmented (Thompson, 1976: fig. 51b).

Ringicula

*R. nitida Verrill, 1873—Bay of Baleeira, Por-
tugal, 5/1988, ex A. Waren [D,H];
Bouchet, 1975; Gosliner, 1981.

R. buccinea (Brocchi, 1814)—Fretter, 1960.

R. conformis Monterosato, 1875—Pelseneer,
1924; Gosliner, 1981; Thompson et al.,
1985.

В. sp.—HBOI sta. BE-295b, off Ft. Pierce In-
let, St. Lucie County, Florida, 126 m,
5/1979 [D—radula only]; Minichev, 1967.

Shell external. Foot simple. Operculum ab-
sent. Parapodia present. Mantle cavity di-
rected anteriorly. Plicatidium-type gill two-
sided, attached to mantle by suspensory
membranes. Ciliated strips beginning bluntly
at mantle margin; pallial caecum absent.
Jaws absent, but cuticularized, denticulate
epithelium present in oral cavity. Radula with
formula 1.0.1 (Fretter, 1960; Bouchet, 1975;
R. nitida, pers. obs.) or 1.1.1 (Thompson et
al., 1985; R. sp., pers. obs.). Rachidian tooth
(when present) rhomboid, with median inden-
tation (Thompson et al., 1985). Lateral teeth
single, with enlarged sickle-shaped cusps.
Marginals absent. Tooth size uniform
throughout ribbon. Esophageal gizzard ab-
sent. Stomach with pouch-like chamber. In-
testinal typhlosole short. Nervous system
streptoneurous (Pelseneer, 1924; Fretter,
1960). Circumesophageal nerve ring post-
pharyngeal. Cerebral/pleural ganglia fused in
R. nitida, separate but closely apposed (equi-
distant) in R. buccinea and R. conformis (fide
Pelseneer, 1924; Fretter, 1960). Cerebral
commissure long; pedal commissure long.
Visceral nerve loop formula: LAB- -V-P-A-L
(“the subesophageal ganglion is the only one

PHYLOGENETICS OF CEPHALASPIDEA 433

on the visceral loop which has migrated for-
wards”; Fretter, 1960: 545; histological
cross-sections too poor to trace). Genital
ganglion/nerve? Eye with solid, spherical
lens, embedded midway between surface
and nerve ring (?; S/C not available), directed
dorsolaterally; sometimes absent (Pelseneer,
1924). Reproductive system androdiaulic
(Pelseneer, 1924) or monaulic (Gosliner,
1981; pers. obs.). Gonad with separate male
and female acini. Internal sperm duct absent
and external ciliated groove present (Fretter,
1960; Gosliner, 1981; pers. obs.), or vice
versa (Pelseneer, 1924). Copulatory organ re-
tractile, with tubular glandular tissue (“pros-
tate” of authors), without confirmed sperm
storage area; penis simple, presence/ab-
sence of ejaculatory duct unconfirmed. Prox-
imal receptaculum seminis; distal “Бигза
copulatrix”” (Fretter, 1960; Gosliner, 1978,
1981; sperm sacks could not be confirmed in
histological sections). Egg mass and larval
development unknown.

Remarks. The preserved specimens avail-
able for this study did not section well, per-
haps due to poor fixation; for this reason,
many of the anatomical features reported in
the literature could not be confirmed. The
type species of Ringicula is Auricula ringens
Lamarck, 1804 (by subsequent designation
of Gray, 1847); it was not used in this study
because it is an Eocene fossil from the Paris
Basin, and thus could not provide morpho-
logical characters from soft anatomy.

Hydatina

*H. physis (Linné, 1758) [= H. vesicaria Light-
foot, 1786] (type species) —Hobe Sound,
Tequesta, Florida, 4/1977 [L]; northern
Colombia, 4/1983, ex L. Hill [D,S];
НВОМ 065:01938, РММ-872, Tequesta,
Florida, May 1983 [D,L,S]; МММ, Ambon,
Indonesia, n.d. [D,H]; Rudman, 1972a;
Winner, 1984; Zehra 8 Perveen, 1992.

H. velum (Gmelin, 1791)—Eales, 1938; Nat-
arajan, 1970.

Shell external. Foot simple. Operculum ab-
sent. Parapodia present. Mantle cavity di-
rected laterally. Plicatidium-type gill two-
sided, attached to mantle by suspensory
membranes. Ciliated strips beginning bluntly
at mantle edge, extending into long, free pal-
lial caecum, winding parallel to visceral mass
within shell whorl. Jaws present, comprised

of flattened elements with strong denticles.
Radula with formula (10-20).(0-1).(10-20).
Rachidian tooth absent (pers. obs.) or vesti-
gial with median denticle (Rudman, 1972a) or
flat, plate-like (Eales, 1938) when present.
Laterals subequal, blade-like, hooked, with
varying numbers of denticles (0-5), decreas-
ing in number outwardly in a single radular
row. Distinct marginal teeth absent. Tooth
size uniform throughout ribbon. Oral tube ex-
tremely long; evertible. Esophageal gizzard
with gizzard plates absent. Stomach with
pouch-like chamber. Intestinal typhlosole ab-
sent. Nervous system slightly streptoneur-
ous. Circumesophageal nerve ring prepha-
ryngeal. Cerebral/pleural ganglia fused.
Cerebral commissure short; pedal commis-
sure extremely long. Visceral nerve loop for-
mula: LA- -B-V- - -PAL (Rudman, 1972a) or
LA- - -BV- - -PAL (pers. obs.; Eales, 1938).
Genital ganglion on nerve emanating from
visceral ganglion; genito-rectal nerve absent.
Eye with solid, spherical lens, close to sur-
face (S/C = 0.16), directed dorsolaterally. Re-
productive system androdiaulic. Gonad with
hermaphroditic acini. Internal sperm duct
present; external ciliated groove absent.
Copulatory organ nonretractile, with ejacula-
tory duct continuous with internal sperm
duct; no penial gland or sperm storage area.
[Ghiselin's (1966: 346) statement that the
prostate of Hydatina ‘has been displaced to
the base of the penis” reflects the posterior
displacement of the copulatory organ (to
near the common genital opening) more than
an anterior shift of the prostate; this is not a
penial gland in the sense found in Haminoea
or Bulla.] Proximal ‘‘exogenous sperm sack’;
distal gametolytic gland (Rudman, 1972a).
Prostate pallial, of dense glandular tissue
along ciliated lumen (Rudman, 1972a). Egg
mass ribbon-shaped, anchored with a mucus
string, with 20-30 eggs/capsule; Zehra 8
Perveen (1992) reported 4-6/capsule. Veliger
PMO black. Chromosome number 15 (Nat-
arajan, 1970).

Scaphander

*S. lignarius (Linné, 1758) (type species)—
Guiart, 1901; Perrier 8 Fischer, 1911;
Fretter, 1939; Lloyd, 1952; Hurst, 1965;
Brace, 1977a; Vitturi et al., 1985.

$. clavus Dall, 1889—Ev. Marcus & Er. Mar-
cus, 1967a.

S. punctostriatus (Mighels, 1841)—Gosliner,
1978.

434 MIKKELSEN

S. watsoni Dall, 1881—off Brazil, 5/1987, ex
J. Leal [Н]; Gulf of Mexico, 6/1989, ex J.
Goy [D,S,M].

Shell external. Foot supplemented by pos-
terior pallial lobe. Operculum absent. Para-
podia present. Mantle cavity directed later-
ally. Plicatidium-type gill two-sided, attached
to mantle by suspensory membranes. Ciliated
strips flexed posteriorly at mantle margin, ex-
tending into long, free pallial caecum. Jaws
absent, but oral cavity lined with smooth cu-
ticle. Radula with formula 1.(0-1).1. Rachid-
ian tooth rhomboid, without median indenta-
tion, caducous. Lateral teeth single, with
enlarged sickle-shaped cusps. Marginals ab-
sent. Tooth size uniform throughout ribbon.
Esophageal gizzard with three calcified giz-
zard plates. Gizzard spines absent. Stomach
of flow-through type, without pouch-like
chamber. Intestinal typhlosole short. Nervous
system slightly streptoneurous. Circume-
sophageal nerve ring prepharyngeal. Cere-
bral/pleural ganglia separate, equidistant.
Cerebral commissure long; pedal commis-
sure long. Visceral nerve loop formula: L- -A-
BV-P- -AL [left pallial considered fused with
pleural by Brace (1977a) and by Schmekel
(1985), but here treated as present through
position of left pallial nerve and existence of
vestigial somata (pers. obs.; Brace, 1977a)].
Genital ganglion on visceral loop between vis-
ceral and supraesophageal ganglia. Eye with
solid, spherical lens (slightly irregularly
shaped, but bicolored), deeply embedded
(S/C = 1.7), directed ventrolaterally. Repro-
ductive system monaulic. Gonad with her-
maphroditic acini. Internal sperm duct absent;
external ciliated groove present. Copulatory
organ retractile, with terminal spherical glan-
dular mass (““prostate” of authors) containing
a large internal lumen and ducts to glandular
tissue; without sperm storage area. Penis
largely invaginated when retracted, contain-
ing an ejaculatory duct; tip simple. Proximal
receptaculum seminis; distal gametolytic
gland. Egg mass and larval development un-
known. Chromosome number 18 (Vitturi et al.,
1985).

Philine

*P. aperta (Linné, 1767) (type species) —Ox-
witch, United Kingdom, 3/1984, ex N.
Yonow [D,M]; ZMUC, Lemche material
(embedded), Gullmarfjord, Bohuslán,
Sweden, 7/1947 [H]; ZMUC, Lemche

material (embedded), no data, 5/1948
[H]; Tchang Si, 1931; Brown, 1934; Fret-
ter, 1939; Thorson, 1946; Lloyd, 1952;
Hurst, 1965; Vitturi et al., 1985.

P. angasi (Crosse & Fischer, 1865)—Rud-
man, 1972b.

P. denticulata (J. Adams, 1800)—Horikoshi,
1967.

р. falklandica
1972b.

P. gibba Strebel, 1908—Odhner, 1926; Rud-
man, 1972b; Seager, 1978, 1979.

P. japonica Lischke, 1872—Inaba, 1959.

P. orientalis A. Adams, 1854—USNM
858415, Hong Kong, 5/1987 [D,H,S].

P. quadripartita Ascanius, 1772—Patterson,
1969.

P. spp. [unidentified—Williams, 1975.

Powell, 1954—Rudman,

Shell internal within posterior half of body.
Foot supplemented by posterior pallial lobe.
Operculum absent. Parapodia present. Man-
tle cavity directed laterally. Plicatidium-type
gill two-sided, attached to mantle by suspen-
sory membranes. Ciliated strips beginning
bluntly at mantle margin, extending into short
pallial caecum. Jaws absent, but oral cavity
cuticularized. Radula with formula (1-6).1.(0-
1).1.(1-6). Rachidian tooth absent т P.
aperta, present as nondenticulate plate in P.
falklandica and P. gibba (fide Rudman,
1972b; Seager, 1978). Lateral teeth single,
with enlarged sickle-shaped cusps. Margin-
als present in P. gibba (fide Rudman, 1972b;
Seager, 1978) and several other species,
hook-like, smaller, and less robust than lat-
erals. Tooth size uniform throughout ribbon.
Esophageal gizzard with three gizzard plates
present. Gizzard plates fully calcified in P.
aperta, not calcified in P. falklandica and P.
gibba (Rudman, 1972b; Seager, 1978; but
not verified here). Gizzard spines absent.
Stomach of flow-through type, without
pouch-like chamber. Intestinal typhlosole
short. Nervous system slightly streptoneur-
ous; barely so in P. aperta and P. orientalis,
wherein right connective is only slightly more
dorsal than left; Williams (1975) showed dis-
tinct crossing at the far posterior end of the
visceral loop in two unidentified species. Cir-
cumesophageal nerve ring prepharyngeal.
Cerebral/pleural ganglia separate, equidis-
tant. Cerebral commissure long; pedal com-
missure long. Visceral nerve loop formula:
L- -A-BV-P- -AL (in P. falklandica and P. an-
gasi, fide Rudman, 1972b; in P. gibba, fide
Seager, 1978), or L--(A)-BV- - -РАЁ (in P.

PHYLOGENETICS OF CEPHALASPIDEA 435

арепа and P. orientalis, pers. obs.; left pallial
missing, location of left pallial nerve indi-
cated). Because of the generally plesiomor-
phic nature of P. falklandica (fide Rudman,
1972b), the nervous system is coded as in
this species. Genital nerve emanating from
visceral ganglion (in P. falklandica and others;
Rudman, 1972b; Gosliner, 1978) or genital
ganglion on visceral loop between visceral
and supraesophageal ganglia (in P. aperta, P.
angasi, and others; Rudman, 1972b; Brace,
1977a; pers. obs.). Eye with solid, irregular
lens, with (?) “anchors,” deeply embedded
(S/C = 0.6), directed ventrolaterally. Repro-
ductive system monaulic. Gonad with her-
maphroditic acini. Internal sperm duct ab-
sent; external ciliated groove present.
Copulatory organ retractile, in P. aperta and
P. orientalis with long single-stranded tube
(“prostate” of authors) emanating from rear
of penial sheath, comprised of three types of
histological morphology: strongly staining
glandular tissue (most distal), weakly staining
glandular tissue (intermediate), and two-part
conducting tube for autosperm transport
connecting to internal duct at base of penis.
External ciliated groove continuing into penial
sheath, then closing to form an autosperm
intake duct (“уаз deferens” of Lloyd, 1952)
that exits to lie within the hemocoel alongside
the penial sheath, storing oriented sperm in
tube and small vesicle (“зретт vesicle” of
Lloyd, 1952), connecting to elongated tube at
junction of glandular tissue and two-part
conducting tube. Two-part conducting tube
comprised of two lunate, internally ciliated
halves, each half with central “typhlosole,”
and receiving contents of glandular tube on
one side, of autosperm intake duct on other
side. Penis with ejaculatory duct continuous
with coelomic two-part conducting tube; tip
elaborated. Proximal receptaculum seminis;
distal gametolytic gland. Egg mass ball-
shaped, anchored with a mucus string, with
single eggs/capsule [Tchang Si, 1931 (al-
though 1-3 eggs/capsule noted in enlarged
capsules at the end of the egg string; Lloyd,
1952)]. Veliger PMO red to jet black (Thorson,
1946; Horikoshi, 1967). Chromosome num-
ber 17 (Inaba, 1959; Patterson, 1969) or 18
(Vitturi et al., 1985).

Cylichna

*C. cylindracea (Pennant, 1777) (type spe-
cies) —ZMUC, Lemche material (embed-
ded), Gullmarfjord, Bohuslán, Sweden,

8/1949 [H]; Sweden, ex A. Waren [M,S];
Lemche, 1956; Hurst, 1965.

С. verrillii Dall, 1889—HBOM 065:02338,
Johnson-Sea-Link | dive #920, off Ft.
Pierce, Florida, 10/1980 [$].

C. alba (Brown, 1827)—Lemche, 1956.

С. magna Lemche, 1941—Lemche, 1956.

Shell external. Foot supplemented by pos-
terior pallial lobe. Operculum absent. Parap-
odia present. Mantle cavity directed laterally.
Plicatidium-type gill two-sided, attached to
mantle by suspensory membranes. Ciliated
strips flexed at mantle margin, extending into
short pallial caecum (Lemche, 1956: figs. 27,
41). Jaws present, comprised of rod-shaped
elements, strongly bent, and bearing long
denticles. Radula with formula (2-9).1.1.1.(2-
9). Rachidian tooth rhomboid with median in-
dentation. Lateral teeth single, with enlarged
sickle-shaped cusps. Marginals hook-
shaped, smaller, and less robust than later-
als. Tooth size uniform throughout ribbon.
Esophageal gizzard with three uncalcified
gizzard plates present. Gizzard spines ab-
sent. Stomach of flow-through type, without
pouch-like chamber. Intestinal typhlosole
short (Lemche, 1956: fig. 168). Nervous sys-
tem slightly streptoneurous (Lemche, 1956;
Hurst, 1965). Circumesophageal nerve ring
prepharyngeal. Cerebral/pleural ganglia sep-
arate, equidistant (Lemche, 1956: figs. 55-
57). Cerebral commissure long; pedal com-
missure long. Visceral nerve loop formula:
L-(A)- -BV-P- -AL [no left pallial detected by
Lemche (1956), but here treated as present
through the position of the left pallial nerve
(Lemche, 1956: fig. 56)]. Genital ganglion on
visceral loop between visceral and suprae-
sophageal ganglia (Lemche, 1956). Eyes ap-
parently absent in C. alba and C. magna (fide
Lemche, 1956); very small eye present in C.
cylindracea, without lens, deeply embedded
(S/C = 1.1), directed ventrolaterally. Repro-
ductive system monaulic. Gonad with sepa-
rate male and female acini (Lemche, 1956).
Internal sperm duct absent; external ciliated
groove present. Copulatory organ retractile,
with terminal glandular bulb ('“prostate” of
authors) emanating from rear of penial
sheath. Penis largely invaginated when re-
tracted (termed “‘rudiment’’ and only occa-
sionally present by Lemche, 1956), without
ejaculatory duct; tip simple. Terminal glandu-
lar bulb with tall cells lining a rather extensive
lumen, opening into penial sheath at base of
penis; without confirmed sperm storage area.

436 MIKKELSEN

Proximal receptaculum seminis; distal game-
tolytic gland. Egg mass and larval develop-
ment unknown.

Acteocina

*A. canaliculata (Say, 1826) (type species)—
PMM-814, Biscayne Bay, Dade County,
Florida, 2/1982 [L]; PMM-826, Haulover
Canal, Brevard County, Florida, 4/1982
[0]; PMM-841, Ft. Pierce, St. Lucie
County, Florida, 5/1982 [L]; PMM-864,
Haulover Canal, Brevard County, Flor-
ida, 12/1982 [D,H,M]; PMM, Indian River,
Florida, 1983 [$]; PMM-900, Peanut Is-
land, Palm Beach County, Florida,
9/1985 [0]; Franz, 1971; Ev. Marcus,
1977b; Gosliner, 1979; Mikkelsen 8 Mik-
kelsen, 1984.

A. atrata Mikkelsen & Mikkelsen, 1984—
PMM-817, Ft. Pierce, Florida, 2/1982
[L,M]; PMM-818, Haulover Canal,
Brevard County, Florida, 2/1982 [LU];
PMM-826, Haulover Canal, Brevard
County, Florida, 4/1982 [L]; Mikkelsen 8
Mikkelsen, 1984.

A. bidentata (Orbigny, 1841)—Er.
1958.

А. candei (Orbigny, 1841)—Er. Marcus, 1958.

Marcus,

Shell external. Foot supplemented by pos-
terior pallial lobe. Operculum absent. Parap-
odia present, small. Mantle cavity directed
laterally. Ciliated strips beginning bluntly at
mantle margin, extending into short pallial
caecum. Plicatidium-type gill two-sided, at-
tached to mantle by suspensory membranes.
Jaws present, comprised of blunt, rod-
shaped elements. Radula with formula 1.1.1.
Rachidian tooth rhomboid with median in-
dentation; rachidian absent in some species
(Gosliner, 1994). Lateral teeth single, with
enlarged sickle-shaped cusps. Marginals
absent. Tooth size uniform throughout rib-
bon. Esophageal gizzard with three partially
calcified gizzard plates present. Gizzard
spines absent. Stomach of flow-through
type, without pouch-like chamber. Intestinal
typhlosole? Nervous system slightly strep-
toneurous. Circumesophageal nerve ring
prepharyngeal. Cerebral/pleural ganglia sep-
arate, equidistant. Cerebral commissure
long; pedal commissure long. Visceral nerve
loop formula: L-(A)- -BV-P--AL (left pallial
ganglion absent, but here treated as present
through the position of the left pallial nerve).
Genital ganglion on visceral loop between

visceral and supraesophageal ganglia. Eye
with solid, spherical lens, embedded midway
between surface and nerve ring (?; S/C not
available), directed ventrolaterally. Repro-
ductive system monaulic. Gonad acini her-
maphroditic. Internal sperm duct absent; ex-
ternal ciliated groove present. Copulatory
organ retractile, with glandular tissue (“ргоз-
tate” of authors) with ciliated lumen; sperm
storage not confirmed; penis with external
sperm duct only, without ejaculatory duct.
Proximal receptaculum seminis (Gosliner,
1979); distal gametolytic gland. Egg mass
ball-shaped, anchored with a mucus string,
with single eggs/capsule. Veliger PMO color-
less.

Bulla

*B. striata Bruguière, 1792—PMM-931,
Spanish Harbor Keys, Florida Keys,
1/1988 [D,H,M]; НВОМ 065:02023,
Hobe Sound, Tequesta, Florida, 3/1979
[H,S]; Biscayne Bay, Florida, 1/1988
[D,S]; PMM-996, Indian River, Bessie
Cove, Martin County, Florida, 5/1994
[D,L]; Tchang Si, 1931; Er. Marcus, 1957;
Brace, 1977a; Vitturi et al., 1985; Winner,
1985.

В. ampulla Linné, 1758 (type species)—NNM,
Ambon, Indonesia, n.d. [D]; Eales, 1938;
Gosliner, 1978.

B. gouldiana Pilsbry, 1895—Robles, 1975;
Williams, 1975.

B. solida Gmelin, 1791—HBOM 065:00281,
R/V GOSNOLD 237/514, off Sebastian,
Florida, 25 т, 12/1974 [0].

Shell external. Foot supplemented by pos-
terior pallial lobe. Operculum absent. Para-
podia present. Mantle cavity directed later-
ally. Plicatidium-type gill two-sided, attached
to mantle by suspensory membranes. Cili-
ated strips exogyrous; pallial caecum short.
Jaws present, comprised of blunt, rod-
shaped elements. Radula with formula
1.2.1.2.1. Rachidian tooth rhomboid, laterally
elongated, denticulate, with central cusp
smaller than 5-7 subequal cusps on either
side. Lateral teeth ““claw-shaped,” with 4-10
robust cusps; the first with main cusp central;
the second with main cusp at inner edge.
Marginals plate-like, non-denticulate. Tooth
size uniform throughout ribbon. Esophageal
gizzard with three uncalcified gizzard plates
present. Gizzard spines present preceding
and following plates, flattened. Stomach of

PHYLOGENETICS OF CEPHALASPIDEA 437

flow-through type, without pouch-like cham-
ber. Intestinal typhlosole absent. Nervous
system slightly streptoneurous. Circume-
sophageal nerve ring prepharyngeal. Cere-
bral/pleural ganglia separate, equidistant.
Cerebral commissure long; pedal commis-
sure long. Visceral nerve loop formula: L- -
A-BV-P- -AL. Genital ganglion on or near
visceral loop between visceral and suprae-
sophageal ganglia. Eye with solid, spherical
lens, close to surface (S/C = 0.16), directed
dorsolaterally. Reproductive system monau-
lic. Gonad with hermaphroditic acini. Internal
sperm duct absent; external ciliated groove
present. Copulatory organ retractile, with
glandular ciliated coils within spherical vesi-
cle (“prostate” of authors) proximal to a ter-
minal glandular appendix (= sperm storage
bulb?). Penis with ejaculatory duct; tip sim-
ple. Receptaculum seminis proximal, embed-
ded within female gland mass; gametolytic
gland distal with additional sperm-storage
area in expanded base of gametolytic duct.
Egg mass of tangled strings, unanchored,
with single eggs/capsule (rarely 2/capsule;
1-25/capsule reported by Robles (1975)].
Veliger PMO dark red. Chromosome number
17 (Vitturi et al., 1985).

Haminoea

*H. antillarum (Orbigny, 1841) —PMM-933, Ft.
Pierce Inlet, Florida, 4/1988 [D,H,S,M];
Biscayne Bay, Florida, 1/1988 [D,L];
Bandel, 1976.

H. hydatis (Linné, 1758) (type species) —Per-
rier & Fischer, 1914; Fretter, 1939; Berrill,
1931; Tchang Si, 1931; Vitturi et al.,
1985.

H. elegans (Gray, 1825) —PMM-874, Semi-
nole Shoals, Martin County, Florida,
8/1983 [0]; PMM-931, Spanish Harbor
Keys, Florida Keys, 2/1988 [D]; Er. Mar-

CUS, 1958,

H. musetta Marcus 8 Burch, 1965—Burch 4
Natarajan, 1967.

H. solitaria (Say, 1821)—Smallwood, 1903;
Inaba, 1959; Harrigan & Alkon, 1978.

H. succinea (Conrad, 1846) —PMM-912, Na-
ples, Florida, 3/1987 [D,L].

H. virescens (Sowerby, 1833)—Hurst, 1967;
Williams, 1975.

Н. zelandiae Gray, 1843—Rudman, 197 1a.

Shell external. Foot supplemented by pos-
terior pallial lobe. Operculum absent. Para-
podia present. Mantle cavity directed later-

ally. Plicatidium-type gill two-sided, attached
to mantle by suspensory membranes. Cili-
ated strips exogyrous; pallial caecum short.
Jaws present, comprised of blunt, rod-
shaped elements. Radula with formula (5-
63).1.1.1.(5-63). Rachidian tooth quadrate,
tricuspidate with median denticle largest.
Lateral teeth broad, denticulate. Marginals
hook-shaped, decreasing in size outwardly.
Tooth size uniform throughout ribbon.
Esophageal gizzard with three uncalcified,
transversely ridged gizzard plates. Gizzard
spines present preceding and following
plates. Stomach of flow-through type, with-
out pouch-like chamber. Intestinal typhlosole
short. Nervous system slightly streptoneur-
ous. Circumesophageal nerve ring postpha-
ryngeal. Cerebral/pleural ganglia separate,
equidistant. Cerebral commissure long;
pedal commissure long. Visceral nerve loop
formula: L-[?A]-BV-P--AL; position of left
pallial ganglion variable. Genital ganglion on
visceral loop between visceral and suprae-
sophageal ganglia. Eye with solid, spherical
lens (bicolored, but with shallow surface hol-
lows), close to surface (S/C = 0.08, 0.16), di-
rected dorsolaterally. Reproductive system
monaulic. Gonad with hermaphroditic acini.
Internal sperm duct absent; external ciliated
groove present. Copulatory organ retractile,
with terminal two-part spherical glandular
mass (“‘prostate’’ of authors), the most prox-
imal with central channel and ducts to glan-
dular tissue, the most distal with large inter-
nal lumen for storage of unoriented sperm.
Penis with external groove only, without ejac-
ulatory duct. Spermatophores reported (Per-
rier & Fischer, 1914). Proximal receptaculum
seminis; distal gametolytic gland. Egg mass
shape variable, anchored or not, with single
eggs/capsule [in H. antillarum and others; H.
hydatis with 1/capsule (Tchang Si, 1931) or
2/capsule (Berrill, 1931)]. Veliger PMO red to
black. Chromosome number 16 (Inaba,
1959), 17 (Burch & Natarajan, 1967) or 18
(Vitturi et al., 1985).

Smaragdinella

*S. calyculata Broderip & Sowerby, 1829 [= $.
viridis Rang, in Quoy & Gaimard, 1833]
(type species)—USNM 543627, Oahu,
Hawaii, 1937 [Н]; USNM 751607, Easter
Island [D,M,S]; Rumphius Biohistorical
Expedition, sta. 40, Ambon, Indonesia,
12/1990 [D]; Ev. Marcus & Burch, 1965;
Burch & Natarajan, 1967; Natarajan,
1970; Rudman, 1972c; Williams, 1975.

438 MIKKELSEN

Shell external. Foot supplemented by
posterior pallial lobe. Operculum absent.
Parapodia present. Mantle cavity directed
laterally. Plicatidium-type gill two-sided, at-
tached to mantle by suspensory membranes.
Ciliated strips exogyrous; pallial caecum
short. Jaws present, comprised of blunt, rod-
shaped elements. Radula present, with for-
mula (5-6).(13-45).1.(13-45).(5-6). Rachidian
tooth rhomboid, with median denticle only.
Lateral teeth hook-shaped, largest in center
half-row, decreasing in size in- and out-
wardly. Marginals plate-like, lacking hooks.
Tooth size uniform throughout ribbon.
Esophageal gizzard with three uncalcified,
transversely ridged gizzard plates. Gizzard
spines present, V-shaped. Stomach of flow-
through type, without pouch-like chamber
(Rudman, 1972c). Intestinal typhlosole?
Nervous system euthyneurous. Circume-
sophageal nerve ring postpharyngeal. Cere-
bral/pleural ganglia separate, equidistant.
Cerebral commissure long; pedal commis-
sure long. Visceral nerve loop formula: LA- -
BV- -PAL. Genital ganglion immediately ad-
jacent to visceral ganglion, on loop. Eye with
solid, spherical lens, close to surface (S/C
not available), directed dorsolaterally. Repro-
ductive system monaulic. Gonad with her-
maphroditic acini. Internal sperm duct ab-
sent; external ciliated groove present.
Copulatory organ retractile, with terminal
three-part spherical glandular mass (“ргоз-
tate” of authors), the most proximal two with
central channel (and ducts to glandular tis-
sue?), the most distal with multiple compart-
ments for sperm storage. Penis with external
groove, without ejaculatory duct; tip simple.
Proximal receptaculum seminis; distal game-
tolytic gland (Rudman, 1972c). Egg mass and
larval development unknown. Chromosome
number 18 (Burch & Natarajan, 1967; Natara-
jan, 1970).

Retusa

*R. obtusa (Montagu, 1803) (type species)—
Fleet, Dorset, United Kingdom, 2/1986,
ex T.E. Thompson [H,S,M]; Thorson,
1946; Hurst, 1965; Smith, 1967; Gos-
liner, 1978; Berry, 1989.

В. chrysoma Burn, in Burn
1974—Burn 4 Bell, 1974.

В. operculata Minichev, 1966—Minichev,
1966.

R. semisulcata Philippi, 1836—Huber, 1993.

В. truncatula (Bruguiere, 1792)—ZMUC,
Lemche material (embedded), Isefjord,

8 Bell,

Denmark, 4/1944 [H]; ZMUC, Lemche
material (embedded), no data, 5/1948
[H]; Sao Miguel, Azores, 7/1991 [D; Mik-
kelsen, 1995]; CAS 072614, Ponta de Pi-
rámide, Sáo Miguel, Azores, 7/1988, [0];
Vayssiere, 1893; Rasmussen, 1944;
Thorson, 1946; Burn & Bell, 1974; Lu-
que, 1983.

В. semisulcata Philippi, 1836—Huber, 1993.

Shell external. Foot supplemented by pos-
terior pallial lobe. Operculum absent in adult
R. obtusa and most other species; present in
three species—R. chrysoma (fide Burn & Bell,
1974), R. operculata (fide Minichev, 1966),
and R. truncatula (fide Burn & Bell, 1974; Lu-
que, 1983; but not in some populations, Mik-
kelsen, 1995). Parapodia absent. Mantle cav-
ity directed laterally. Ciliated strips flexed
posteriorly at mantle margin. Pallial caecum
short (Gosliner, 1978). Gill absent. Vayssiere
(1893) ambiguously described a gill in R.
truncatula, but he could in fact have been
describing some other pallial structure such
as the extensive kidney. Smith (1967) did not
mention a gill in his excellent description of
the larval development of R. obtusa. Jaws,
radula, and buccal mass absent. Oral tube
lined by ciliated cells only, without any trace
of cuticularization. Esophageal gizzard with
three uncalcified tuberculate gizzard plates
present. Gizzard spines absent. Stomach of
flow-through type, without pouch-like cham-
ber. Intestinal typhlosole short. Nervous sys-
tem slightly streptoneurous (Hurst, 1965; Hu-
ber, 1993). Circumesophageal nerve ring
“prepharyngeal” according to relative posi-
tion of buccal ganglia. Cerebral/pleural gan-
glia fused in R. obtusa, but separate (and
equidistant) in other species [e.g., В. trunca-
tula (pers. obs.); R. semisulcata (fide Huber,
1993)]. Cerebral commissure long; pedal
commissure long. Visceral nerve loop for-
mula variable in the genus: L- - -BV-P- -AL (R.
obtusa, R. truncatula, left pallial not seen;
pers. obs.; Vayssière, 1893); LA- - -BV- - -PAL
(R. semisulcata; fide Huber, 1993). [See Hu-
ber (1993) for summary of other variability in
visceral loop ganglia.] Genital ganglion on
visceral loop between visceral and suprae-
sophageal ganglia. Eye absent in R. obtusa;
in R. truncatula, with solid, spherical lens,
deeply embedded (S/C = 0.86), directed ven-
trolaterally. Reproductive system monaulic.
Gonad with hermaphroditic acini. Internal
sperm duct absent; external ciliated groove
present. Copulatory organ retractile, with

PHYLOGENETICS OF CEPHALASPIDEA 439

glandular tissue (‘‘prostate’’ of authors);
sperm storage area (Ghiselin, 1966; Gosliner,
1978) unconfirmed. Penis with external
groove only, without ejaculatory duct. Proxi-
mal receptaculum seminis; distal gametolytic
gland. Egg mass ball-shaped, unanchored,
with single eggs/capsule (Smith, 1967; Berry,
1989). Veliger PMO black (Smith, 1967) or
colorless (Rasmussen, 1944: fig. 13).

Cylindrobulla

*С. beauii (P. Fischer, 1856) (type species)—
Key Largo, Florida Keys, 7/1987, ex
АЛ. Virnstein [H]; Belize, ex RSS.
Houbrick [H]; HBOM 065:01745, Shark
Channel, Florida Keys, 9/1981 [S]; Er.
Marcus & Ev. Marcus, 1970 [in part].

С. п. sp.—Bermuda, 8/1990, ex О. De Freese
[D,H,L,S]; HBOM 065:01893, Key Largo,
Florida Keys, 7/1982 [S].

Shell external. Foot simple. Operculum ab-
sent. Parapodia absent. Mantle cavity di-
rected laterally. Plicatidium-type gill one-
sided, attached to mantle over its full length.
Ciliated strips beginning bluntly at mantle
edge; no pallial caecum. Single anterior
adductor muscle present. Jaws absent, but
thickened cuticular epithelium in oral tube.
Аааа uniseriate, with formula 0.1.0.
Rachidian tooth quadrate-cuspidate, with
denticles flanking main cusp. Tooth size uni-
form throughout ribbon. Descending limb/as-
cus present. Pharyngeal pouches present.
Esophageal gizzard with gizzard plates ab-
sent. Stomach of flow-through type, without
pouch-like chamber. Intestinal typhlosole en-
tire length from stomach to rectum. Nervous
system slightly streptoneurous. Circume-
sophageal nerve ring postpharyngeal. Cere-
bral/pleural ganglia fused. Cerebral commis-
sure long; pedal commissure short. Visceral
nerve loop formula: L-A-B-V-PA- -L. Genital
nerve emanating from visceral ganglion; no
ganglion seen in cross-sections. Eye with
solid, oblong, irregular lens, with “anchors,”
embedded deeply (S/C = 1.3, 1.1), directed
ventrolaterally. Reproductive system an-
drodiaulic [not oódiaulic as according to
Ghiselin (1966), who based his conclusion on
incorrect data presented for Ascobulla ulla by
Ev. Marcus 4 Er. Marcus (1956), corrected by
Ev. Marcus (1972)]. Gonad acini? Both inter-
nal sperm duct and external ciliated groove
present. Copulatory organ retractile, without
glandular tissue or sperm storage area. Penis

with ejaculatory duct continuous with internal
sperm duct; tip simple. Prostate pallial, of
“flower” configuration. Two distal sperm
sacks: larger bursa copulatrix (containing un-
oriented sperm) and smaller receptaculum
seminis (containing oriented sperm); game-
tolytic gland absent. Egg mass of tangled
strings, unanchored, with single egg/capsule
(D. De Freese, pers. comm.). Veliger PMO or-
ange.

Ascobulla

*A. ulla (Marcus 8 Marcus, 1970) (type spe-
cies)—Ft. Pierce Inlet, Florida, 8/1990,
ex D. De Freese [H]; PMM-991, Sebas-
tian Inlet, Florida, 7/1993 [H]; PMM-992,
Sebastian Inlet, Florida, 8/1993 [0];
Marco, Florida, no date, CAS 77802 [0];
D. De Freese (L; pers. comm.); Ev. Mar-
cus 8 Er. Marcus, 1956; Ev. Marcus,
1972.

Shell external. Foot simple. Operculum ab-
sent. Parapodia absent. Mantle cavity di-
rected laterally. Plicatidium-type gill one-
sided, attached to mantle over its full length.
Ciliated strips beginning bluntly at mantle
edge; no pallial caecum. Single anterior ad-
ductor muscle present. Jaws absent, but
thickened cuticular epithelium in oral tube.
Radula uniseriate, with formula 0.1.0.
Rachidian tooth dagger-shaped, with mar-
ginal serrations/denticulations. Tooth size in-
creasing throughout ribbon. Descending
limb/ascus present. Pharyngeal pouches
present. Esophageal gizzard with gizzard
plates absent. Stomach of flow-through type,
without pouch-like chamber. Intestinal
typhlosole absent. Nervous system slightly
streptoneurous. Circumesophageal nerve
ring postpharyngeal. Cerebral/pleural ganglia
fused. Cerebral commissure long; pedal
commissure short. Visceral nerve loop for-
mula: L-A-B-V-PA- -L. Genital nerve emanat-
ing from visceral ganglion; no ganglion seen
in cross-sections. Eye with solid, oblong, ir-
regular lens, with ‘‘anchors,”’ deeply embed-
ded (S/C = 1.3), directed ventrolaterally. Re-
productive system androdiaulic. Gonad with
hermaphroditic acini. Both internal sperm
duct and external ciliated groove present.
Copulatory organ retractile, without glandular
tissue or sperm storage area. Penis with
ejaculatory duct continuous with internal
sperm duct; tip simple. Prostate pallial, of
“flower” configuration. Two distal sperm

440 MIKKELSEN

sacks: larger bursa copulatrix (containing un-
oriented sperm) and smaller receptaculum
seminis (containing oriented sperm); game-
tolytic gland absent. Egg mass kidney-
shaped, without anchor (K. B. Clark, pers.
comm.), with single eggs per capsule (D. De
Freese, pers. comm.). Veliger PMO unknown.

Volvatella

*V. bermudae Clark, 1982—Bermuda,
8/1979, ex К.В. Clark [Н]; Clark, 1982.

V. fragilis Pease, 1860 (type species)—Evans,
1950.

V. cincta Nevill & Nevill, 1869—DMNH 25113,
Ceylon, 1957 [S].

V. ficula Burn, 1966—Burn, 1966; Jensen &
Wells, 1990.

V. laguncula Sowerby, 1894—Thompson,
1979.

V. pyriformis Pease, 1868—Evans, 1950.

V. ventricosa Jensen & Wells, 1990—Jensen
& Wells, 1990.

У. vigourouxi (Montrouzier, 1861)—Baba,
1966 (fide Clark, 1982); Jensen 4 Wells,
1990.

Shell external. Foot simple. Operculum ab-
sent. Parapodia absent. Mantle cavity di-
rected laterally. Plicatidium-type gill one-
sided, attached to mantle over its full length.
Ciliated strips beginning bluntly at mantle
edge; no pallial caecum. Single anterior ad-
ductor muscle present. Jaws absent, but
thickened cuticular epithelium in oral tube.
Radula uniseriate, with formula 0.1.0.
Rachidian tooth dagger-shaped, with mar-
ginal serrations/denticulations. Tooth size in-
creasing throughout ribbon. Descending
limb/ascus present. Pharyngeal pouches
present. Esophageal gizzard with gizzard
plates absent. Stomach of flow-through type,
without pouch-like chamber. Intestinal
typhlosole absent. Nervous system slightly
streptoneurous (Burn, 1966). Circumesoph-
ageal nerve ring postpharyngeal. Cerebral/
pleural ganglia fused. Cerebral commissure
long; pedal commissure short. Visceral nerve
loop formula variable: L-A- -BV-PA- -L (in V.
bermudae, pers. obs.), -PAL (in V. bermudae,
fide Clark, 1982). [LAB- in V. vigourouxi and
V. ventricosa (fide Baba, 1966; Jensen 4
Wells, 1990) is probably an error, mistaking
the left pallial for the subintestinal]. Genital
ganglion on nerve emanating from visceral
ganglion. Eye with solid, irregular lens, with

“anchors,” midway between surface and
nerve ring (S/C = 0.3), directed ventrolaterally
in V. bermudae [and probably, based on lo-
cation, in other species, e.g., V. pyriformis
(fide Evans, 1950), V. laguncula (fide Thomp-
son, 1979)], but dorsolaterally in V. fragilis
(fide Evans, 1950). Reproductive system an-
drodiaulic. Gonad acini? Both internal sperm
duct and external ciliated groove present.
Copulatory organ retractile, without glandular
tissue or sperm storage area. Penis with
ejaculatory duct continuous with internal
sperm duct; tip equipped with stylet. Pros-
tate pallial, of “flower” configuration. Two dis-
tal sperm sacks: larger bursa copulatrix
(containing unoriented sperm) and smaller
receptaculum seminis (containing oriented
sperm); gametolytic gland absent. Egg mass
string-shaped, unanchored (K.B. Clark, pers.
comm.), with single eggs per capsule (Clark,
1982). Larval development unknown.

Akera

"A. bullata О.Е. Müller, 1776 (type species) —
ZMUC, Lemche material (embedded),
Frederikssund, Denmark, 10/1950 [H];
Sweden, Smedjan, Gullmarn, 8/1909, ex
A. Waren [0,$,М]; Sweden, Tjärnö
Marinzoologiska Station, 10/1988, ex A.
Waren [0]; МОС, Roskeeda Bay, west-
ern Ireland, 7/1974 [L] (as named giant
variety A. farrani “Norman” Pilsbry,
1896); Guiart, 1901; Tchang Si, 1931;
Thorson, 1946; Morton 4 Holme, 1955;
Morton, 1972; Williams, 1975; Thomp-
son, 1976; Brace, 1977a.

А. bayeri Marcus & Marcus, 1967—Ev. Mar-
cus, 1970.

Shell external. Foot supplemented by pos-
terior pallial lobe. Operculum absent. Para-
podia present. Mantle cavity directed later-
ally. Ciliated strips beginning bluntly at mantle
edge, continuing into long pallial caecum
that is adherent to visceral mass. Plicatidium-
type gill two-sided, attached to mantle by
suspensory membranes. Two adductor mus-
cles present, one anterior, one posterior (Mor-
ton, 1972). Jaws present, comprised of blunt,
rod-shaped elements. Radula with formula
(19-44).(2-8).1.(2-8).(19-44). Rachidian tooth
rhomboid, with median cusp flanked by
smaller denticles. Lateral teeth hook-like, 2-8
with marginal denticles. Marginals hook-like,
smooth, more slender and elongate than lat-
erals, increasingly so outwardly. Tooth size

PHYLOGENETICS OF CEPHALASPIDEA 441

uniform throughout ribbon. Esophageal giz-
zard with numerous, uncalcified gizzard
plates of varying sizes. Gizzard spines present
preceding (anterior to) plates only; filter cham-
ber following gizzard equipped with fleshy
cartilaginous-capped “spines” only. Stom-
ach of flow-through type, without pouch-like
chamber, small blind caecum present (Morton
& Holme, 1955) or absent (pers. obs.). Intes-
tinal typhlosole absent. Nervous system
slightly streptoneurous. Circumesophageal
nerve ring postpharyngeal. Cerebral/pleural
ganglia separate, hypoathroid. Cerebral com-
missure short; pedal commissure long. Vis-
ceral nerve loop formula: L- -A-BV-PA- -L
(pers. obs., in part; Brace, 1977a). Genital
ganglion appressed to visceral ganglion along
visceral loop. Eye with solid, oblong-oval lens,
close to surface (S/C = 0.08), directed vent-
rolaterally. Reproductive system monaulic.
Gonad with hermaphroditic acini. Internal
sperm duct absent; external ciliated groove
present. Copulatory organ retractile, without
glandular tissue or sperm storage area. Penis
with external groove only, without ejaculatory
duct; tip simple. Prostate? Proximal recep-
taculum seminis, distal gametolytic gland.
Egg mass of tangled strings, unanchored,
with single eggs/capsule. Veliger PMO not
described but figured as colorless (Thorson,
1946: fig. 148B, С).

Aplysia

*А. brasiliana Rang, 1828—HBOM
065:00272, R/V GOSNOLD 229/408, off
St. Lucie Inlet, Florida, 13 m, 4/1974 [0,1];
HBOM 065:00414, Jim Island flat, Ft.
Pierce, Florida, 2/1974 [D,H,S]; HBOM
065:03034, Mullet Key, Florida, 1/1979
[D,M].

californica Cooper,
‚ 1977.

. dactylomela Rang, 1828—Switzer-Dunlap
& Hadfield, 1977.

depilans Gmelin,

Bebbington, 1969.
. fasciata Poiret, 1789—Thompson & Beb-
bington, 1969.

. grandis (Pease, 1860)—Ostergaard, 1950.
. juliana Quoy & Gaimard, 1832—Switzer-
Dunlap & Hadfield, 1977.

punctata Cuvier, 1803—Guiart, 1901;
Saunders & Poole, 1910; Eales, 1921;
Tchang Si, 1931; Howells, 1942;
Thorson, 1946.

1863—Kriegstein,

1791—Thompson 4

SO Ss Ss SS

A. parvula Morch, 1863—Williams, 1975; Vit-
turi et al., 1985.
A. sp.—Kandel, 1979.

Shell internal (but with small aperture in
mantle). Foot simple. Operculum absent.
Parapodia present. Mantle cavity directed
laterally. Ciliated strips and pallial caecum
absent. Plicatidium-type gill two-sided, at-
tached to mantle by suspensory membranes.
Jaws present, comprised of blunt rod-
shaped elements. Radula with formula
16.1.16. Rachidian tooth rhomboid, with me-
dian cusp flanked by smaller denticles. Lat-
eral teeth with denticulate cusps, changing
gradually in cusp configuration from center to
edge of ribbon so that distinction between
lateral and marginal teeth inexact. Tooth size
uniform throughout ribbon. Esophageal giz-
zard with numerous, uncalcified gizzard
plates, varying in size. Gizzard spines present
preceding plates and also in posterior filter
chamber. Stomach of flow-through type,
without pouch-like chamber. Intestinal
typhlosole absent. Nervous system slightly
streptoneurous. Circumesophageal nerve
ring postpharyngeal. Cerebral/pleural ganglia
separate, hypoathroid (but pleural ganglion
only slightly closer to pedal than to cerebral
ganglion). Cerebral commissure short; pedal
commissure long. Visceral nerve loop for-
mula: L---ABVPA---L; Williams (1975)
claimed that part of the left pallial ganglion is
fused with the pleural and part with the sube-
sophageal. Genital ganglion on nerve ema-
nating from visceral ganglion. Eye with solid,
spherical lens, close to surface (S/C not
available), directed dorsolaterally. Reproduc-
tive system oddiaulic. Gonad with hermaph-
roditic acini. Internal sperm duct absent; ex-
ternal ciliated groove present. Copulatory
organ retractile, without glandular tissue or
sperm storage area. Penis with external
groove only, without ejaculatory duct; tip
simple. Prostate uncertain (Eales, 1921;
Thompson & Bebbington, 1969). Proximal re-
ceptaculum seminis, distal gametolytic
gland. Egg mass of tangled strings, unan-
chored; multiple eggs/capsule. Veliger PMO
colorless (Thompson, 1976; Kriegstein, 1977;
Switzer-Dunlap & Hadfield, 1977), although
Saunders & Poole (1910: 513) noted ‘‘drops
of coloured liquid” within its vacuoles. Chro-
mosome number 16 or 17 in numerous ex-
amined species (Inaba, 1959; Patterson,
1969; Natarajan, 1970; Vitturi et al., 1985).

442 MIKKELSEN

Gegania

*G. valkyrie Powell, 1940—NMNZ M.36712,
New Zealand, east of North Cape,
2/1974 [H (1 specimen sectioned by G.
Haszprunar; Haszprunar, 1985b));
MNHN, New Caledonia, 505-515 m,
9/1985 [H,S]; Climo, 1975; Haszprunar,
1985b; Bieler, 1988.

Shell external. Foot simple (Climo, 1975).
Operculum present. Parapodia absent
(Climo, 1975). Mantle cavity directed anteri-
orly. Ciliated strips beginning bluntly at man-
tle margin, extending into a short pallial
caecum. Plicatidium-type gill one-sided, at-
tached to mantle (and rectum) throughout
length. Jaws absent, but cuticularized epi-
thelium present (Haszprunar, 1985b). Radula
with formula 1.2.1.2.1 (Climo, 1975). Rachid-
ian tooth rhomboid, with median cusp
flanked by smaller denticles. Lateral teeth
hook-shaped, denticulate. Marginals plate-
like. Tooth size uniform throughout ribbon.
Esophageal gizzard absent. Stomach of flow-
through type, without pouch-like chamber.
Intestinal typhlosole absent. Nervous system
streptoneurous. Circumesophageal nerve
ring prepharyngeal [but Haszprunar (1985b)
claimed the pharynx can shift for feeding].
Cerebral/pleural ganglia separate [not fused
as Claimed for right pleural by Haszprunar
(1985b)], slightly epiathroid. Cerebral com-
missure short; pedal commissure long. Vis-
ceral nerve loop formula: L-A-B-V-P-AL
(Haszprunar, 19856; pers. obs.). Genital
nerve emanating from visceral ganglion
(Haszprunar, 1985b). Eye with “hollow,” ir-
regular lens, close to surface, directed dor-
solaterally. Reproductive system hermaphro-
ditic, with male and female tracts largely
separate. Gonadal acini with separate male
and female follicles. Copulatory organ and
external ciliated groove absent. Internal

sperm duct present, but opens into mantle
cavity (Haszprunar, 1985b). Prostate absent
(Haszprunar, 1985b). Two proximal allo-
sperm storage sacks in tandem, gametolytic
gland connected by short duct to receptac-
ulum seminis, thence with common duct
opening into mantle cavity in vicinity of
vesicula seminalis or ampulla (Haszprunar,
1985b: fig. 1). Egg mass ball-shaped (‘gelat-
inous strap””), unanchored, with single eggs/
capsule (Climo, 1975). Larval development
planktotrophic according to protoconch mor-
phology (R. Bieler, pers. comm., and 1988:
fig. 14), otherwise unknown.

All-Zero Outgroup

Based (in part) on the “larger outgroup,” the
caenogastropods.

Shell external. Operculum present. Para-
podia absent. Foot simple. Mantle cavity di-
rected anteriorly. Ciliated strips absent. Gill
(ctenidium) two-sided. Jaws present; oral cu-
ticle smooth. Radula of uniform tooth size.
Rachidian tooth rhomboid; lateral/marginal
teeth of more than one form. Esophageal giz-
zard absent. Stomach with pouch-like cham-
ber. Intestinal typhlosole partial. Nervous
system streptoneurous. Circumesophageal
nerve ring prepharyngeal. Cerebral/pleural
ganglia separate. Cerebral and pedal com-
missures long. Visceral nerve loop formula:
LA-B-V-P-AL. Genital nerve emanating from
visceral ganglion. Eye with solid, spherical
lens, close to surface, directed dorsolaterally.
Reproductive system dioecious. Homologue
of internal sperm duct present (but open or
closed status undetermined); ejaculatory
duct and lateral external ciliated groove ab-
sent. Copulatory organ nonretractile, without
autosperm storage bulb. Receptaculum
seminis proximal; bursa copulatrix (homo-
logue of gametolytic gland) distal.

MALACOLOGIA, 1996, 37(2): 443-511

MORPHOLOGY AND PHYLOGENETIC RELATIONSHIPS OF CERTAIN
PYRAMIDELLID TAXA (HETEROBRANCHIA)

John B. Wise

Houston Museum of Natural Science, One Hermann Circle Drive,
Houston, Texas 77030, U.S.A.

ABSTRACT

The marine gastropod family Pyramidellidae is poorly known. Although numerous and world-
wide, the anatomies of only a few species are known, and our understanding of this family’s
taxonomy and systematics is based almost entirely on shell characters.

Eight pyramidellid genera and 12 species were dissected, sectioned, and examined with
SEM. Traditionally used gastropod characteristics are either absent (e.g., radula) or of little use
(e.g., reproductive system minus the penis), because they are undiversified morphologically in
the taxa examined herein. Characters of gut, mantle cavity, and penial complex proved most
useful in developing an understanding of how the taxa in the present study may be defined.

Phylogenetic analysis of 13 taxa and 28 characters yielded six equally parsimonious cla-
dograms of 67 steps and a consistency index of 68%. New systematic standards are proposed
for defining (on the basis of synapomorphies) three of the four traditional pyramidellid subfam-
ilies, the new subfamily Sayellinae, and the new genera Houbricka and Petitella.

To test the hypothesis that protoconch configuration is a reflection of developmental mode
and not phylogeny, protoconch characters were eliminated from a second phylogenetic anal-
ysis. This yielded one cladogram, which when a taxon’s developmental mode and host(s) are
known, support the contention that protoconch shape is a result of developmental mode.

Historically shell characters, to the exclusion of soft-part anatomy, have been used to assign
taxa to the various pyramidellid genera. Therefore, it might be tempting to rely more on ana-
tomical characters and treat shell characters a priori as homoplasious. However, this study
shows that members of the subfamily Pyramidellinae could only be distinguished by concho-
logical characters, whereas in other taxa soft-part anatomy proved the most phylogenetically
useful. The present study indicates that all characters (= total evidence) should be used in any

phylogenetic analysis.

Key words: morphology, phylogeny, Pyramidellidae, synapomorphies, cladograms.

INTRODUCTION

The Pyramidellidae Gray, 1840, are marine
ectoparasitic gastropods that occur in boreal
to tropical waters worldwide, and from the
intertidal zone to several thousand meters.
Pyramidellids feed on a variety of invertebrate
hosts (Robertson, 1957; Ankel & Christensen,
1963; Robertson & Orr, 1961; Fretter & Gra-
ham, 1962; Scheltema, 1965; Boss & Merrill,
1965; Bullock & Boss, 1971; Robertson &
Mau-Lastovicka, 1979; Boss, 1982). They
pierce the host's tissues with a buccal stylet
and remove host body fluids by the muscular
action of their buccal pump (Ankel, 1949a;
Fretter & Graham, 1949; Maas, 1965; Wise,
1993).

The systematic position of the Pyrami-
dellidae has been controversial for over 130
years (Boss, 1982). This controversy, al-
though caused in part by the lack of informa-

443

tion about this family, is also due to changing
views about gastropod phylogeny (for a
review of the current state of gastropod phy-
logeny and systematics, see Bieler, 1992).
Traditionally, gastropods have been divided
into three subclasses: Prosobranchia, Opis-
thobranchia, and Pulmonata, with the proso-
branchs as primitive gastropods giving rise to
both opisthobranchs and pulmonates. In this
scheme, pyramidellids occupy an intermedi-
ary position between the prosobranchs and
opisthobranchs (Boss, 1982). Because they
have a spirally coiled calcareous shell into
which the entire body is retractable, a foot
with an operculum, a long proboscis, and an
anteriorly oriented mantle cavity, most early
authors placed them in the Prosobranchia,
but because they also have such character-
istics as a pallial kidney, subepithelial eyes on
the median side of the tentacles, an ovotestis,
and a heterostrophic protoconch, later au-

444 WISE

TABLE 1. A list of authors and the subclasses to
which they assigned the Pyramidellidae.

Author Subclass

Mörch, 1865
Pelseneer, 1899
Thiele, 1929-35
Wenz, 1938-44
Thorson, 1946

Opisthobranchia
Prosobranchia
Prosobranchia
Prosobranchia
Prosobranchia

Fretter & Graham, 1949 Opisthobranchia
Risbec, 1955 Prosobranchia
Boettger, 1955 Euthyneura*
Knight et al., 1960 Opisthobranchia
Taylor & Sohl, 1962 Opisthobranchia
Ghiselin, 1966 Opisthobranchia
Maas, 1965 Opisthobranchia
Hyman, 1967 Opisthobranchia
Golikov 8 Prosobranchia

Pectinibranchia
Opisthobranchia
Prosobranchia
Sinsitrobranchia
Opisthobranchia
Prosobranchia
Prosobranchia
Heterobranchia
Heterobranchia

Starobogatav, 1975
Thompson, 1976
Minichev &

Starobogatav, 1979
Salvini-Plawen, 1980
Gosliner, 1981
Robertson, 1985
Haszprunar, 1985b, 1988a
Ponder & Warén, 1988

*Placed pyramidellids within Cephalaspidea, with the
Acteonidae giving rise to the Pyramidellidae.

thors placed them in the Opisthobranchia
(Table 1). At present, many authors view the
three classic subclasses as artificial, and sev-
eral revisionary schemes have been pro-
posed, although none have met broad accep-
tance (Brusca & Brusca, 1990). For example,
in Haszprunar's (1985b) system, the Gas-
tropoda are divided into two subclasses: the
Caenogastropoda (= Prosobranchia) and the
Heterobranchia (= Opisthobranchia, Pulmo-
nata, and “allogastropods””). The Pyramidel-
lidae are placed within the Heterobranchia,
superorder Allogastropoda, which unites the
Architectonicoidea, Pyramidelloidea, and the
fossil Nerineoidea. Most recently, Haszprunar
(1988a, 1990) and Ponder & Warén (1988) as-
sign the pyramidellids to the order Heteros-
tropha of the subclass Heterobranchia on the
basis of several purported, but cladistically
untested synapomorphies (e.g., lateral and
rhinophoral nerves, giant neurons, ciliated
strips, heterostrophy, sperm morphology,
and chalazae). In this system, the pyramidel-
lids are basal heterobranchs and represent
an evolutionary link between the two sub-
classes (Gosliner, 1981; Robertson, 1985;
Haszprunar, 1985a, b, c, 1988a, b, 1990;
Healy, 1988a, b, 1993).

Confusion about the systematics of the
pyramidellids and uncertainty about their role
as ectoparasites is due, largely, to the lack of
information about them. To date, the anatom-
ical knowledge of a few species serve as the
paradigm for a very large family (Ankel, 1949a,
b, 1959; Fretter & Graham, 1949, 1962; Fret-
ter, 1951; Risbec, 1955; Maas, 1963, 1965;
Höisaeter, 1965; Brandt, 1968; Kristensen,
1970; Robertson, 1974, 1978, 1985; Haszpru-
nar, 1985a, b, c, 1988a, b; Ponder, 1973,
1987; White, 1985; Wise, 1993).

The monophyly of the Pyramidellidae is
supported by characters of the alimentary
tract (e.g., buccal stylet) and mantle cavity
(e.g., pigmented mantle organ) (Haszprunar,
1988a). However, relationships among mem-
bers ofthe Pyramidellidae are unclear. In most
classifications, the family is subdivided into
four subfamilies: Cyclostremellinae Moore,
1966; Odostominae Pelseneer, 1928; Py-
ramidellinae Gray, 1840; and Turbonillinae
Simroth, 1907 (Ponder and Warén, 1988).
Nordsieck (1972) presented an alternative
view. Traditionally, the assignment of taxa to
a subfamily and how groups within these sub-
families have been defined has been based on
shell characters (Tryon, 1886; Dall 8 Bartsch,
1904, 1906, 1909; Bartsch, 1909, 1917, 1955;
Nomura, 1936, 1937, 1938, 1939, 1940; Laws,
1937a-d, 1938, 1939, 1940, 1941; Laseron,
1959; Nordsieck, 1972; Aartsen, 1977, 1981,
1987; Gofas et al. 1981; Linden 4 Eikenboom,
1992). Shell characters, however, have been
shown to be convergent when snails live in
similar habitats and may be unreliable indi-
cators of phylogenetic relationships (Davis,
1979; Kool, 1993).

The objectives of this study are to develop
a more comprehensive understanding of
pyramidellid anatomy, add to our limited
knowledge of this group's biology, and pro-
vide a phylogenetic framework upon which to
build a more comprehensive classification for
the family.

MATERIAL AND METHODS
Sample Material

Eight genera and 12 species (representing
three of the four pyramidellid subfamilies)
were collected alive for dissection, fixation,
and histological examination (Table 2). The
subfamily Cyclostremellinae was not exam-
ined because no specimens of the animals
were available (i.e., only shells were available).

PYRAMIDELLID MORPHOLOGY AND PHYLOGENY 445

TABLE 2. Collection localities for taxa examined this study.

Taxa

Boonea стс (Carpenter, 1864)
Boonea seminuda* (С. В. Adams 1839)

Locality

Palos Verdes, Los Angeles (34°12’N, 119°20’W), California
Wild Harbor (41°33’N, 70°36’W) & Bass River (41°40’N,

70°11’W), Massachusetts

Odostomia babylonia (C. B. Adams, 1845)
Odostomia didyma Verrill & Bush, 1900
Sayella hemphillii* (Dall, 1884)

Petitella crosseana* (Dall, 1885)
Pyramidella sulcata (A. Adams, 1854)

Indian Fill Key (24°54’N, 80°42’W), Florida

Indian Fill Key, Florida

Cedar Key (29°08’N, 83°02’W), Florida

Ft. Pierce (27°35’N, 80°19’W), Florida

Pago Bay (13°25’N, 144°48’W) & Tumon Bay (13°31’N,

144°48’W), Guam

Pyramidella crenulata (Holmes, 1859)
Pyramidella mitralis A. Adams, 1854
Turbonilla hemphilli Bush, 1899
Houbricka incisa* (Bush, 1899)
Tathrella iredalei* Laseron, 1959

*Туре species of genus

At the beginning of this study, it was my in-
tention to add the number of genera examined
by using available museum material. Holo-
types (shells only) for each species were
examined. However, museum specimens
proved unusable because they had been
placed in alcohol without first cracking the
shell and the animal’s bodies were not pre-
served.

Snails from Florida were collected through-
out the year by one of two methods. In the
first, the topmost substratum (approximately
2-4 cm) from inter- to subtidal sand and mud
flats was placed in a 0.5 mm sieve, rinsed with
seawater to remove silt and mud, and sorted
under a dissecting microscope. The second
method involved rinsing the underside of em-
bedded rocks or coral rubble with seawater,
and examining this debris under a dissecting
microscope for pyramidellids. In Massachu-
setts (October 1991), Boonea seminuda were
found on the slipper shell Crepidula fornicata.
In Guam (July 1990), Pyramidella mitralis
(Otopleura mitralis in previous literature) and
P. sulcata were collected during night dives in
both Tumon and Pago bays. During the day,
these snails remain submerged within the
sand, but at night they were easily collected
as they crawled on top of the substratum.
Tathrella iredalei was found on the shells of
the various Tridacna spp. (giant clam) at
Guam’s Fadian Hatchery. Boonea cincta
(Chrysallida cincta in most previous works)
was found (August 1990) on the opercula of
Tequla eiseni found inter- to subtidally on the
rocks at Palos Verdes, Los Angeles County,
California. Snails were kept alive in bowls of
aerated seawater.

Ft. Pierce 8 Cedar Key, Florida

Pago 8 Tumon bays, Guam

Ft. Pierce, Florida

Ft. Pierce, Florida

Fadian Hatchery (13°26’N, 144°49’W), Guam

Morphology
Light Microscopy
(a) Observations of Living Animals

Living snails were observed and their habits
noted. Photographs were taken with a Pentax
35 mm camera mounted on a Zeiss Tessavar
dissecting microscope.

(b) Gross Dissection

Snails were prepared for dissection by first
cracking their shells with a vise. Snails were
dissected in toto, and structures of the ali-
mentary tract and penial complex were ex-
cised and examined using either a compound
or a dissecting microscope equipped with an
ocular micrometer. Whole snails and their
parts were routinely stained with toluidine
blue to facilitate distinguishing the various or-
gans and organ systems. Photographs were
taken with a Polaroid camera (using type 52
Polapan Land film) mounted on a Nikon La-
bophot compound microscope.

(c) Histology

Snails were removed from their shells ei-
ther by decalcification or by cracking the
shell with a vise. The first method, utilizing a
commercial decalcifier (Decalcifying solution,
Krajian, J. T. Baker) to dissolve the shell, was
used when serial sections of the entire snail
were desired. Tissues were fixed in 10% for-
malin buffered in filtered seawater. Speci-
mens were embedded in paraffin, sections
were cut at 4-6 um, and stained with hema-
toxylin and eosin-Y (Sheehan & Hrapchak,

446

WISE

TABLE 3. Character-state distributions for 12 pyramidellid taxa and 28 characters.

Characters
1 2
Taxa MAS SS OA SA SS the) O A 2s 4 5 6 7 &
OUT OOOO OOOO OO Oy OO Os O bo) O OF OOO OOOO OOO
GING 2 4 a OOO a ae a 2071770207 отт (0 2 À ? ©
Ем 9 O 12.0202 03025 1 ZH ото Oo 8 02 1 1 002300
JUDI 0001000011321 TAO O 2 1%Y0
DIDY 227070907 Oa tie tS 2707 17707737 O O 210
SEME? 3) 2) 2) 10) 0 10) 117272020207 Оооо
ans sea 22 0 0 0 1 2° Oo oh 2 a oo 2 0 oy i too © 1 © ©
SUL OO ооо ооо ao) aa aoa BS 1 OR O OR
< @ тт OOO OO tt @O ia Wisi 10000
MIT (0 OOS 22 WO ON ON O OO a a ao a a ab as 1 1 oO OC O C
AO a OM? Юон т 24 i 2 oO OO 2ii i
INC 1 OU oo) ON Oa OO OO aw @ a dtd dy oO oO O 1 oO
IRED то? 0 001.204.071 II OR aI ae т aOR OI À

(Abbreviations: OUT = Amathinidae, CINC = Boonea cincta, SEMI = Boonea seminuda, JUDI = Odostomia babylonia, DIDY
= Odostomia didyma, SEMP = Sayella hemphillii, CROS = Petitella crosseana, SUL = Pyramidella sulcata, CREN = P.
crenulata, MIT = P. mitralis, HEMP = Turbonilla hemphilli, INC = Houbricka incisa, IRED = Tathrella iredalei).

1980). Specimens were often secured within
small pieces of Cucumis sativus (common
cucumber), also fixed in 10% formalin buff-
ered in seawater. This was done prior to em-
bedding when the snail’s orientation was par-
ticularly important. Photographs were taken
with a 35 mm camera (using Kodak T-Max
film 100 ASA) mounted on a Nikon Labophot
compound microscope.

Scanning Electron Microscopy
(a) Hard Parts

Shells and opercula were cleaned by
sonication, air dried, coated with gold-pal-
ladium, and examined with either a Cam-
bridge S-100, Selectron 250, or Hitachi
S-570 scanning electron microscope op-
erating at 4-10 KEV.

(b) Soft Parts

Tissue specimens were fixed in 2%
glutaraldehyde buffered in 0.025 M sodium
cacodylate in seawater. Postfixed tissues
were thoroughly rinsed in sodium cacody-
late buffered in filtered seawater. Speci-
mens were dehydrated in a graded series
of ethyl alcohol, critical point dried, and
coated and examined as above.

Phylogenetic Analysis
Twenty-eight characters were analyzed for

eight genera and 12 species of pyramidellids.
Characters were obtained from shell and soft-

part anatomy. At no time were characters
eliminated or included on the basis of any
preconceived ideas of how they might influ-
ence the outcome of the phylogenetic anal-
yses. The distribution of the states of the char-
acters is shown in Table 3.

The method of character analysis used to
determine relationships of the taxa was phy-
logenetic systematics (i.e., cladistics) (Hen-
nig, 1966; Lipscomb, 1984; Schuh & Farris,
1981; Farris, 1982, 1983). These relationships
are expressed in cladograms that were con-
structed using the computer program Hen-
nig86 (Farris, 1988). No a priori character
weighting was employed. Successive weight-
ing (Hennig86 option w xs;) was used to
choose between equally parsimonious cla-
dograms that were produced when the data
set was reanalyzed without the protoconch
characters. Successive weighting selects cla-
dograms that require the fewest number of
characters to have homoplasies (i.e., trees
with the shortest length and fewest changing
characters) (Carpenter, 1988; Lipscomb,
1993).

Characters were polarized using the out-
group comparison method (Hennig, 1966;
Watrous & Wheeler, 1981; Schoch, 1986). The
family Amathinidae was chosen as the out-
group, because it appears to be the sister
group (i.e., most closely related) to the Pyra-
midellidae and is the only other family within
the superfamily Pyramidelloidea.

Transformation series were determined (af-
ter polarization) for all multistate characters

PYRAMIDELLID MORPHOLOGY AND PHYLOGENY 447

TABLE 4. Transformation(s) for each multistate
character. Italicized transformation series, although
proposed initially were rejected, as they were in-
congruent with cladogram groupings constructed
of other homologies.

Character Transformation
1 0-1,2-3
$: 0-1,2
4 0-2-1-3
5 0-1-2
6 0-1-2
9 0-1-2; 0-1,2
13 0-3,2-1
14 0-1,2; 0-2-1
19 0-1-2-3
22 0-1,2,3; 0-1-2-3
25 0-1-2
26 0-2-1

(Table 4) using the homology method outlined
by Lipscomb (1992).

The relative quality of the phylogenetic re-
sults was judged using the consistency index
(Cl), a measure of the degree to which char-
acter state changes on a cladogram are min-
imal (Kluge & Farris, 1969), and the retention
index (Rl), a measure of the amount of ho-
mology hypothesized by the data set that is
retained on the tree (Farris, 1989).

Autapomorphies were eliminated from all
analyses. Although autapomorphies are very
useful in defining terminal taxa, they provide
no information about how taxa are related to
each other. Moreover, by including them in
the analyses the Cl is superficially inflated
(Farris, 1989).

Institutional abbreviations are as follows:

AMNH American Museum of Natural His-
tory, New York, U.S.A.

ANSP Academy of Natural Sciences of
Philadelphia, Pennsylvania, U.S.A.

AMS Australian Museum, Sydney, Aus-

| tralia

BMNH The Natural History Museum, Lon-
don, U.K.

CASIZ California Academy of Sciences,
San Francisco, U.S.A.

MCZ Museum of Comparative Zoology,
Cambridge, Massachusetts, U.S.A.

USNM National Museum of Natural His-
tory, Smithsonian Institution, Wash-
ington, D.C., U.S.A.

PM Peabody Museum, Yale University,

U.S.A.
Voucher specimens on deposit at USNM.

Superfamily Pyramidelloidea Gray, 1840
Family PYRAMIDELLIDAE Gray, 1840

Diagnosis: Shell sharply lanceolate to roughly
planispiral. Shell sculpture variable. Smooth,
heterostrophic protoconch oriented 90-150?
to teleoconch and often partially submerged
within succeeding adult whorl. Shell aperture
elongate-lenticular to ovate, with or without
columellar folds. Operculum paucispiral, with
subcentric nucleus. Alimentary tract com-
prised of acrembolic proboscis, buccal sac,
buccal pump, esophagus (undifferentiated or
divided into anterior and posterior sections),
and a pair of salivary glands. Buccal sac con-
taining piercing stylet. Simultaneously her-
maphroditic. Euthyneurous nervous system
highly concentrated and epiathroid.

Remarks: The Pyramidellidae is a large
pandemic family. Depending on the author
consulted, it contains 35-75 genera and 800-
1000 or more species. The current taxonomy
is based primarily on shell characters and 1$
both disputatious and conjectural (Abbott,
1974; Boss, 1982).

Discussion: Shell: Shell shapes range from
planispiral to acutely lanceolate. Pyramidel-
lids, although generally small (average shell
length 6 mm), may attain lengths of 50 mm.
Shell sculpture, when present, varies and can
be microscopic and/or macroscopic, with ax-
ial and spiral lines, axial ribs, and nodes. Su-
tures may be deep, shallow, shouldered, or
crenulate. The heterostrophic protoconch is
smooth and oriented 90-150° to the teleo-
conch. The protoconch configurations vary
among genera. The shell aperture is generally
elongate to ovate, with or without columellar
folds, and palatal teeth may be present within
the outer lip. The thin, paucispiral operculum
has a subcentric nucleus. When columnar
folds are present, the operculum may be
notched to accommodate them.

Head-foot: Pyramidellids have a well-de-
veloped head, a pair of cephalic tentacles,
and a large foot with an operculum that
tapers posteriorly to either a blunt or acute
apex. The epidermis of the tentacled head,
mantle, and foot is lined with one layer of
simple columnar or cuboidal cells. These
cells have basal nuclei and are ciliated on the
ventral surface of the foot, the lower antero-
dorsal portion of the propodium, and the

448 WISE

mentum. Head-foot and mantle contain large
basophilic, subepidermal gland cells. These
cells discharge granulated droplets of mucus
in the cytoplasm, and the droplets are dis-
charged directly through the cell membrane
into the space between the epidermal cells.
This mucus coats the external surface of the
mantle and head-foot. When present, the
posterior pedal gland lies in a medial position
just dorsal and parallel to the ventral surface
of the foot. It has an invaginated layer of cil-
iated epithelial cells surrounding a lumen.
Gland cells containing sulfated mucins that
are stained dark purple by hematoxylin and
eosin fill the pedal gland. The pedal gland
opens on the postero-ventral surface of the
foot. Snails that produce an attachment
thread anchor themselves to the substratum
or to their host. The pedal sinus complex,
located within the lower portion of the foot
consists of a series of sinuses surrounded by
nucleated connective tissue. Muscle fibers
radiate from the columella muscle into the
head-foot and are interspersed throughout
the gland cells and hemolymph sinuses.

Pyramidellid tentacles have often been de-
scribed as rabbit-ear or donkey-ear in shape
(Fretter & Graham, 1949), but such descrip-
tions oversimplify their variability and com-
plexity of the structure. Members of the
Odostominae have a tentacular pad com-
posed of a distinctive cluster of long cilia lo-
cated inside and subterminal to the tentacle
apices (Ponder, 1973). Fretter & Graham
(1949) suggested these tentacular pads were
sensory in nature and constructed of many
fused cilia. Darkly pigmented eyes with a lens
are subepithelial and on the median side of
the tentacles. Eyes are usually round to
ovate, but may be lenticular. Spacing or dis-
tance between the eyes varies among spe-
cies.

The variably-shaped mentum is located
just ventral to the head and extends, shelf-
like, over the propodium (Fig. 11A-K shows
the mentum shapes of the snails examined
this study). Its function is usually locomotion.
К is the first part of the crawling snail in con-
tact with the substratum. Histologically, the
mentum is indistinguishable from the foot,
but it is innervated by the cerebral rather than
pedal ganglia (Huber, 1987).

The mantle and its organs are similar for all
pyramidellids (Fig. 1A-C). The long, wide an-
terior mantle (= skirt) narrows posteriorly to
meet the visceral mass. Its right, anterior por-
tion forms a short canal or siphon.

A

mae re EIER
о

VCS

FIG. 1. Diagram of pyramidellid mantle cavities and
organs: А. Subfamily Odostominae (bar = 500 um);
B. Subfamily Pyramidellinae (bar = 1 mm); C. Sub-
family Turbonillinae (bar = 500 um) (а = anus, des =
dorsal ciliated strip, glvcs = gland ventral ciliated
strip, h = heart, mae = mantle edge, те = mentum,
о = osphradium, рад = pallial gonoduct, pk = pallial
kidney, pmo = pigmented mantle organ, vcs = vis-
ceral ciliated strip, vm = visceral mass).

Pallial cavity: All pyramidellids have ventral
and dorsal ciliated strips (sensu Fretter &
Graham, 1949, 1962) on the right side of the
mantle cavity (Figs. 1A-C, 2A, ЗА). The dorsal

PYRAMIDELLID MORPHOLOGY AND PHYLOGENY 449

strip hangs from the mantle roof immediately
dorsal to the ventral strip. These strips join on
the mantle roof at the posterior end of the
mantle cavity. Both of these strips are con-
structed of a single layer of ciliated columnar
cells secured to a basal lamina (Fig. 2B). The
beating of the cilia is responsible for the in-
take and left-to-right movement of water
within the mantle cavity.

A gland underlies the ventral strip (Fretter 8,
Graham, 1962: 126). The ventral ciliated strip
gland may extend the entire length of the
strip (e.g., Pyramidella, Fig. 1B) or may un-
derlie only 20-25% of its most anterior por-
tion (e.g., Odostomia and Boonea, Fig. 1A).
The gland is comprised of large cells filled
with a viscid substance (Fig. 2C), which is
sometimes released when the snail is dis-
turbed (Table 5).

The pallial kidney is a long, tubular, narrow
organ suspended from the mantle roof (Figs.
1А-С, ЗА). It extends anteriorly from the heart
at the visceral mass-mantle cavity junction,
to immediately posterior to the pigmented
mantle organ. Histologically, it consists of a
series of thin-walled, slightly basophilic
chambers or tubules (Fig. 3B). The papilla-
like nephridiopore is located subterminally on
the antero-ventral surface of the kidney (Fig.
ЗС, D).

A pigmented mantle organ of large, rect-
angular, and often multi-colored cells, is
present (Figs. 3A, 4A). Genera examined
have one of three shapes: (1) small and ob-
long (Fig. 1A), (2) large and rectangular, sur-
rounded by a field of transparent cells (Fig.
1В), and (3) very large and elongate (with
wide anterior and attenuated posterior ends),
composed of many large transparent cells
mixed with a few white opaque cells (Fig. 1C).
In several genera, this organ produces and
releases an exudate when the snail is dis-
turbed (Table 5). Both Fretter (1951) and Pon-
der (1987) identified this structure as the hy-
pobranchial gland. However, histologically
and positionally, it is unlike the hypobranchial
gland of other gastropods (Robertson, 1985;
present study).

Only members of the Pyramidellinae have
a foliobranch gill (sensu Robertson, 1974).
This gill, first described by Risbec (1955), is
composed of folds oriented perpendicular to
and between the ciliated strips on the right
side of the mantle roof (Fig. 4B, C). This
highly folded structure ostensibly functions in
gas exchange; however, it is not homologous
with the gastropod ctenidium (Ponder, 1987;

FIG. 2. A. SEM microphotograph of the ventral cil-
iated strip of Pyramidella sulcata (bar = 80um); В.
Longitudinal section of the ventral ciliated strip of
P. sulcata (bar = 40 um); C. Longitudinal section of
the gland of Pyramidella sulcata underlying ventral
ciliated strip (bar = 200 um) (c = cilia, des = dorsal
ciliated strip, glc = gland cells, vcs = ventral ciliated
strip).

450

WISE

TABLE 5. Snail exudate origin and characteristics.

Taxa PMO Cells GLVCS Exudate Secreted by
CINC bright yellow & a few brown orange cream-colored bright yellow pmo
8 red
SEMI bright yellow 8 a few green, clear, pink, white or light bright yellow pmo
or brown orange
JUDI bright yellow & a few black, red cream-colored bright yellow pmo
or brown
DIDY bright yellow, red, brown, orange, cream-colored bright yellow pmo
8 black
SEMP transparent 4 white with red, darkly pigmented with milky-blue glvcs
yellow & orange a few red
CROS yellow & orange or black & white black & white light blue pmo
SUL transparent & opaque yellow with a few red light blue glvcs
& white
CREN transparent & opaque yellow with a few red light blue glvcs
& white
MIT opaque & clear, with a few red yellow with a few red light blue glvcs
& yellow & white
HEMP clear ringed by yellow yellow & white yellow glvcs & pmo
INC clear, yellow & red blue atop transparent bright yellow pmo
matrix
IRED yellow & a few white yellow bright yellow glvcs & pmo

ABBREVIATIONS: CINC = Chrysallida cincta, ЗЕМ! = Boonea seminuda, JUDI = Odostomia babylonia, DIDY = О. didyma,
SEMP = Sayella hemphillii, CROS = Petitella crosseana, SUL = Pyramidella sulcata, CREN =P. crenulata, MIT = Pyramidella
mitralis, HEMP = Turbonilla hemphilli, INC = Houbricka incisa, IRED = Tathrella iredalei; glvcs = ventral ciliated strip, pmo

= pigmented mantle organ.

Haszprunar, 1988a). Typically, the gastropod
ctendium is located on the left side of the
mantle cavity, has a complex series of affer-
ent and efferent blood vessels (lacking in
pyramidellids), and is composed of filaments
attached to a central axis (also absent in
pyramidellids).

Fretter & Graham (1949) described the lo-
cation of the anus in Odostomia spp. and
Chrysallida spp. as on the extreme left side at
the inner most end of the mantle cavity. How-
ever, in all the taxa examined in this study
(e.g., Boonea cincta, Odostomia babylonia, O.
didyma), the rectum terminates as an anal
papilla extending from beneath the common
genital duct at the posterior end of the right
side of the mantle floor (Figs. 1A, C, 4D). Fret-
ter (1951), however, described the position of
the anus in Turbonilla elegantissima and T.
jeffreysii as in the taxa | examined.

The simple osphradium is composed of
white, elliptical cells located beneath the ep-
ithelium on the extreme left side of the mantle
roof. In the subfamily Pyramidellinae, part of
the osphradium extends across the mantle to
terminate at the right anterior corner of the
mantle (Fig. 6B).

Alimentary tract: In the Odostominae, the
location of the introvert-proboscis aperture is
medial, on the ventral side of the head, dorsal

to the mentum base. In both the Turbonillinae
and Pyramidellinae, this aperture is medial
and at the anterior apex of the mentum. In
the retracted condition, the introvert extends
posteriorly, to pass through the nerve ring
and enter the cephalic hemocoel. Although
the configuration and number of alimentary
structures is variable for pyramidellids, a sim-
ilar ground plan is shared by all taxa (Fig. 5A):
there is an acrembolic proboscis (= introvert),
buccal sac (containing sucker, stylet with
cuticular sheath, and stylet bulb), one or two
esophagi, and a pair of salivary glands. In
some genera (e.g., Boonea and Odostomia),
a separate oral tube connecting the mouth
and buccal pump is present (Fretter & Gra-
ham, 1949; Wise, 1993).

Reproductive system: Pyramidellids are si-
multaneous hermaphrodites with both ovary
and testis within the lobules of the single go-
nad (= ovotestis). The gonad is located on the
concave side of the upper visceral coils. The
reproductive system is monaulic (i.e., POS-
sessing an undivided pallial gonoduct). Gen-
era examined here have a common pallial
gonoduct, extending anteriorly beneath the
mantle floor to open on the right side of the
head anterior to the right tentacular base
above the dorsum of the foot. In some pyra-
midellid genera, the aperture of the common

PYRAMIDELLID MORPHOLOGY AND PHYLOGENY 451

FIG. 3. A. SEM microphotograph of pigmented mantle organ, pallial kidney, and ciliated strips of Pyra-
midella mitralis (bar = 250 um); B. Longitudinal section of the kidney of P. mitralis (bar = 50 um); C. SEM
microphotograph of anterior portion of the kidney in Fig. В. (with nephridiopore) (bar = 50 um); D. Enlarge-
ment of nephridial opening (bar = 10 um) (c = cilia, cs = ciliated strips, ne = nephridiopore, pk = pallial

kidney, pmo = pigmented mantle organ, tu = tubules).

pallial gonoduct is reported to be on the
mantle floor or the neck (Fretter & Graham,
1949, 1962; Robertson, 1978). Moreover,
Fretter 8 Graham (1949, 1962) described an
open, ciliated sperm groove connecting the
gonoduct aperture to the penial aperture in
Odostomia unidentata. In contrast, no cili-
ated sperm groove or ciliated sperm convey-
апсе is present in any of the taxa | examined,
as was determined by Robertson (1978) for
Boonea spp. and Fargoa spp. Although, as
suggested by Hadfield & Switzer-Dunlap
(1984), in the absence of a ciliated sperm
groove, a closed vas deferens should be
present, no vas deferens were found in the
taxa examined herein. The penis aperture is
medial and immediately ventral to the men-
tum. Until recently (Ponder, 1987), it was be-
lieved that all pyramidellids possess a penis
that lies beside the proboscis within the
nerve ring (Fig. 22B, D). However, six of the
eight genera examined herein have their pe-
nes within the head-foot and outside and
ventral to the nerve ring (Figs. 22А, С, 12A-

С). Penis configuration is highly variable
among these genera. Pyramidellids, opistho-
branchs, pulmonates, and some of the su-
perfamilies within the order Heterostropha
share a modified spermatozoon that pos-
sesses a distinctive acrosome and a complex
mitochondrial derivative (= paracrystalline
matrix and glycogen components surround-
ing the axoneme) (Healy, 1988a, b, 1993).
Within the visceral mass, the ovotestis is
connected to the seminal vesicle via the nar-
row hermaphroditic duct. A short duct con-
nects the seminal vesicle to the coelomic
gonoduct (Fig. 5B). This area of the gonoduct
is the fertilization chamber and the conver-
gence points for the seminal receptacle, al-
bumin and mucous glands, and pallial gland
(Fig. 5B) (Ponder, 1987). Fretter & Graham
(1949, 1962) described two mucous glands
for Odostomia spp. and Chrysallida spp. and
only one for Turbonilla elegantissima. The
proximal portion of the pallial duct (= pallial
gland) appears to function in the encapsula-
tion of the fertilized eggs prior to oviposition.

452 WISE

~

FIG. 4. A. Transverse section of the pigmented mantle organ and gill of P. sulcata showing cells composing
the gill’s filamentous folds (bar = 75 um); B. SEM microphotograph of the gill of Pyramidella sulcata (bar =
100 um); С. SEM microphotograph of the rectum and anal opening of P. mitralis (bar = 40 шт) (а = anus,
cs = ciliated strip, gf = gill fold, glc = gland cells, pmo = pigmented mantle organ).

The anterior, distal portion of the pallial gon-
oduct may function as a prostate (Fretter &
Graham, 1949, 1962).

Nervous system: (Fig. 6A) In pyramidellids,
the nervous system (minus the osphradial
ganglion) is comprised of a highly concen-
trated ring within the head that encircles the
alimentary tract and, in some genera, the pe-
nis (Fretter & Graham, 1949; Huber, 1987;
this study). The nervous system is further de-
scribed as being epiathroid, because the
pleural ganglia lie adjacent to cerebral gan-
glia (Haszprunar, 1988a). The outlying os-
phradial ganglion is connected to the suprae-
sophageal by a long nerve extending across
the nerve ring immediately anterior to the
proboscis. In the Pyramidellinae, the portion
of the osphradium that extends across to the
right side of the mantle is innervated by a
nerve arising from the osphradial nerve, lo-
cated approximately one-half the distance to
the osphradial ganglion (Fig. 6B). The pres-

ence of the osphradium and its ganglion on
the snail’s left side suggests its euthyneurous
condition (= untwisted visceral loop) is a re-
sult of concentration of the nerve ring and not
detorsion (Fretter & Graham, 1949; Haszpru-
nar, 1985c, 1988a).

The arrangement of the nervous system for
the taxa examined in the present study is as
described by Fretter & Graham (1949) and
Huber (1987, 1993), with some exceptions:
(1) Members of the subfamily Pyramidellinae
examined here have a nerve extending from
the osphradial nerve (originating at the su-
praesophageal ganglion) to innervate a por-
tion of the osphradium that extends to the
right anterior mantle corner just posterior of
the mantle edge (Fig. 6B); (2) labial ganglia
described by Fretter & Graham (1949) for
constituents of the Odostominae, were not
apparent in any of the taxa examined this
study; (3) the subesophageal ganglion has
been depicted as spheroid and identical to

PYRAMIDELLID MORPHOLOGY AND PHYLOGENY 453

B ES

| abel \

= visceral mass
mantle cavity

FIG. 5. A. Diagram of generalized pyramidellid
feeding structures; B. Diagram of generalized pyra-
midellid reproductive system (abgl = albumin
gland, bp1 = buccal pump 1, bp2 = buccal pump 2,
bs = buccal sac, es = esophagus, fc = fertilization
chamber, hd = hermaphroditic duct, mg! = mucous
gland, ov/t = ovotestis, р = proboscis, ра! = pallial
gland, sd = salivary gland duct, sgl = salivary
gland, sr = seminal receptacle, s = stylet, su =
sucker, Sv = seminal vesicle).

the supraesophageal ganglion (Fretter & Gra-
ham, 1949; Haszprunar, 1988a, 1990). In all
of the taxa examined in this study, the esoph-
ageal ganglia are asymmetrical, with the
subesophageal ganglion being oblong and
the supraesophageal spheroid (Fig. 6A); (4)
the osphradial nerve, as it extends to the
osphradium from the supraesophageal gan-
glion, passes beside and anterior to the pro-
boscis and not posterior to the proboscis
(Fretter & Graham, 1949; Haszprunar 1988a)
(Fig. 6A); and (5) the rhinophoral and lateral
nerves, described by Huber (1987, 1993) for
various pyramidellids, and used by Haszpru-
nar (1988a, 1990) to hypothesize that the
Pyramidellidae are sister taxa to the opistho-
branchs, are not homologous with those
nerves of the same name in the opistho-
branchs described by Huber (1987, 1993). In
some opisthobranchs (e.g., Architectibran-

FIG. 6. A. Diagram of generalized pyramidellid ner-
vous system. Buccal ganglia not illustrated; B. Di-
agram of the nervous system of Pyramidella sp.
showing the nerve that splits from the main os-
phradial nerve to innervate that portion of osphra-
dium located on the left side of the mantle (bar = 1
mm) (cg = cerebral ganglia, п = nerve, о = osphra-
dium, од = osphradial ganglion, on = osphradial
nerve, pdg = pedal ganglia, plg = pleural ganglia,
sbg = subesophageal ganglion, spg = supraesoph-
ageal ganglion, vg = visceral ganglion).

chia), the rhinophoral nerves innervate the
posterior portion of the Hancock’s organ,
whereas in others (e.g., Aplysiomorpha and
Bullomorpha) these nerves innervate the
rhinophores. These structures are not con-
sidered to be homologous (pers. comm.,
Gosliner, 1992). Moreover, all known pyra-
midellids do not possess either a Hancock’s
organ or rhinophores (the nerves Huber
[1987] identified as rhinophoral in the pyra-
midellids he examined, innervate the lateral
walls of the head—an area he suggested
probably contains sensory cells). Huber
(1987) stated that the lateral nerves, which
originate on the pedal ganglia near the
pleuro-pedal connective, occur in pyramidel-

454 WISE

lids, opisthobranchs, and pulmonates. In the
Architectibranchia, these nerves innervate
the lateral body walls, as well as parts of the
columella muscle, whereas in Aplysiomorpha
and Bullomorpha, Huber only listed them as
present. In primitive pulmonates (e.g., Ar-
chaepulmonata), the lateral nerves innervate
portions of the pharynx, while in pyramidel-
lids, he determined they innervate the mus-
cles of the lateral body wall. As with the rhi-
nophoral nerves, it is not possible to consider
the lateral nerve homologous across the taxa
he examined. Moreover, examination of just
the opisthobranchs (illustrated by Huber,
1987) revealed that the rhinophoral and lat-
eral nerves are interchangeable, and there-
fore fail the positional test of homology as
defined by Remané (1955).

Life history: The literature contains a lim-
ited amount of life-history data (Lebour,
1932; Thorson, 1946, 1950; Ramussen,
1944, 1951; Amio, 1963; Robertson, 1967,
White, 1985). Pyramidellids for which longev-
ity is known have a life span of one year (Ras-
mussen, 1944; Wells, 1959; Nishino et al.
1983; White et al., 1985; McFadden & Myers,
1989; pers. obser.). The spawning season for
snails living in temperate to subtropical re-
gions is generally 3-4 months. Cumming
(1988, 1993) discusses the spawning behav-
¡or of a single tropical species in a mariculture
setting. Pyramidellids deposit their eggs in a
continuous string called a chalazae. The cha-
lazae is composed of eggs joined end to end
by threads that are continuous with the wall
of the cocoon. These strings are molded into
a gelatinous egg mass.

Character Descriptions

SHELL CHARACTERS

(1) Protoconch angle

0—120-125° (Figs. 9C, 19F)
1—90-95° (Fig. 25C)
2—130-135° (Fig. 14C)
3—140-145° (Fig. 15D)

Remarks: The protoconch is the larval shell
formed prior to metamorphosis into the juve-
nile stage. The protoconch angle is the angle
at which the protoconch axis is oriented to the
axis of the adult shell.

(2) Orientation of protoconch

O—sinistrally heterostrophic (Figs. 9C,
25С, 29D, 8 24H)

1—dextrally heterostrophic (Figs. 7C, 14C,
15D, & 18D)

Remarks: In state 0, the protoconch generally
lies across the teleoconch, with its earliest
portion partially submerged (Fig. 9C) or com-
pletely exposed (Fig. 25C). In state 1, the ear-
liest portion of the protoconch is completely
submerged within the teleoconch and coils
upwards. The terms sinistrally and dextrally
heterostrophic are adopted from Fretter et al.
1986 (p. 557, fig. 377) and are used here to
described the relationship of the protoconch
to the teleoconch.

(3) Number of protoconch whorls

0—2.0 (Fig. 23E)
1—2.5-3.0 (Fig. 28C)
2—1.0-1.5 (Fig. 18D)

Remarks: The number of protoconch whorls
were determined using the method outlined
by Robertson (1976).

(4) Columellar fold

O—absent—No apparent
fold(s) at aperture (Fig. 29B)

1—a single, prominent and acute fold on
the upper one-half of the columella
perpendicular to the columella axis
(Fig. 9B)

2— а single, less acute fold, which begins
basally on the columella to extend, at
an oblique angle to the upper half of
the columella (Fig. 15C)

3—one large fold on the upper half of the
columella dorsal to two smaller folds,
all three perpendicular to the col-
umella (Fig. 19C)

columella

Remarks: The columella is the solid or hollow
pillar formed by the adaxial walls of the whorls
and surrounds the axis of the coiled shell
(Knight et al., 1960). The columellar fold is a
spirally wound ridge on the columella that 1$
readily seen at the shell's aperture. Slight
swellings on the columella deep within the
body whorl and, therefore, not visible at the
aperture were not considered to be homolo-
gous to columellar folds in this study (e.g., as
in Tathrella iredalei (Fig. 29F).

In the outgroup, Amathinidae, genera with
limpet-like shells (e.g., Amathina) lack a col-
umella. Other genera (e.g., Clathrella) have a
littoriniform shell with a columella. Only the
latter genera were used to polarize charac-
ters of the columella.

PYRAMIDELLID MORPHOLOGY AND PHYLOGENY 455

(5) Columellar fold ridges

O—columellar smooth or unridged (ЭВ,
15C)

1—a single ridge with non-overlapping or
imbricate plates (23D)

2—2-4 ridges of many imbricate plates

(19D&E).
(6) Palatal teeth
O—absent

1—3 to 4 (Fig. 23C)
2—6 to 8 (Fig. 19C)

Remarks: Fully formed palatal teeth, or folds,
are located deep inside the outer lip of the
shell’s aperture. Because the palatals located
immediately inside the aperture are undevel-
oped and depend on the age of the snail, they
were not used as characters.

Operculum Character

(7) Operculum notched to accommodate
columellar fold.

O—absent (Figs. 13F, 28F)
1—present (Fig. 191)

Remarks: All genera examined herein pos-
sess an operculum composed of a brown,
hardened, and proteinaceous substance.
Only taxa of the subfamily Pyramidellinae
have an operculum that is notched to accom-
modate the largest columellar fold.

External Anatomy Characters

(8) Tentacles

O—comnate (Figs. 21A-C, 8A-D)
1—not соппае (Figs. 21D, 17А, В)

Remarks: Connate tentacles are joined along
their anterior base.

(9) Tentacle shape

O—triangular and laterally folded (Fig.
21А)

1—subtriangular and ventro-laterally
folded (Figs. 8A, D, 21B, C)

2—cylindrical (Figs. 21D, 17A, B)

(10) Tentacular pads

O—absent
1—present

Remarks: Tentacular pads are composed of
fused cilia that are located subterminally, at
the apex of the tentacles. These pads are only

found in members of the subfamily Odostom-
inae.

(11) Attachment thread

O—absent
1—present.

Remarks: Attachment threads may be pro-
duced by the pedal gland to secure the snail
to the substratum or its host. Pedal threads
are absent in taxa that are infaunal.

(12) Bifurcate mentum

O—present (Fig. 11D-I)
1—absent (Fig. 11А-С)

Remarks: The mentum is a shelf-like projec-
tion immediately dorsal to the propodium. The
mentum, in all but one of genera examined in
this study, extends beyond the foot to the
substratum when the snail is moving. Its
shape is highly variable among examined
genera.

(13) Anterior mentum edge
O— incised (Fig. 11D, F)
1—retuse (Fig. 11E, С)
2—emarginate (Fig. 11H, |)
3—unnotched (Fig. 11А-С)

Alimentary Tract Characters

(14) Introvert-proboscis aperture

O—on the dorsal surface of the mentum
base

1—medial on the mentum tip

2—ventral side of the head dorsal to the
mentum

Remarks: This aperture is the opening
through which the acrembolic (= completely
invaginable) proboscis passes upon protrac-
tion or retraction.

(15) Four-way junction

O—absent (Figs. 8A-D, 17B)
1—present (Figs. 21A-D, 17A)

Remarks: The four-way junction is formed by
the convergence of the anterior esophagus,
posterior esophagus, and paired salivary
glands.

(16) Buccal pump

O—buccal pump without blind sac or cae-
cum (Fig. 17B)

1—with blind sac (Figs. 8A-D, 17A,
21А-О)

456 WISE

Remarks: The buccal pump 1$ that portion of
the gut between the buccal sac and esoph-
agus.

(17) Distal portion of buccal pump (= bp2)

0—laterally flattened (8A, 17A, 21A)
1—circular (8C)

Remarks: Descriptions refer to shape of distal
end of buccal pump when viewed in cross-
section.

(18) Buccal pump

0—outside the proboscis sheath
1—within the proboscis sheath

Remarks: When the proboscis is retracted
the buccal pump may lie within or outside of
the proboscis sheath.

(19) Salivary gland ducts

0—not within or attached to buccal pump
or esophagus (Fig. 17В)

1—attached to the exterior of anterior
esophagus (Figs. 17A, 21A,C, D)

2—within the walls of anterior esophagus
(Fig. 21B)

3— within the walls of buccal pump (Fig.
8A-D)

Remarks: The ducts of the salivary glands ex-
tend anteriorly to penetrate the stylet bulb,
where they unite to form a single duct entering
the stylet.

(20) Salivary gland terminus

O—attachment absent (Fig. 8A-D)
1—attachment present (Figs.
21A-D)

17A-B,

Remarks: The salivary glands are elongate,
slender organs composed of few to numerous
cells bordering a central ciliated lumen. The
posterior end of each gland terminates in a
vesicle-like bladder that can be attached to
the posterior esophagus by a fine thread of
muscle or connective tissue.

(21) Buccal stylet
O—absent
1—present

Remarks: The stylet, purportedly made of
chitin, is enclosed within a sheath.

Pallial Cavity Characters

(22) Pigmented mantle organ

O—absent
1—small, oblong and anterior of the kid-
ney (Fig. 1A)

2—large, rectangular and oblong mass
comprised of pigmented cells sur-
rounded by transparent cells just an-
terior of the kidney (Fig. 1C)

3—very large, wide anterior with attenu-
ated posterior that extends to the
juncture of the dorsal and ventral cili-
ated strips (Fig. 1B)

Remarks: The pigmented mantle organ which
hangs from the mantle roof just to the right of
center, produces an opaque to colorful, vis-
cous substance that 1$ released when the
snail is disturbed (Table 5).

(23) Secondary gill

O—absent
1—present

Remarks: When present, the gill is located
between the opposing ciliated strips (Fig. 4B).
The outgroup has a “gill” to the right of the
dorsal ciliated strip and is not homologous to
the pyramidellid gill. It is in a different position
in the mantle cavity and is comprised of thin
sheets or leaves, in contrast to fairly thick
ridges as in the pyramidellids. Moreover, nei-
ther is homologous with the gastropod
ctenidium, on the basis of position, blood
supply, and structure.

(24) Osphradium

O—subtriangular, left side only

1—subtriangular on the left side and ex-
tends across mantle to right mantle
corner, just inside mantle edge

(25) The gland beneath the ventral ciliated
strip

O—underlies entire ventral ciliated strip
(Fig. 1B)

1—underlies 50-60% of the strip (begin-
ning anteriorly and extending posteri-
orly)

2—underlies only 20-25% of the anterior
portion of strip (beginning anteriorly
and extending posteriorly) (Fig. 1A)

(26) Exudate

O—secreted from the ventral ciliated strip
gland

1—secreted from the pigmented mantle
organ

PYRAMIDELLID MORPHOLOGY AND PHYLOGENY 457

2—secreted from both the ventral ciliated
strip gland and the pigmented mantle
organ

Remarks: A viscid exudate is secreted when
the snail is disturbed (Table 5). The exudate 15
believed to be repugnant to repel potential
predators (Robertson, 1985; pers. obser.)

Penial Characters
(27) Penis position
O—outside the nerve ring (Figs. 22A, C,
12A-C)
1—а portion of the penis lies within the
nerve ring (Figs. 22B, D)

(28) Penial sperm sac

O—absent
1—present (Figs. 22B, D)

Phylogenetic Analysis

Classical evolutionary taxonomic schemes
use conchological characters to subdivide the
Pyramidellidae into four subfamilies (Abbott,
1974; Boss, 1982). This study is the first to use
cladistic (= phylogenetic) methodology to test
the validity of three of the subfamilies and to
construct hypotheses about the relationships
of the species. The following results and dis-
cussion are presented in two parts. The first
focuses on the relationships of the taxa as
indicted by the cladograms, and briefly, the
taxonomic implications of these relationships.
The second examines conflicting hypotheses
of relationships when soft-part anatomy and
protoconch characters are considered sepa-
rately. To aid in the discussion of the cla-
dograms, clades are labelled as units I-IV
(Figs. 30-35).

Relationships within the Pyramidellidae

Six equally parsimonious cladograms (Figs.
30-35) are produced using the data set of 28
Characters and 13 taxa (Table 3). All trees
have a length of 67 steps, a consistency index
(Cl) of 68, and a retention index (RI) of 81. Final
transformation series for multistate charac-
ters are listed in Table 4.

A clade uniting Odostomia babylonia, O.
didyma, Boonea seminuda, В. cincta is found
(unit | in Figures 30-35) in all trees. Boonea
cincta has previously been placed in the ge-
nus Odostomia, subgenus Chrysallida. This
clade corresponds to the subfamily Odos-
tominae. In traditional classifications (Dall &
Bartsch, 1904; 1909), taxa were assigned to
the Odostominae if their shells were short,

subconic or ovate in shape, unsculptured or
cancellate, possessed few whorls, and had a
single columellar fold. Here | propose that the
Odostominae (represented by the taxa in unit
I; Figs. 30-35) are monophyletic and that their
definition be expanded by seven synapomor-
phies: columellar fold (state 4-1), tentacular
pads (state 10-1), mentum not bifurcated
(state 12-1), shape of anterior mentum edge
(state 13-3), introvert aperture location (state
14-2), position of salivary gland ducts within
the alimentary tract (state 19-3), and the size
of the gland beneath ventral ciliated strip
(state 25-2). Also defining this clade are two
convergent character states: tentacle shape
(state 9-1) and pedal thread (state 11-1).

Within unit I, relationships among O. baby-
lonia, В. seminuda, and the O. didyma/B.
cincta sister group, while resolved in Figures
30, 32, and 34, are unresolved in Figures 31,
33, and 35. In all cladograms (Figs. 30-35),
Boonea cincta and Odostomia didyma are
shown to be more closely related to each
other than either is to B. seminuda and O.
babylonia. In these trees then, the genus
Odostomia is not monophyletic. This appar-
ent paraphyly is dependent upon protoconch
character states 1-3 that group В. cincta and
O. didyma. This sister taxa grouping is con-
trary to characters of soft-part anatomy,
which indicate that O. babylonia and O.
didyma are most closely related. Moreover,
on the basis of soft-part anatomy (particularly
due to similarities of the alimentary tract) Boo-
nea cincta and B. seminuda are sister taxa.
Consequently, Chrysallida cincta Carpenter is
transferred to the genus Boonea.

In all cladograms (Figs. 30-35), units И-М
(sister taxa to the Odostominae [unit 1]) form a
monophyletic group on the basis of five char-
acter states. These synapomorphies include:
introvert-proboscis aperture medial on men-
tum tip (state 14-1), portions of the alimentary
tract forming a four-way junction (state 15-1),
buccal pump within the proboscis sheath
(state 18-1), salivary gland ducts attached to
the exterior of the anterior esophagus (state
19-1), and the salivary gland's terminal end
attached to the posterior esophagus by a fine
thread (state 20-1).

Unit Il, composed of Sayella hemphillii
and Petitella crosseana (formerly placed in
Sayella) are united as sister taxa in all cla-
dograms (Figs. 30-35) because they share
two synapomorphies: protoconch angle
(state 1-3) and columellar fold configuration
(state 4-2) and four convergences: dextral hy-
perstrophy (state 2-1), number of protoconch

458 WISE

whorls (state 3-2), tentacles not connate
(state 8-1) and tentacle shape (state 9-2).
These characters are, however, in conflict
with characters (15-19) of the alimentary
tract. Although grossly their respective head
and foot are similar, these two species are
anatomically very different. Furthermore,
when protoconch character (state 3-1) is
eliminated from the phylogenetic analysis,
these taxa are no longer united and their close
kinship no longer supported (Fig. 36).

This study shows that Sayella hemphillii
and Petitella crosseana, both originally as-
signed to the same genus, and to the Odos-
tominae solely on the basis of overall shell
shape and the number of columellar folds, do
not belong in the same genus nor should ei-
ther be considered any longer a member of
this subfamily. Sayella hemphillii is anatomi-
cally very different from all described odos-
tomian species, and the anatomy of P. cros-
seana is unlike any described pyramidellid.
Consequently, Sayella crosseana 1$ assigned
to the new genus Petitella, and both taxa are
placed in the new subfamily Sayellinae be-
cause they are distinct from our current con-
structs of existing pyramidellid subfamilies as
shown here.

The clade composed of Pyramidella sul-
cata, P. crenulata, and P. mitralis (unit Ill; fig-
ures 30-35) is equal to the subfamily Pyra-
midellinae. Originally, pyramidellids were
allotted to this subfamily if their shell shape
was elongate-conic, the shell surface was
polished, the adult whorls were flat-sided and
the columella had 1-3 columellar folds (Ра! €
Bartsch, 1904, 1909). As stated above, shell
characters of this nature may provide con-
fusing and poorly defined guidelines for as-
signing taxa to phylogenetically meaningful
groups. Here the monophyly of the Pyramidel-
linae is proposed on the basis of five synapo-
morphies: presence of three columnar folds
(state 4-3), notched operculum (state 7-1),
size and shape of pigmented mantle organ
(state 22-3), presence of a secondary gill
(state 23-1), and the configuration of the os-
phradium (state 24-1) (Figs. 30-35). Within
unit Ш, Pyramidella mitralis and P. sulcata are
sister taxa relative to P. crenulata, because
they share the same number of columellar fold
ridges (state 5-2) and shell apertural palatal
teeth (state 6-2). Paradoxically, the members
of this clade cannot be distinguished from one
another on the basis of the soft-part anatomy
examined in this study. Pyramidella sulcata
can only be separated from P. mitralis by over-

all shell shape (Figs. 14, 19) and minute per-
forations of the shell (Fig. 19A). Shell charac-
ters also separate Pyramidella crenulata from
P. sulcata; the former is much smaller, and its
sutures are crenulated (Fig. 18B). These shell
characters, because they are autapomor-
phies, were not included in the analysis.

Taxa of the subfamily Pyramidellinae (unit
ill), as discussed above, are only separated
into species on the basis of shell characters.
Anatomically these taxa are nearly identical.

Turbonilla hemphilli, Houbricka incisa (for-
merly Turbonilla incisa), and Tathrella iredalei
are united in a clade (unit IV; Figures 30-35)
that, at least in part, corresponds to the sub-
family Turbonillinae. Pyramidellids were as-
signed to the subfamily Turbonillinae if their
shell shape was lanceolate, their adult whorls
were numerous and the whorls had pro-
nounced axial ribs (Dall & Bartsch, 1904,
1909). Here the monophyly of the Turbonilli-
nae is proposed and supported by four syn-
apomorphies: incised anterior mentum edge
(state 13-1), exudate origin (state 26-2), po-
sition of the penis through the nerve ring (state
27-1), and presence of a sperm sac (state
28-1).

In all trees (unit IV) (Figs. 30-35), Houbricka
incisa is separated from the Turbonilla hemp-
hilli/Tathrella iredalei sister group on the basis
of several anatomical differences: anterior
mentum edge (state 13-0), pigmented mantle
organ shape (state 22-1), origin of repugna-
tory exudate (state 26-0), and penial complex
(states 27-0 and 28-0). Therefore, the genus
Turbonilla is paraphyletic in all trees (Figs. 30-
35). Prior to this study, the anatomy of both
species was unknown, and both were in-
cluded in the same genus because of their
lanceolate shell and protoconch configura-
tion. Houbricka incisa is anatomically very dif-
ferent from Turbonilla hemphilli or any of the
other taxa currently assigned to this genus.
Therefore, the new genus Houbricka, with
Turbonilla incisa Bush as the type species, is
proposed herein. Tathrella iredalei is provi-
sionally retained in the subfamily Turbonillinae
because it possesses synapomorphies that
seem to justify this placement.

Protoconch Characters vs.
Anatomical Characters

Members of the Pyramidellidae are gener-
ally only known from their shells (Abbott,
1974; Fretter et al. 1986, Haszprunar, 1988a).
Moreover, the lack of understanding of this
family’s biology and anatomy is often attrib-

PYRAMIDELLID MORPHOLOGY AND PHYLOGENY 459

uted to the difficulty associated with studying
such small snails (Haszprunar, 1988b). Con-
sequently, with a better understanding of their
anatomy, it is not surprising that oversplit tax-
onomic groupings based on shell characters
alone may be in conflict (= incongruent) with
anatomical characters. When the protoconch
characters 1-3 are suppressed (Hennig86
option cc];), 20 equally parsimonious cla-
dograms are produced. After successive
weighting (Hennig86 option xs w:), one tree
remains with a length of 57 steps, a Cl of 81,
and a RI of 90. In this cladogram both O.
didyma/O. babylonia and В. seminuda/B.
cincta are sister taxa (Fig. 36). Contrary to the
relationships suggested by the protoconch
characters, feeding structures suggest that O.
babylonia and O. didyma are most closely re-
lated (= sister taxa). They essentially share the
same gut configuration, the only difference 15
that the salivary gland ducts in O. babylonia
exit the gut just prior to the stylet bulb (Fig.
8C). Boonea seminuda and B. cincta share the
same gut configuration and differ only in the
length of buccal pump one (bp1). The bp1 of
В. cincta 1$ equal to the length of the buccal
pump two (bp2), whereas in B. seminuda the
bp1 is 1.5 times the bp2 (Fig. 8B).

These conflicting hypotheses of relation-
ships may be explained by the fact that pro-
toconch shape 1$ a reflection of developmen-
tal mode and not phylogeny. Historically,
protoconch configuration has been correlated
with the type of larval development rather than
phylogeny (Reid, 1989). For constituents of
the Odostominae in the present study, our
knowledge of larval development is limited to
B. cincta and B. seminuda. Boonea cincta
undergoes direct development (i.e., non-
planktonic) and recently metamorphosed in-
dividuals crawl away from the egg mass within
25 to 28 days (LaFollette 1979). Boonea sem-
inuda is lecithotrophic and in the plankton for
approximately two weeks (Robertson, 1978).
Species with non-planktotrophic develop-
ment and no planktonic stage typically may
have a large smooth protoconch (i.e., highly
inflated) of few whorls and generally are not
distinguishable from the adult shell. Boonea
cincta meets two of the three criteria; how-
ever, its smooth protoconch is easily delim-
ited from the highly sculptured adult shell (Fig.
7C). Generally, snails that are lecithotrophic
have a protoconch of a few whorls of inter-
mediate size (i.e., less inflated). This condition
is present in В. seminuda (Fig. 9C-E). There-
fore, developmental modes are different in

these two species, as are their respective pro-
toconch shapes. The protoconch characters
are incongruent with the other characters and
rejected as synapomorphies. What of O.
babylonia and O. didyma? Their protoconch
shapes are very different. Is this, too, a re-
flection of developmental mode and not phy-
logeny? The only way to determine this is to
discover their developmental modes, which
when treated as characters, can be mapped
onto a cladogram of the working phylogeny
for the group and checked for congruence
with other characters.

What of convergences in gut or feeding-
structure anatomy as a result of feeding on
similar hosts? The hosts for both B. seminuda
and B. cincta are known. Boonea seminuda
parasitizes a number of hosts across its range
(e.g., Crepidula fornicata and Aequipecten
gibbus), whereas В. cincta feeds on a number
of trochiid gastropods (e.g., Norrisia norrisi)
and Haliotis spp. These hosts are all different,
indicating that feeding structure anatomy is a
reflection of phylogeny and not adaptation.
The hosts for O. babylonia and O. didyma are
not known.

Protoconch and adult shell characters (e.g.,
similar columellar fold configurations) also
united S. hemphillii and Petitella crosseana as
sister taxa in the initial cladograms, whereas
anatomical characters separate the two (Fig.
36). Although they both have a grossly similar
head-foot (e.g., tentacles not connate, tenta-
cles cylindrical, similar shaped and emargin-
ate mentum, and location of introvert open-
ing—none of these are unique to the two),
they are very dissimilar anatomically. They
have very different alimentary tract (Fig. 17A,
B) and penial configurations (Fig. 12A, B). The
paraphyly of the genus Turbonilla, as depicted
in Figures 30-35, is even more apparent when
the protoconch characters were eliminated
from the phylogenetic analysis (Fig. 36). How-
ever, until both developmental modes and
hosts are known for these species, it is im-
possible to eliminate the protoconch charac-
ters as completely phylogenetically uninfor-
mative and/or choose between competing
hypotheses of relatedness (i.e., between phy-
logeny and convergent adaptation).

Historically, shell characters, to the exclu-
sion of soft-part anatomy, have been used to
assign taxa to the various pyramidellid gen-
era. Consequently, it might be tempting to rely
more on anatomical characters and treat con-
chological characters a priori as homoplastic
and uninformative. However, as stated above,

460 WISE

| was only able to distinguish members of the idence, sensu Jones et al., 1993) should be
subfamily Pyramidellinae on the basis of shell used in a phylogenetic analysis. In addition, it
morphology, whereas in other taxa, soft-part is only by testing these characters for con-
anatomy proved most informative in deter- gruence with other characters, that the often
mining relatedness. Therefore, the present complex relationships between taxa can be
study indicates that all characters (= total ev- resolved.

FIG. 7. Shell morphology of Boonea seminuda. SEM microphotographs: A. Apertural and dorsal views of
shell (bar = 1 mm); B. Aperture and columellar fold (bar = 400 um); C. Tilted, frontal view of protoconch (bar
= 100 um); D. Lateral view of protoconch (bar = 100 um); E. Tilted, lateral view of protoconch (bar = 100 um);
Е. Unattached surface of operculum; G. Attached surface of operculum (bar = 200 um).

PYRAMIDELLID MORPHOLOGY AND PHYLOGENY 461

р
| fl
B Man
ps [A ~ S \ bp2
KANN m
N u PA \ у и
PE dos i \ sd ( \ SS
A AN Ss \ A ù
e JA
Y
O
D GAS
ppl VA bp2
== i «| LAO ale
: = oe i LE 8 es
Le A) a sd с
ря = sb N

FIG. 8. Diagram of pyramidellid alimentary tracts. A. Boonea стс (bar = 200 um); В. Boonea seminuda
(bar = 500 um); С. Odostomia babylonia (bar = 150 um); D. O. didyma (bar = 150 um). (bp1 = buccal pump
1, bp2 = buccal pump 2, bs = buccal sac, es = esophagus, p = proboscis, sb = stylet bulb, sd = salivary
gland duct, sgl = salivary gland, su = sucker).

462 WISE

FIG. 9. Shell morphology of Boonea cincta. SEM microphotographs: A. Apertural and dorsal views of shell
(bar = 1 um); B. Frontal view of protoconch (bar = 100 um); C. Lateral view of protoconch and first two adult
whorls (note smooth protoconch vs. cross-hatched teleoconch) (bar = 200 um); D. Apical view of proto-
conch (bar = 100 um); E. Lateral view of protoconch showing demarkation of larval and adult shells (bar =
150 um); F. Unattached surface of operculum; G. Attached surface of operculum (bar = 200 um).

PYRAMIDELLID MORPHOLOGY AND PHYLOGENY 463

FIG. 10. Living snails of the subfamily Odostominae. A. Boonea seminuda (bar = 650 um); В. Одозютиа
babylonia (bar = 200 um); C. Odostomia didyma (bar = 150 um).

464 WISE

FIG. 11. A. Unnotched mentum of Boonea seminuda and Boonea cincta; B. Unnotched mentum of Odos-
tomia babylonia; С. Unnotched mentum of Odostomia didyma; D. Retuse mentum of Pyramidella mitralis,
P. sulcata, and P. crenulata; E. Incised mentum of Turbonilla hemphilli; F. Retuse mentum of Houbricka
incisa; G. Incised mentum of Tathrella iredalei; H. Emarginate mentum of Sayella hemphillii; |. Emarginate
mentum of Petitella crosseana.

PYRAMIDELLID MORPHOLOGY AND PHYLOGENY 465

pe

FIG. 12. Diagram of retracted penis configuration and position in: A. Boonea seminuda (bar = 250 um); B.
Sayella hemphilli (bar = 300 um); С. Petitella crosseana (bar = 200 um) (f = foot, me = mentum, nr = nerve
ring, op = operculum, р = proboscis, pe = penis).

466 WISE

FIG. 13. Shell morphology of Odostomia babylonia. SEM microphotographs: A. Apertural and dorsal views
of shell (bar = 600 um); B. Enlargement of shell's shoulder and suture (bar = 40 um); C. Apical view of
protoconch (bar = 100 um); D. Lateral view of protoconch showing partially exposed earliest portion (bar
= 100 um); E. Lateral view of protoconch tilted to reveal demarkation of larval and adult shells (bar = 40 um);
F. Unattached surface of operculum; G. Attached surface of operculum (bar = 150 um).

PYRAMIDELLID MORPHOLOGY AND PHYLOGENY 467

FIG. 14. Shell morphology of Odostomia didyma. SEM microphotographs: A. Apertural and dorsal views of
shell (bar = 500 um); B. Protoconch and oldest adult whorl (bar = 100 um); C. Lateral view of protoconch
(bar = 100 um); D. Apical view of protoconch (bar = 100 um); E. Frontal view of protoconch and first adult
whorl (note deep shoulder) (bar = 100 um); F. Unattached surface of operculum; G. Attached surface of
operculum (bar = 150 um).

468 WISE

FIG. 15. Shell morphology of Sayella hemphillii. SEM microphotographs: А. Apertural and dorsal views (bar
= 1 mm); B. Young specimen (note difference between this shell and shell in figure A.) (bar = 1 mm); C. Body
whorl cracked open to reveal columellar fold (bar = 400 um); D. Apical view of protoconch (bar = 100 um);
E. Frontal view of protoconch (bar = 100 um); F. Lateral view of protoconch (bar = 100 um); G. Attached
surface of operculum; H. Unattached surface of operculum (bar = 400 um).

PYRAMIDELLID MORPHOLOGY AND PHYLOGENY 469

FIG. 16. A. Sayella hemphillii (bar = 1 mm); В. Pe-
titella crosseana (bar = 500 um).

A p bp] |
/ |
} bs | —sd
Ji |
® E
> sb,
ee ==
(0% pes
KY, pe
\ —
sol
B
p
— bs bp
_2 0 © ' |
e =="
| = A xs 64
50 $
—es
sel

FIG. 17. A. Diagram of the alimentary tract of
Sayella hemphillii (bar = 1 mm); В. Diagram of ali-
mentary tract of Petitella crosseana (bar = 500 um)
(aes = anterior esophagus, bp = buccal pump, bp1
= buccal pump 1, bp2 = buccal pump 2, bs = buc-
cal зас, es = esophagus, р = proboscis, рез =
posterior esophagus, sb = stylet bulb, sd = salivary
gland duct, sgl = salivary gland).

470 WISE

FIG. 18. Shell morphology of Petitella crosseana. SEM microphotographs: A. Apertural and dorsal views
(bar = 1 mm); В. Enlargement of the shell’s shoulder and suture (bar = 20 um); С. Lateral view of protoconch
and earliest adult whorls (bar = 100 um); D. Frontal view of protoconch (bar = 100 um); E. Apical view of
protoconch (bar = 100 um); F. Unattached surface of operculum; G. Attached surface of operculum (bar =
200 um).

PYRAMIDELLID MORPHOLOGY AND PHYLOGENY 471

FIG. 19. Shell morphology of Pyramidella sulcata. A. Apertural and dorsal view of shell (bar = 4 mm); SEM
microphotographs: B. Suture and surface microstructure (note growth lines) (bar = 600 um); C. Shell
aperture with columellar folds and mature, well-developed palatal teeth (bar = 2 mm); D. Ridges of largest
columellar fold (bar = 200 um); E. Columellar fold ridges, composed of imbricated plates (bar = 10 um);
F. Lateral view of partially eroded protoconch (bar = 200 um); G. Apical view of protoconch (bar = 200 um);
|. Unattached surface of operculum (bar = 1 mm).

472 WISE

FIG. 20. Living snails of the subfamily Pyramidellinae. A. Pyramidella sulcata (bar = 3 mm); B. Pyramidella
crenulata (bar = 2 mm) C. Pyramidella mitralis (bar = 3 mm).

PYRAMIDELLID MORPHOLOGY AND PHYLOGENY 473

B
р (a aes
= | EN bp2 N La
Кб EN = 7 — |
A AVENA N
A UL bs UN NS
Е sb sd я So
|
== р
ALA
D

FIG. 21. Diagram pyramidellid alimentary tracts. A. Pyramidella mitralis, P. sulcata, P. crenulata (bar = 1
mm); В. Turbonilla hemphilli (bar = 500 um); С. Houbricka incisa (bar = 500 um); D. Tathrella iredalei (bar =
300 um) (aes = anterior esophagus, bp1 = buccal pump 1, bp2 = buccal pump 2, bs = buccal sac, p =
proboscis, pes = posterior esophagus, sb = stylet bulb, sd = salivary gland duct, sgl = salivary gland).

474 WISE

m
\\/ op—\\ 7
vu Y
U у

FIG. 22. Diagram of pyramidellid retracted penes in situ. А. Pyramidella sulcata, Р. crenulata, and P. mitralis
(bar = 1 mm); В. Turbonilla hemphilli (bar = 500 um); С. Houbricka incisa (bar = 400 um); D. Tathrella iredalei
(bar = 300 um) (f = foot, me = mentum, nr = nerve ring, ор = орегсшит, р = proboscis, ре = penis, rmu =
retractor muscle, ss = sperm sac).

PYRAMIDELLID MORPHOLOGY AND PHYLOGENY 475

FIG. 23. Shell morphology of Pyramidella crenulata. A. Apertural and dorsal views of shell (bar = 3 mm); SEM
microphotographs: В. Sutures and exterior surface of shell (bar = 500 um); С. Portion of broken body whorl
revealing mature palatal teeth and columellar folds (bar = 400 um); D. Single ridge of non-overlapping and
overlapping plates on largest columellar fold (bar = 40 um); E. Lateral view of protoconch (bar = 100 um);
F. Dorsal view of protoconch (bar = 125 um); G. Unattached surface of operculum; H. Attached surface of
operculum (bar = 430 um).

476

ЕС. 24. Shell morphology of Pyramidella mitralis. A. Apertural and dorsal views of shell (bar = 4 mm); SEM
microphotographs: B. Sutures and surface sculpture of shell (bar = 200 um); C. Shell cut away to expose
columellar folds (bar = 400 um); D. Ridges of largest columellar fold (bar = 100 um); E. Lateral view of
protoconch (bar = 200 um); Е. Apical view of protoconch (bar = 125 um); G. Enlargement of titled, lateral
view of protoconch with a portion of earliest whorl visible (bar = 40 um); H. Attached surface of operculum
(bar = 1 mm).

PYRAMIDELLID MORPHOLOGY AND PHYLOGENY 477

FIG. 25. Shell morphology of Turbonilla hemphilli. A. Apertural and dorsal views of shell (bar = 1.5 mm); SEM
microphotographs: В. Enlargement of whorl immediately dorsal to body whorl (note ribbing) (bar = 125 um);
C. Frontal view of protoconch (bar = 100 um): D. Lateral view of protoconch (bar = 100 um); E. Lateral view
of protoconch showing portion of earliest whorl (bar = 100 um); F. Unattached surface of operculum; G.
Attached surface of operculum (bar = 250 um).

478 WISE

FIG. 26. Living snails of the subfamily Turbonillinae.
A. Turbonilla hemphilli (bar = 1.5 mm); В. Houbricka
incisa (bar = 1 mm); С. Tathrella iredalei (bar = 800
um).

FIG. 27. A. SEM microphotograph of anterior por-
tion of penis of Turbonilla hemphilli (note medial
groove and lateral cuticular hooks) (bar = 75 um);
B. SEM microphotograph of anterior tip of penis of
Houbricka incisa (bar = 10 um).

PYRAMIDELLID MORPHOLOGY AND PHYLOGENY 479

FIG. 28. Shell morphology of Houbricka incisa. SEM microphotographs: A. Apertural and dorsal views (bar
= 1.25 mm); В. Enlargement of the whorl immediately dorsal to body whorl (note costae and intercostal
grooves) (bar = 150 um); C. Lateral view of earliest larval whorls (bar = 100 um); D. Lateral view of
protoconch and earliest adult whorl (bar = 100 um); E. Apical view of protoconch (bar = 100 um); F.
Unattached surface of operculum; G. Attached surface of operculum (bar = 175 um).

480 WISE

FIG. 29. Shell morphology of Tathrella iredalei. A. Apertural and dorsal views (bar = 1 mm); SEM micro-
photographs: B. Body whorl (bar = 1 mm); C. Shell broken to reveal swellings on columella located within
the whorl immediately dorsal to body whorl (bar = 400 um); D. Frontal view of protoconch (bar = 50 um);
E. Apical view of protoconch (bar = 100 um); F. Lateral view of protoconch with early whorl partially
exposed (bar = 100 um); G. Unattached surface of operculum; H. Attached surface of operculum (bar = 400
um).

PYRAMIDELLID MORPHOLOGY AND PHYLOGENY 481

A, EIA, о

out semi judi cinc didy semp cros cren sul mit inc hemp ired

FIG. 30. One of six cladograms resulting from analysis of character matrix in Table 3 (OUT = Amathinidae,
CINC = Chrysallida cincta, ЗЕМ! = Boonea seminuda, JUDI = Odostomia babylonia, DIDY = Odostomia
didyma, SEMP = Sayella hemphillii, CROS = Petitella crosseana, SUL = Pyramidella sulcata, CREN = P.
crenulata, MIT = Pyramidella mitralis, HEMP = Turbonilla hemphilli, INC = Houbricka incisa, IRED = Tathrella
iredalei).

482 WISE

out semi judi cinc didy semp cros cren

sul inc hemp ired
26-1
- 19-0 25-1
- 18-0 11-1
- 16-0 * 9-2
* 17-2 - 15-0 8-1
17-2
16 18 23
13-1
13-2 26-2
1-2 9-2 27-1
15 * 2-17 8-1 28-1
* 3-2 4-2
* 3-2
a 2—1
1-3
17-1
2
13-3
12-1
* 11-1 a
10-1 17
* 9-1
4-1
14-1
15-1
18-1
14 19-1
Sa 20-1
13 16-1
21-1
22-1

FIG. 31. One of six cladograms resulting from analysis of character matrix in Table 3 (OUT = Amathinidae
СМС = Chrysallida cincta, SEMI = Boonea seminuda, JUDI = Odostomia babylonia, DIDY = Odostomia
didyma, SEMP = Sayella hemphillii, CROS = Petitella crosseana, SUL = Pyramidella sulcata, CREN = Р.

crenulata, MIT = Pyramidella mitralis, HEMP = Turbonilla hemphilli, INC = Houbricka incisa, IRED = Tathrella
iredalei).

PYRAMIDELLID MORPHOLOGY AND PHYLOGENY 483

Пе a а FERNER.

out semi judi cinc didy cr semp cros inc hemp ired

* 26-1
- 19-0 25-1
- 18-0 pd eas El
- 16-0 22-2 * 9-2
- 15-0 19-2 * 8-1
* 26-1
13-2
р Fer 22 24
4 no

x.
AO
* 3-

нь
NN
D OS
A
pan
ur

A

NM

pnw

or

N N ND

om

ANT

LU

1-1
a 21 3-1
* 9-1

13 16-1

FIG. 32. One of six cladograms resulting from analysis of character matrix in Table 3 (OUT = Amathinidae,
CINC = Chrysallida cincta, SEMI = Boonea seminuda, JUDI = Odostomia babylonia, DIDY = Odostomia
didyma, SEMP = Sayella hemphillii, CROS = Petitella crosseana, SUL = Pyramidella sulcata, СВЕМ = P.
crenulata, MIT = Pyramidella mitralis, HEMP = Turbonilla hemphilli, INC = Houbricka incisa, IRED = Tathrella

iredalei).

484 WISE

re TV Te

out semi judi cinc didy cren sul mit semp cros inc hemp ired

2261
- 19-0 25-1
gen sit
- 16-0 22-2 222
x 172 = HEC 19-2 Seal
202621
17-2

> ja 1-1
on 2
5-1
17-1 4-3
14-2
13-3
12-1
11-1
10-1 17
* 9-1
4-1
14-1
15-1
18-1
14 19-1
SÓ 20-1
13 16-1
21-1
22

FIG. 33. One of six cladograms resulting from analysis of character matrix in Table 3 (OUT = Amathinidae,
CINC = Chrysallida cincta, SEMI = Boonea seminuda, JUDI = Odostomia babylonia, DIDY = Odostomia
didyma, SEMP = Sayella hemphillii, CROS = Petitella crosseana, SUL = Pyramidella sulcata, CREN = P.

crenulata, MIT = Pyramidella mitralis, HEMP = turbonilla hemphilli, INC = Houbricka incisa, IRED = Tathrella
iredalei).

PYRAMIDELLID MORPHOLOGY AND PHYLOGENY 485

FE. ee en
out

semi judi cinc didy inc hemp ired semp cros cren

sul mit
* 26-1
25-1 - 19-0
* 1 - 18-0
ae 2 * - 16-0
* 471 = .15=0
* 26-1

13-2
He ‚9-2 22 24
8-1
<< 4.2 a
16 1-2 19 aren * 3-2 23 5-2
3 = 2-1 26-2 FDA 6-2
* 3-2 27-1 1-3
15 17-2 28-1
4-3
21 2
3-1 71
1-1 22-3
4-2 23-1
13-3 24-1
12-1
-1
0-1 18
* 9-1
4-1
14-1
15-1
18-1
14 19-1
1 20-1
13 16-1
21-1
22-1

FIG. 34. One of six cladograms resulting from analysis of character matrix in Table 3 (OUT = Amathinidae,
CINC = Chrysallida cincta, SEMI = Boonea seminuda, JUDI = Odostomia babylonia, DIDY = Odostomia
didyma, SEMP = Sayella hemphillii, CROS = Petitella crosseana, SUL = Pyramidella sulcata, CREN = Р.

crenulata, MIT = Pyramidella mitralis, HEMP = Turbonilla hemphilli, INC = Houbricka incisa, IRED Tathrella
iredalei).

486 WISE

Pa TARTA, № NE

out semi judi cinc didy inc hemp ired semp cros cren mit
* 26-1
25-1 - 19-0
STE - 18-0
22-2 * 9-2 - 16-0
* 172 19-2 * 8-1 - 15-0
* 26-1
* 17-2 13-2
16 19 292252] 23
SB * 8-1
4-2
18 13-1 3-2 -2
26-2 2-1 -2
1-2 =
3-2 ae
2 =
> и
31 SA
17-1 1—1 22-3
14-2 23-1
13-3 24-1
=n
11-1
0-1 17:
9-1
4-1
14-1
15-1
18-1
14 19-1
> 20-1
13 16-1
21-1
22-1

FIG. 35. One of six cladograms resulting from analysis of character matrix in Table 3 (OUT = Amathinidae,
CINC = Chrysallida cincta, SEMI = Boonea seminuda, JUDI = Odostomia babylonia, DIDY = Odostomia
didyma, SEMP = Sayella hemphillii, CROS = Petitella crosseana, SUL = Pyramidella sulcata, СВЕМ = P.

crenulata, MIT = Pyramidella mitralis, HEMP = Turbonilla hemphilli, INC = Houbricka incisa, IRED = Tathrella
iredalei).

PYRAMIDELLID MORPHOLOGY AND PHYLOGENY 487

out cinc semi judi didy cros inc cren sul mit semp hemp ired

FIG. 36. Cladogram resulting from analysis of character matrix in Table 3 when protoconch characters 1-3
were eliminated (OUT = Amathinidae, CINC = Chrysallida cincta, SEMI = Boonea seminuda, JUDI = Odos-
tomia babylonia, DIDY = Odostomia didyma, SEMP = Sayella hemphillii, CROS = Petitella crosseana, SUL
= Pyramidella sulcata, CREN = P. crenulata, MIT = Pyramidella mitralis, HEMP = Turbonilla hemphilli, INC
= Houbricka incisa, IRED = Tathrella iredalei).

488 WISE

TAXA DIAGNOSES AND DESCRIPTIONS

Family Pyramidellidae Gray, 1840
Subfamily Odostominae Pelseener, 1928
Genus Boonea Robertson, 1978

Boonea Robertson, 1978:364. Type-species:
Jaminia seminuda C. B. Adams, 1839,
by original designation.

Diagnosis: Shell thick, chalky white, conical,
3-5 mm in length, with 4-5 adult whorls.
Whorls with or without spiral cords, axial ribs
or both. Body whorl 50% of shell length. Um-
bilicus minute or absent. Protoconch smooth,
sinistrally heterostrophic oriented 120-130?
to teleoconch, partially submerged in first
adult whorl. Aperture auriform, with single
acute columellar fold. Operculum tan or
brown, auriform, paucispiral, with subcentric
nucleus. Head-foot white and often lentigi-
nous. Foot narrowing posterior to propodium,
widening and narrowing again posteriorly to a
blunt tip. Posterior pedal gland producing at-
tachment thread. Tentacles subtriangular,
connate, ventro-laterally folded; tentacular
pads present. Eyes subepithelial, on median
side of tentacles. Mentum unnotched, not bi-
furcate. Introvert-proboscis aperture on ven-
tral side of head, dorsal to mentum base. In-
trovert joining buccal sac, which is composed
of sucker, mouth, sheathed stylet with sepa-
rate opening, oral tube, and stylet bulb. Buc-
cal sac joining buccal pump, which 1$ divided
into anterior (bp1) and posterior sections
(bp2). Esophagus originating on ventral sur-
face of bp1-bp2 juncture. Salivary gland
ducts entering gut and extending parallel to
one another within walls of bp1 and entering
stylet bulb without exiting alimentary tract.
Globose penis tapering posteriorly and lo-
cated outside and ventral to nerve ring. Un-
cuticularized spermatophores attached to
species-specific location (e.g., snail's neck or
outside last whorl) prior to transfer to mate.

Remarks: Robertson (1978) erected the ge-
nus Boonea, to which he transferred three
western Atlantic species (Boonea seminuda,
B. impressa, and B. bisuturalis) from the ge-
nus Odostomia. The reasons for removing
them were valid, and were based on differ-
ences (е.д., in protoconch shape, operculum
configuration, penial complex, and in the lo-
cation of the gonoduct aperture) between the
Boonea species and the Odostomia species
described by Fretter 4 Graham (1949).

Boonea seminuda (C. B. Adams, 1839)

Jaminia seminuda С. В. Adams, 1839: 280,
pl. 4 (misnumbered; should be pl. 5), fig.
13; Clench & Turner, 1950: 341, pl. 41,
figs. 5-6 (Lectotype: MCZ 186052; type
locality: Dartmouth Harbor, Massachu-
setts).

Odostomia seminuda (С. В. Adams, 1839);
Gould, 1841: 273; Perry & Schwengel,
1955: 122-123, pl. 23, fig. 164; Andrews,
1971: 129, no figure designation, photo
оп р. 129; Odé & Spears, 1972: 2, fig. 3;
Abbott, 1974: 292, fig. 3487.

Chemnitzia seminuda (С. В. Adams); Stimp-
son, 1851: 112

Odostomia (Chrysallida) willisi Bartsch, 1909:
97, 99, pl. 13, fig. 42.

Odostomia (Chrysallida) seminuda Bartsch,
1909: 97, pl. 13, figs. 45, 48.

Odostomia (Chrysallida) toyatani Henderson
8 Bartsch, 1914: 417-418, pl. 13, fig. 2.

Boonea seminuda (C. B. Adams, 1839); Rob-
ertson, 1978: 364, figs. 3, 10-30.

Description: Shell (Fig. 7): Thick, conical,
white, 6 mm in length, composed of 4-5 adult
whorls. Each whorl with 4 spiral cords parallel
to the whorl suture. Cords of upper whorls
crossed by perpendicular axial ribs to give a
cancellate or latticed appearance. Body whorl
50% of shell length and only upper one-half of
spiral cords crossed by axial ribs (Fig. 7A).
Intersection of ribs and cords delineate a se-
ries of deep, rectangular depressions. Whorl
sutures and grooves between spiral cords
striated. Auricular aperture ovate, with thick,
scalloped outer lip and fluted base. Single,
prominent, acute columellar fold on upper half
of columella perpendicular to the columella
axis (Fig. 7B). Smooth, sinistrally heteros-
trophic protoconch oriented 120° to teleo-
conch axis, submerged 40-45% in first adult
whorl, with earliest portion of protoconch par-
tially exposed (Fig. 7С-Е). Operculum brown,
auricular, paucispiral, with subcentric nu-
cleus, but lacking a notch to accommodate
columellar fold (Fig. 7F, G).

Head-foot (Fig. 10A): Opaque, lentiginous
with scattered white cells (particularly abun-
dant on head posterior to eyes). Anterior
portion of foot with slight medial indenta-
tion and rounded lateral edges. Foot nar-
rowing posterior to propodium, then widen-
ing to gradually taper to a blunt apex. Pedal
gland opening at anterior ends of medial
groove on ventral surface of foot. Attach-

PYRAMIDELLID MORPHOLOGY AND PHYLOGENY 489

ment thread present. Tentacles subtriangu-
lar, connate, ventro-laterally folded; tentacu-
lar pads present. Black eyes beneath epithe-
lium on median side of tentacles. Mentum
unnotched, not bifurcate (Fig. 11A). Digestive
tissue cells of the visceral mass light brown
(with black flecks), yellow to light orange
(with brown flecks), or light grey (with black
flecks). Reproductive organs opaque to
transparent.

Alimentary tract (Fig. 8B): п the retracted
condition, the introvert-proboscis extends
posteriorly from its aperture on the ventral
side of the head, dorsal to the mentum base
to enter the cephalic hemocoel. Introvert join-
ing buccal sac, which is connected to buccal
pump. Buccal pump divided into anterior
(bp1) and posterior sections (bp2), with bp1
one and one-half times longer than bp2; bp1
narrow, round in cross-section, thickened
along the last one-third of its length; bp2
wider, laterally flattened, distally rounded.
Long, coiled esophagus, with irregular sur-
face, originating on the ventral surface of the
alimentary tract at bp1-bp2 juncture, extend-
ing into visceral mass and joining stomach.
Convoluted salivary gland ducts penetrating
alimentary tract immediately anterior to bp1-
bp2 juncture, extending parallel to one an-
other within the walls of bp1, and entering
stylet bulb without exiting gut. Salivary
glands not attached distally to esophagus.

Pallial cavity (Fig. 1A): Mantle and mantle
organs typical for members of Odostominae.
Mantle edge finely crenulate. Ventral and
dorsal ciliated strips joining on mantle roof at
posterior end of mantle cavity. Small, oblong
pigmented mantle organ composed primarily
of cells filled with bright yellow exudate and a
few cells containing clear, brown, and green
contents; exudate released when snail is dis-
turbed (Table 5). Small pink, white, or light
orange ventral ciliated strip gland underlying
20-25% of the ventral ciliated strip. Gill ab-
sent.

Reproductive system: Typical of pyra-
midellids examined (Fig. 5B). Penis in pocket
outside and ventral to nerve ring (Fig. 12A).
Penis with small, rounded anterior end, nar-
rowing, then widening to become large and
bulbous posteriorly. Penis attached anteriorly
and posteriorly to floor of its pocket by mus-
cle fibers. Anterior penial opening extending
posteriorly into a cavity framed by a single
layer of heavily ciliated cuboidal cells. In
some specimens, this cavity filled with a
brown, glandular substance. Penis opening

to outside via a medial aperture beneath
mentum. Producing uncuticularized sper-
matophores attached to right posterior sec-
tion of neck prior to exchange with a mate
(Robertson, 1978).

Nervous system: Typical for known pyra-
midellids (Fig. 6A).

Ecology and Distribution: Boonea seminuda
occurs from Prince Edward Island, Canada,
south to Florida and Texas (Robertson, 1978)
and has several different hosts within its range
(Robertson, 1978; Robertson & Mau-Lasto-
vicka, 1979). This species is known to spawn
from mid-June to early October in Massachu-
setts and early July and August in North Caro-
lina (Robertson, 1978). Egg masses have also
been found in North Carolina coastal waters in
January (Wells & Wells, 1961).

Boonea cincta (Carpenter, 1864)

Chrysallida cincta Carpenter, 1864: 659, no
fig. (Holotype: USNM 15730, type local-
ity: Santa Barbara, California).

Odostomia (Chrysallida) cincta (Carpenter);
Dall & Bartsch, 1909: 152-153, pl. 15: fig.
2, 2a; Abbott, 1974: 293 fig. 3495; LaFol-
lette 1977: 19, 21, 22, figs. 1-4.

Odostomia (Chrysallida) vicola Dall & Bartsch,
1909: 153, pl. 16, fig. 11 (Holotype:
USNM 206899; type locality: San Pedro
Bay, California).

Odostomia (Chrysallida) hipolitensis Dall &
Bartsch, 1909: 155, pl. 16, fig. 8 (Holo-
type: USNM 162770; type locality: Punta
San Hipólito, Baja California Sur).

Odostomia (Chrysallida) риса Dall &
Bartsch, 1909: 160, pl. 16, figs. 10-10а
(Holotype: USNM 1627630; type locality:
San Pedro, California).

Odostomia (Chrysallida) promeces Dall &
Bartsch, 1909: 164, pl. 18, fig. 2 (Holo-
type: USNM 162777; type locality: Bahía
Todos Santos, Baja California).

Odostomia (Chrysallida) pulcherrima Dall 8
Bartsch, 1909: 164, pl. 17, fig. 7 (Holo-
type: USNM 206900; type locality: Ter-
minal Island, California).

Odostomia (Chrysallida) утса Dall 8
Bartsch, 1909: 165, pl. 17, fig. 4 (Holo-
type: USNM 162726; type locality: San
Pedro, California).

Odostomia (Chrysallida) santorium Dall &
Bartsch, 1909: 167, pl. 18, fig. 1 (Holo-
type: USNM 46499; type locality: Punta
San Hipólito, Baja California Sur).

490 WISE

Odostomia (Chrysallida) sapia Dall 8 Bartsch,
1909: 167, pl. 18, figs. 3, 3a (Holotype:
USNM 162775; type locality San Diego,
California).

Odostomia (Chrysallida) deceptrix Dall 8
Bartsch, 1909: 169, pl. 17, fig. 1 (Holo-
type: USNM 206904; type locality: Punta
Abreojos, Baja California Sur).

Odostomia (Chrysallida) contrerasi Baker,
Hanna 4 Strong, 1928: 231, pl. 12, fig. 13
(Holotype: CASIZ 066090; type locality:
“Gulf of California”).

Remarks: | examined the holotypes of B.
cincta and all of its synonyms listed above
and determined that they are all conspecific.
This decision was based on the work of La-
Follette (1977), who showed that the shell
sculpture of C. cincta can be smooth or can-
cellate, as exemplified by the sculpture of
the forms named Odostomia santorium and
O. promeces respectively.

Boonea cincta, formerly Chrysallida cincta,
is anatomically very similar to Boonea sem-
inuda the type species of the genus. | have
not examined Chrysallida torrita, the type
species of Chrysallida; therefore, it 1$ not
possible to determine if Boonea is a synonym
of the older name Chrysallida.

Description: Shell (Fig. 9): Thick, conical,
white, 4 mm in length, composed of 3-4 adult
whorls. Upper whorls with 4 spiral cords par-
allel to suture. Upper 3 cords crossed by per-
pendicular axial ribs to give a cancellate or
latticed appearance, while fourth cord 1$
smooth (Fig. 9A, C). Body whorl 50% of shell
length, with only upper one-half of spiral cords
crossed by axial ribs (Fig. 9A). Intersection of
ribs and cords delineate a series of deep rect-
angular depressions. Whorl sutures and
grooves between spiral cords striated. Auric-
ular aperture ovate, with thick, scalloped
outer lip. Single, prominent, acute columellar
fold on upper half of the columella, perpen-
dicular to the columella axis. Protoconch
smooth, dextrally heterostrophic, oriented
130° to teleoconch axis, submerged 30-35%
in first adult whorl, with earliest portion of pro-
toconch completely submerged (Fig. 9B-E).
Operculum, brown, auricular, paucispiral,
with subcentric nucleus, but lacking a notch
to accommodate columellar fold (Figs. 9F, С).

Head-foot: Opaque with scattered white
cells. Anterior portion of foot (= propodium)
with slight medial indentation and rounded
lateral edges. Foot narrowing posterior to

propodium, then widening to gradually taper
to a blunt posterior apex. Pedal gland open-
ing at posterior end of ventral surface of foot.
Attachment thread present. Tentacles subtri-
angular, ventro-laterally folded, joining ante-
riorly across midline (= connate); tentacular
pads present. Black eyes beneath epithelium
on median side of tentacles. Mentum un-
notched, not bifurcate (Fig. 11A). Visceral
mass of yellow and orange (with flecks of
brown) digestive tissue cells, translucent to
transparent reproductive organs.

Alimentary tract (Fig. 8A): Retracted intro-
vert-proboscis extending posteriorly from its
aperture on the ventral side of the head, dor-
sal to the mentum base and entering cephalic
hemocoel. Introvert joining the buccal sac,
which is connected to buccal pump. Buccal
pump divided into anterior (bp1) and poste-
rior sections (bp2) of equal length; bp1 nar-
row, round in cross-section; bp2 wider, lat-
erally flattened, distally rounded. Esophagus
originating on ventral surface of alimentary
tract at bp1-bp2 juncture, extending into the
visceral mass, where it joins the stomach.
Esophagus long, coiled, with an irregular or
tuberculate surface. Salivary gland ducts
penetrating alimentary tract immediately an-
terior to bp1-bp2 juncture, extending parallel
to one another within its walls, entering stylet
bulb without exiting gut. Salivary glands not
attached distally to alimentary tract.

Pallial cavity (Fig. 1A): Mantle and mantle
organs typical for members of the Odostom-
inae. Mantle edge finely crenulate. Ventral
and dorsal ciliated strips joining on mantle
roof at posterior end of mantle cavity. Small,
oblong pigmented mantle organ composed
primarily of cells filled with bright yellow ex-
udate and a few cells containing brown, or-
ange, or red contents; exudate released
when the snail is disturbed (Table 5). Small,
cream-colored gland beneath ventral ciliated
strip extending posteriorly from anterior edge
of ventral ciliated strip to 20-25% of strip’s
length. Gill absent.

Reproductive system: Typical of pyra-
midellids examined in this study (Fig. 5B). Pe-
nial complex unknown. Egg masses contain
approximately 25-35 eggs laid in irregular
gelatinous mass on host.

Nervous system: Typical of known pyra-
midellids (Fig. 6A).

Ecology and Distribution: Boonea cincta 1$ a
common eastern Pacific species, occurring
from Santa Barbara to the Gulf of California.

PYRAMIDELLID MORPHOLOGY AND PHYLOGENY 491

It parasitizes a number of gastropods (e.g.,
Norrisia norrisi, Astraea undosa, and A. gib-
berosa; LaFollette, 1977) and can often be
found on the dorsal surface of the operculum
of the trochid Tegula eiseni (LaFollette, 1977;
this study). This species’ shell was first de-
scribed based on a single, probably imma-
ture, specimen collected at Santa Barbara,
California (Carpenter, 1864). The shell sculp-
ture of B. cincta varies from highly cancellate
to almost entirely smooth (LaFollette, 1977).
Living snails collected for this study were all
cancellate. This species undergoes direct de-
velopment (veliger stage at approximately ten
days within the cocoon) with juveniles leaving
the egg mass at about 27 days (LaFollette,
1979).

Genus Odostomia Fleming, 1813

Odostomia Fleming, 1813: 76. Type-species:
Turbo plicata Montagu, 1803, by original
designation.

Diagnosis: Shell white or yellowish, short,
conical, 3-5 mm in length, with 4-6 adult
whorls. Whorls smooth to cancellate. Body
whorl 50-60% of shell length. Umbilicus small
or absent. Protoconch smooth, dextrally or
sinistrally heterostrophic oriented 120-150°
to teleoconch, partially submerged in first
adult whorl. Aperture ovate, with single acute
columella fold. Operculum brown, ovate, pau-
cispiral, with subcentric nucleus. Head-foot
white, with numerous opaque white cells (es-
pecially on tentacles and sides of head). Foot
narrowing posterior to propodium, widening
and then becoming attenuate posteriorly.
Pedal gland producing attachment thread.
Tentacles subtriangular, connate, ventro-lat-
erally folded; tentacular pads present. Eyes
subepithelial on median side of tentacles.
Mentum unnotched, not bifurcate. Introvert-
proboscis aperture on ventral side of head,
dorsal to mentum base. Introvert joining buc-
са! sac, which is composed of sucker, mouth,
sheathed stylet with separate opening, oral
tube, and stylet bulb. Buccal sac joining buc-
cal pump, which is divided into anterior (bp1)
and posterior sections (bp2). Esophagus orig-
inating on ventral surface of bp1-bp2 junc-
ture. Salivary gland ducts entering alimentary
tract and extending parallel to one another
within walls of bp1. Just posterior to buccal
sac, ducts exiting alimentary tract and enter-
ing stylet bulb.

Remarks: The genus Odostomia is one of four
genera recognized by Dall & Bartsch (1904,
1909) to which they assigned 40 subgenera
on the basis of a small number of convergent
shell characters (Abbott, 1974).

Odostomia babylonia (C. B. Adams, 1845)

Cheminitzia babylonia C. B. Adams, 1845: 6;
Clench & Turner, 1950: 259 (type lost,
fide Clench & Turner; type locality: Ja-
maica).

Odostomia (Cingulina) babylonica [sic] (C. B.
Adams); Bush, 1899: 176.

Odostomia (Cingulina) babylonica [sic] (C. B.
Adams); Verrill & Bush, 1900: 534, pl. 65,
fig. 11.

Odostomia (Miralda) judithae Usticke, 1959:
86-87, pl. 4, fig. 16, (Holotype: AMNH
198476, type locality: Sugar Bay, St.
Croix, U. S. Virgin Islands).

Cingulina judithae (Usticke); Usticke, 1969:
31.

Pyramidelloides judithae (Usticke); Usticke,
1971: 28.

Cingulina babylonia (C. B. Adams); Abbott,
1974: 301; DeJong & Coomans, 1988:
120, pl. 19; fig. 637.

Liamorpha babylonia (C. B. Adams); Faber,
1988: 81.

Remarks: Abbott (1974: 301), DeJong 4
Coomans (1988: 20) and Faber (1988: 81) all
considered Odostomia judithae Utiscke to be
a junior synonym of O. babylonia С. В. Adams.
Although Adams’ type material is lost, his de-
scription of this highly sculptured species is
unmistakable. Moreover, this species was ac-
curately figured by Verrill & Bush (1900).

Description: Shell (Fig. 13): Polished, trans-
parent, conical, 2 mm in length, composed of
3-4 adult whorls (Fig. 13A). Whorls posterior
to body whorl with two strong spiral cords:
one subsutural (a part ofthe shoulder at each
suture) and one equally dividing each whorl.
Numerous, irregular ridges perpendicular to
shoulder edge and often extending to the
suture (Fig. 13B). Body whorl 50% of shell
length. Each whorl with tightly spaced, lay-
ered, nearly orthocline growth lines that are
crossed by numerous microscopic spiral lines
(Fig. 13B). Aperture ovate, thick, ribbed at
outer lip. Single, prominent, acute columellar
fold on upper half of the columella, perpen-

492 WISE

dicular to columella axis. Protoconch smooth,
sinistrally heterostrophic, oriented 120° to te-
leoconch, submerged 30-35% in first adult
whorl, with earliest portion of protoconch par-
tially exposed (Fig. 13С-Е). Operculum light
brown, lenticular, paucispiral, with a subcen-
tric nucleus. Operculum lacking notch to ac-
commodate columellar fold (Fig. 13F, G).

Head-foot (Fig. 10B): White and opaque
to translucent. Anterior portion of foot with
slight medial indentation and rounded an-
tero-lateral edges. Foot narrowing posterior
to propodium, widening, becoming posteri-
orly attenuate. Pedal gland opening medial-
ly on posterior end of ventral surface of
foot. Attachment thread present. Aggregate
of large, white subepithelial cells lying just
anterior to operculum. Tentacles subtriangu-
lar, connate, ventro-laterally folded; tentacu-
lar pads present. Eyes black, large, round,
subepidermal close together on median side
of tentacles. Mentum unnotched attached to
foot laterally, not bifurcate (Fig. 11B). Visceral
mass of pale orange, light brown, dark blue,
or burgundy digestive tissue cells and
opaque reproductive organs.

Alimentary tract (Fig. 8C): Retracted intro-
vert-proboscis extending posteriorly from its
aperture on the ventral side of the head, dor-
sal to the mentum base and entering the
cephalic hemocoel. Introvert joining buccal
sac, which is connected to buccal pump.
Buccal pump is divided into anterior (bp1)
and posterior sections (bp2); bp1 elongate
spherical; bp2 one and one-half times length
of bp1, wider, oblong, circular in cross-sec-
tion. Esophagus originating on the ventral
surface of alimentary tract at bp1-bp2 junc-
ture, extending into the visceral mass to join
stomach. Esophagus long, coiled, with a tu-
berculate surface. Short, uncoiled salivary
gland ducts penetrating alimentary tract at
the distal end of bp1. Ducts extending paral-
lel to one another within the walls of bp1,
exiting the alimentary tract immediately pos-
terior to buccal sac and entering stylet bulb.
Salivary glands not attached distally to ali-
mentary tract.

Pallial cavity (Fig. 1А): Mantle and mantle
organs as in other Odostominae. Mantle
edge finely crenulate. Ventral and dorsal cil-
iated strips joining on mantle roof at posterior
end of mantle cavity. Small, oblong pig-
mented mantle organ composed primarily of
large cells containing bright yellow exudate
and a few cells filled with brown, red, or black
contents; exudate released when the snail is

disturbed (Table 5). Cream-colored, ventral
ciliated strip gland extending posteriorly from
beneath the anterior edge of the ventral cili-
ated strip to 20-25% of the strip’s length. Gill
absent.

Reproductive system: Typical of pyra-
midellids examined this study (Fig. 5B). Pe-
nial complex unknown.

Nervous system: Typical of known pyra-
midellids (Fig. 6A).

Ecology and Distribution: Odostomia babylo-
та is found littorally to just sublittorally, in-
habiting the underside of embedded rocks
and coral rubble, from West Indies (Abbott,
1974: 301). Host unknown.

Odostomia didyma Verrill & Bush, 1900

Odostomia (Cyclodostomia) didyma Мег! €
Bush, 1900: 533, pl. 65, fig. 14. (Holo-
type: PM 15706; type locality: Bermuda).

Odostomia didyma Мет! 8 Bush; DeJong 4
Coomans, 1988: 122, pl. 19, fig. 641.

Description: Shell (Fig. 14): Vitreous, thick,
conic, 2 mm in length, composed of 3-4 adult
whorls. Whorls with thick spiral cord just
above suture. Sloping subsutural shelf ventral
to each suture (Fig. 14C). Body whorl 50% of
shell length. Entire shell etched by micro-
scopic prosocline growth lines. Aperture
rhomboid with thick, basally flared outer lip.
Single, prominent, acute columellar fold on
upper half of the columella, perpendicular to
columella axis. Protoconch smooth, dextrally
heterostrophic, oriented 130° to teleoconch,
submerged 30-35% in first adult whorl, with
earliest portion of protoconch completely
submerged (Fig. 14B-E). Operculum light
brown, lenticular, paucispiral, with subcentric
nucleus. Operculum lacking notch to accom-
modate columellar fold (Figs. 14F, G).
Head-foot (Fig. 10C): Transparent to
opaque. Golden-yellow pigmentation present
on dorsal periphery of foot, mantle floor par-
allel to dorsal ventral ciliated strip, and the
length of tentacles laterally. White cells, al-
though scattered around head-foot, concen-
trated just posterior to eyes. Anterior portion
of foot with sharp antero-lateral projections
and deep medial indentation. Foot constrict-
ing immediately behind propodium, widen-
ing, narrowing again and tapering to a blunt
posterior apex. Pedal gland opening in mid-
die of a groove extending along the posterior

PYRAMIDELLID MORPHOLOGY AND PHYLOGENY 493

half of ventral surface of the foot. Attachment
thread present. Tentacles long, subtriangu-
lar, connate, ventro-laterally folded; tentacu-
lar pads present. Eyes black, subepithelial,
bean-shaped lying close together on median
side of tentacles. Mentum very short, un-
notched, not bifurcate, with lateral attach-
ments nearly even with its anterior edge (Fig.
11C). Visceral mass composed of brown,
grey, black, or burgundy digestive tissue
cells and opaque reproductive organs.

Alimentary tract (Fig. 8D): Retracted intro-
vert-proboscis extending posteriorly from its
aperture on the ventral side of the head, dor-
sal to the mentum base to enter cephalic
hemocoel. Introvert joining buccal sac, which
is connected to buccal pump. Buccal pump
divided into anterior (bp1) and posteri-
or sections (bp2); bp1 elongate, spherical;
bp2 one and one-half times length of the
bp1, wider, oblong, circular in cross-section.
Esophagus originating on alimentary tract at
juncture of bp1-bp2, and extending posteri-
orly to join the stomach within the visceral
mass. Esophagus long, coiled, with irregular
surface. Salivary gland ducts penetrating al-
imentary tract at distal end of bp1, continuing
anteriorly and parallel to one another within
the walls of the bp1, entering the stylet bulb
without exiting alimentary tract. Salivary
glands not attached distally to alimentary
tract.

Pallial cavity (Fig. 1А): Mantle and mantle
organs as in other Odostominae. Mantle
edge finely crenulate. Ventral and dorsal cil-
iated strips joining on mantle roof at posterior
end of mantle cavity. Small, oblong pig-
mented mantle organ composed primarily of
dark and variably colored cells filled with a
combination of: (a) yellow, black, red, and or-
ange, (5) yellow and dark brown, or (с) yellow,
brown, red or black contents. Snail exuding a
large amount of bright yellow exudate when
disturbed (Table 5). Cream-colored gland be-
neath ventral ciliated strip extending posteri-
orly from the strip’s anterior edge to 20-25%
its length. Gill absent.

Reproductive system: Typical of pyra-
midellids in this study (Fig. 5B). Penial com-
plex unknown. Producing cuticularized sper-
matophores, that, while commonly attached
by their bulbous end to the parietal wall of the
shell, were also seen attached to operculum
and immediately inside the shell's aperture.
Fresh spermatophores golden-brown.

Nervous system: Characteristic of known
pyramidellids (Fig. 6A).

Ecology and Distribution: Odostomia didyma
is found intertidally to subtidally on the un-
derside of embedded rocks and coral rubble,
from Bermuda to Curacao, Netherlands An-
tilles (Dejong 4 Coomans, 1988). Host un-
known.

Subfamily Sayellinae, new subfamily

Genus Sayella Dall, 1885

Sayella Dall, 1885: 286. Type-species: Leu-
сота hemphillii Dall, 1884, by original
designation.

Diagnosis: Shell brown, pupoid to elongate-
pupoid, with subsutural white band, 4-5 mm
in length, with 4-5 adult whorls. Sides of
whorls convex to straight. Body whorl 40-
50% of shell length. Umbilicus absent. Pro-
toconch smooth, dextrally heterostrophic,
partially submerged in first adult whorl, ori-
ented 140” to teleoconch. Aperture auricular,
with single columellar fold originating basally
on columella, extending at an oblique angle to
upper half of the columella. Operculum light
brown, lenticular, with subcentric nucleus.
Head-foot and mantle darkly pigmented. Foot
broad anteriorly, tapering posteriorly to blunt
point. Attachment thread absent. Tentacles
stout, cylindrical, not connate; tentacular
pads absent. Eyes black subepithelial, on me-
dian side of tentacles. Mentum emarginate,
short, with shallow longitudinal medial cleft.
Introvert-proboscis aperture medial at ante-
rior mentum tip. Introvert joining buccal sac
posteriorly; buccal sac composed of sucker,
sheathed stylet, mouth-stylet aperture, and
stylet bulb. Buccal sac extending posteriorly
to join buccal pump, which is divided into
anterior (bp1) and posterior sections (bp2).
Anterior esophagus originating at ventral sur-
face of buccal pump, continuing posteriorly
to join posterior esophagus and paired sali-
vary glands, forming a four-way junction. Sal-
ivary gland ducts attached to exterior of
esophagus. Anterior to anterior esophagus-
buccal pump junction, ducts detached and
entering stylet bulb. Salivary glands attached
distally to alimentary tract. Anteriorly tapered,
hooded penis outside and ventral to nerve
ring.

Remarks: Dall (1883) originally believed S.
hemphillii to be a freshwater ellobiid and as-
signed it to the genus [еисота. Later, Dall
(1885) placed this species and $. crosseana

494 WISE

(Dall, 1885) in his new subgenus Sayella within
the genus Melampus Montfort, 1810. Subse-
quently, Sayella was transferred to the Pyra-
midellidae by Morrison (1939) based on his
study of S. chesapeakea Morrison, 1939.

Sayella hemphillii (Dall, 1884)

Leuconia hemphillii Dall, 1884: 323, pl. 10,
fig. 6. (Holotype: USNM 36016; type lo-
cality: Cedar Key, Florida).

Melampus (Sayella) hemphillii (Dall); Dall,
1885: 286, pl. 18, fig. 11.

Sayella livida Rehder, 1935: 129, pl. 7, fig. 7;
Abbott, 1974: 300, fig. 3649 (Holotype:
USNM 125556; type locality: Corpus
Christi Bay, Texas); Harry, 1984: 68-70,
72, 74:

Odostomia (Syrnola) cf. livida Rehder, 1935;
Andrews, 1977: 127-128, unnumbered

fig.

Synonymic Remarks: Rehder (1935) noted
that the shells of S. /ivida and S. hemphillii are
very similar and suggested that examination
of more specimens may show that the two are
conspecific. Examination of the $. livida ho-
lotype and material collected in Florida indi-
cates that they are, in fact, the same species.

Description: Shell (Fig. 15): Elongate pupoid
(Fig. 15A) to pupoid (Fig. 15B), dark red-
brown, with whitish subsutural band at each
whorl, 4-5 mm in length, composed of 4-5
convex whorls. Adult whorls with numerous
microscopic, orthocline growth lines (Fig.
15D). Sutures shallow, simple. Body whorls
40-50% of shell length. In older individuals,
upper whorls and protoconch etched, pitted
or extensively eroded (Fig. 15A). Aperture
ovate in young specimens, elongate-ovate in
mature snails. Single columellar fold originat-
ing at base of columella, extending at an ob-
lique angle to upper half of columella (Fig.
15C). Protoconch smooth, dextrally heteros-
trophic, oriented 140° to teleoconch, sub-
merged 50-55% in adult shell, with earliest
portion of protoconch completely submerged
(Fig. 15D, E). Operculum light brown, lentic-
ular, with subcentric nucleus (Fig. 15G, H),
lacking notch to accommodate columellar
fold.

Head-foot (Fig. 16A): Generally heavily pig-
mented giving snail a “sooty” appearance
(although a few snails were only lightly pig-
mented). White to opaque cells between eyes
and scattered throughout dorsal surface of

foot. Anterior edge of foot convex, with slight
rounded lateral projections. Foot narrowing
posterior to propodium, widening, ending in
bluntly attenuated tip. Attachment thread ab-
sent. Tentacles cylindrical, stout, not con-
nate; tentacular pads absent. Eyes black,
subepithelial, round, on median side of ten-
tacles. Mentum emarginate, short, anteriorly
rounded, with shallow longitudinal medial
cleft (Fig. 11H). Visceral mass composed of
grey, black, or light brown digestive tissue
cells and transparent to translucent repro-
ductive structures.

Alimentary tract (Fig. 17A): Retracted intro-
vert-proboscis extending posteriorly from its
medial aperture at the anterior mentum tip to
enter the cephalic hemocoel and join the
buccal sac. Buccal sac continuing posteriorly
to buccal pump, which is divided into anterior
(bp1) and posterior sections (bp2); bp1 two
times the length of the laterally flattened bp2.
Anterior esophagus originating on ventral
surface of alimentary tract at bp1-bp2 junc-
ture. Anterior esophagus joining posterior
esophagus and paired salivary glands to
form a four-way junction. Posterior esopha-
gus widening posterior to four-way junction
and extending into visceral mass to join the
stomach. Salivary gland ducts arranged in
tight folds affixed to exterior of anterior
esophagus, extending anteriorly, leaving an-
terior esophagus at esophagus-buccal pump
junction, straightening and entering stylet
bulb. Salivary glands attached distally to ali-
mentary tract at posterior esophagus.

Pallial cavity (Fig. 1A, C): Mantle and man-
tle organs typical for pyramidellids. Mantle
edge scalloped. Iridescent ventral and dorsal
ciliated strips joining on the mantle roof at
posterior end of mantle cavity. Small, oblong
pigmented mantle organ composed primarily
of transparent and white cells, with a few
scattered cells filled with red, yellow, and or-
ange contents (Fig. 1A). Pigmented mantle
organ producing no exudate. Gland beneath
ventral ciliated strip underlying entire ventral
ciliated strip, darkly pigmented, with a few
scattered red cells, producing and exuding
copious milky-blue exudate (Table 5).
Cream-colored kidney visible through trans-
parent dorsal pallial roof, because the mantle
area above this organ unpigmented. Gill ab-
sent.

Reproductive system: Typical of pyra-
midellids in this study (Fig. 5B). Penis within
head just anterior and ventral to nerve ring
(Fig. 12B). Penis in a pocket that opens me-

PYRAMIDELLID MORPHOLOGY AND PHYLOGENY 495

dially and ventral to mentum base. Elongate
anterior section of penis cylindrical, with a
subapical swelling that narrows to form a
short terminal nipple. Anterior portion framed
posteriorly by large, pleated fleshy hood that
contains numerous glandular cells. Penis an-
chored to pocket posteriorly and at base of
hood by muscle and connective tissue.

Nervous system: Typical for known pyra-
midellids (Fig. 6A).

Ecology and Distribution: Sayella hemphillii
occurs intertidally to subtidal in the surface
layers of sand and mud flats in the Gulf of
Mexico along the coast Texas and west Flor-
ida (Abbott, 1974). In this study, $. hemphillii
were collected at Cedar Key, Florida, in areas
with large concentrations of the polychaete
Onuphis magna, which is a possible host.

Genus Petitella, new genus

Type-species: Melampus (Sayella) crosseana
Dall, 1885: 286, here designated.

Diagnosis: Shell pupoid to elongate pupoid,
semitransparent and yellow brown around
lower periphery of body whorl, 3-4 mm in
length, composed of 4-5 straight to slightly
convex adult whorls. Whorls with numerous,
nearly orthocline microscopic growth lines.
Body whorl 50% of shell length. Protoconch
smooth, dextrally heterostrophic, submerged
50-55% in first adult whorl, oriented 140-
145” to teleoconch, with earliest portion of
protoconch completely submerged. Aperture
elongate-ovate, with thin outer lip slightly
flared basally. Single, columella fold originat-
ing at base of columella, extending obliquely
to upper half of the columella. Parietal wall
dark golden-brown. Operculum tan, auricular,
with subcentric nucleus. Head-foot opaque to
transparent, with white cells concentrated at
mentum. Foot with broad anterior and slightly
convex rounded lateral edges, tapering pos-
teriorly to blunt apex. Attachment thread ab-
sent. Tentacles stubby, cylindrical, rounded
apically, not connate; tentacular pads absent.
Eyes black subepithelial, spherical, on me-
dian side of tentacles. Mentum emarginate,
antero-laterally rounded with medial longitu-
dinal cleft. Introvert-proboscis aperture at an-
terior tip of mentum. Introvert joining buccal
sac, which is composed of sucker, sheathed
stylet, mouth/stylet aperture, and stylet bulb.
Long, undifferentiated buccal pump posterior
to buccal sac. Buccal pump without blind sac

or caecum. Esophagus extending posteriorly,
entering visceral mass and joining stomach.
Salivary gland ducts penetrating alimentary
tract at stylet bulb. Salivary glands attached
distally to alimentary tract at anterior portion
of esophagus. Retracted penis folded within
pocket that opens ventro-medially to mentum
outside and ventral to the nerve ring.

Petitella crosseana (Dall, 1885)

Melampus (Sayella) crosseana Dall, 1885:
286, pl. 18, fig. 10. (Holotype: USNM
37613; type locality: Egmont Key, Flor-
ida).

Sayella crosseana (Dall); Abbott, 1974: 300.

Remarks: Petitella crosseana was originally
assigned to the genus Sayella on the basis of
shell characters. Anatomically, P. crosseana
is unlike any known sayellids (e.g., Sayella
hemphillii) or for that matter any known pyra-
midellids. Consequently, | propose that it be
placed т a new genus. Etymology: Petitella 15
named for Richard E. Petit in recognition of his
contributions to malacology.

Description: Shell (Fig. 18): Polished, semi-
transparent, pupoid to elongate pupoid, yel-
low brown around lower periphery of body
whorl (Fig. 18A), 3-4 mm in length, composed
of 4-5 straight to slightly convex adult whorls.
Whorls with numerous, nearly orthocline mi-
croscopic growth lines (Figs. 18B, C). Body
whorl 50% of shell length. Upper adult whorls
and protoconch often eroded and pitted. Ap-
erture elongate-ovate, with thin outer lip
slightly flared basally. Single, columella fold
originating at base of columella, extending
obliquely to upper half of columella. Parietal
wall dark golden-brown. Protoconch smooth,
dextrally heterostrophic, submerged 50-55%
in first adult whorl, oriented 140-145” to te-
leoconch, with earliest portion of protoconch
completely submerged (Fig. 18С-Е). Opercu-
lum tan, auricular, with subcentric nucleus
(Fig. 16F,G):

Head-foot (16B): Opaque to transparent,
with white cells concentrated at mentum. An-
terior portion of short foot broad, slightly con-
vex, with rounded lateral edges. Foot taper-
ing posteriorly to blunt apex. Attachment
thread absent. Tentacles cylindrical, stubby,
rounded apically, not connate; tentacu-
lar pads absent. Eyes black, subepithelial,
spherical, on median side of tentacles. Men-

496 WISE

tum emarginate, antero-laterally rounded,
with medial longitudinal cleft (Fig. 111). Vis-
ceral mass of white-opaque reproductive or-
gans and a distinctive branching network of
brown to black digestive tissue cells, this net-
work generally arranged perpendicular to the
coiling axis of the visceral mass and visible
through shell.

Alimentary tract (Fig. 17B): When retracted,
introvert-proboscis extending posteriorly
from its medial aperture at anterior tip of
mentum to enter cephalic hemocoel and join
buccal sac. Long, undifferentiated muscular
conduit posterior to buccal sac functioning
as buccal pump. Buccal pump without blind
sac or caecum. Esophagus extending poste-
riorly, entering the visceral mass and joining
stomach. Salivary gland ducts penetrating al-
imentary tract at stylet bulb. Salivary glands
attached distally to alimentary tract at ante-
rior portion of esophagus.

Pallial cavity (Fig. 1A, C): Mantle and man-
tle organs as in other pyramidellids. Mantle
edge finely scalloped. Small, oblong, pig-
mented mantle organ containing either large
cells filled with a bright yellow exudate and a
few cells containing orange contents, or cells
with black contents and a small number of
cells filled with a white exudate. Pigmented
mantle organ secreting a light blue exudate
when snail disturbed (Table 5). Gland be-
neath ventral ciliated strip composed mostly
of large white cells mixed with a few black
cells, extending the length of the ventral cili-
ated strip. Gill absent.

Reproductive system: Typical of pyra-
midellids herein (Fig. 5B). Penis outside and
ventral to the nerve ring. Retracted penis
folded within a pocket that opens ventro-me-
dially to mentum (Fig. 12C). Posterior end an-
chored to floor of pocket by several retractor
muscles. Penis anteriorly attenuate, posteri-
orly bulbous. Shallow dorsal groove extends
posteriorly from penis anterior to one-half pe-
nis length.

Nervous system: Characteristic of known
pyramidellids (Fig. 6A).

Ecology and Distribution: Petitella crosseana
occurs intertidally to subtidally in the surface
layers of mud and sand flats in the Gulf of
Mexico along the coast of Texas and west
Florida (Abbott, 1974), and in the Atlantic,
South Carolina (Merrill & Petit, 1965) to Florida
(present study), south to the West Indies (Ab-
bott, 1974). Host unknown.

Subfamily Pyramidellinae Gray, 1840
Genus Pyramidella Lamarck, 1799

Obeliscus Humphrey, 1797: 24. [Rejected
work, 1.C.Z.N. Opinion 51].

Pyramidella Lamarck, 1799: 76. Type-spe-
cies: Trochus dolabratus Linnaeus,
1758, by monotypy.

Pyramidellus Montfort, 1810: 499. Type spe-
cies: Trochus dolabratus Linnaeus,
1758, by monotypy.

Aphalista Laseron, 1959: 1876. Type species:
Pyramidella mitralis A. Adams, 1853, by
original designation.

Diagnosis: Shell elongate-conical, porcella-
neous, generally white or brown, with or with-
out bands or spots, reaching 50 mm in length.
Sides of whorls convex to straight. Body
whorl approximately 40% of shell length. Um-
bilicus present. Protoconch smooth, sinis-
trally heterostrophic protoconch oriented
120-125° to teleoconch, partially submerged
in first adult whorl. Aperture elongate-ovate,
with one large and two smaller prominent col-
umellar folds. Operculum brown, elongate-
ovate, notched to accommodate largest col-
umellar fold. Head-foot light yellow, with a
prominent mass of white cells between and
posterior to eyes. Foot wide anteriorly, with
shallow medial indentation and bluntly ta-
pered posterior apex. Attachment thread ab-
sent. Tentacles triangular, connate, medially
notched, laterally folded; tentacular pads ab-
sent. Eyes black, subepithelial on median side
of tentacles. Mentum retuse, broad anteriorly,
with sharp antero-lateral projections and shal-
low longitudinal groove. Introvert-proboscis
aperture opening medially on mentum tip.
Introvert connecting posteriorly to buccal
sac, which is composed of sucker, sheathed
stylet, mouth-stylet aperture and stylet bulb.
Buccal sac joining buccal pump separated
into very short anterior section (bp1) and very
elongate, laterally flattened posterior section
(bp2). Esophagus divided into anterior and
posterior sections that, with the salivary
glands, form a four-way junction. Salivary
gland ducts, attached to exterior of anterior
esophagus, extending anteriorly and entering
stylet bulb. Salivary glands attached distally
to alimentary tract at posterior esophagus.
Ciliated, scoop-shaped penis, with bulbous
posterior end outside and ventral to nerve
ring.

Remarks: The genus Pyramidella was first
proposed by Lamarck in 1799. Historically,
confusion has existed over the use of Pyra-

PYRAMIDELLID MORPHOLOGY AND PHYLOGENY 497

midella because the genus Obeliscus was of-
ten used in its place in the older literature.
However, the name Obeliscus is no longer
considered available because it was pro-
posed by Humphrey (1797), a work rejected
for nomenclatural purposes (l.C.Z.N. Opinion
Sl):

Pyramidella sulcata (A. Adams, 1854)

Obeliscus sulcatus A. Adams, 1854: 807, pl.
171, fig. 34. (Holotype: BMNH 1986:
284; type locality: Tahiti).

Obeliscus monilis A. Adams, 1854: 806, pl.

171, fig. 12.

Obeliscus teres A. Adams, 1854: 807, pl. 171,
IGS. 31. 32;

Obeliscus tessellatus A. Adams, 1854: 808,
ре 17 Ъ ПО. 16.

Pyramidella pratii Bernardi, 1859: 386, pl. 13,
19. 1.

Pyramidella teres (A. Adams);
1669: pl. 1,110.00:

Pyramidella tessellatus (A. Adams); Sowerby,
1865: pl. 1, fig. 4.

Pyramidella sulcata (A. Adams); Tryon, 1886:
301, pl. 72, figs. 79-83; Cernohorsky,
19727200, pl. 57, 10.2, 23; Kay, 1979:
413, fig. 133B.

Wingenella pricena Laseron, 1959: 190-191,
las: 17. 18:

Wingenella eburnea Laseron, 1959: 190, figs.
14-16.

Sowerby,

Synonymic Remarks: The nominal species
Obeliscus teres A. Adams, O. tessellatus A.
Adams, O. monilis A. Adams, were named at
the same time as O. sulcata A. Adams. Tryon
(1886: 301) considered the four to be con-
specific and acted as first reviser in selecting
O. sulcatus as the senior synonym. Pyra-
midella pratii Bernardi, Wingenella eburnea
Laseron, and W. pricena Laseron are also
considered conspecific (Cernohorsky, 1972:
200).

Description: Shell (Fig. 19): White, polished,
elongate-conical, with orange-brown squar-
ish spots, 20-30 mm in length, composed of
12-13 adult whorls. Sides of whorls convex to
straight, with microscopic orthocline growth
lines (Fig. 19B). Sutures deeply channeled.
Body whorl 40% of shell length. Upper whorls
and protoconch, often eroded and pitted (Fig.
19F, H). Aperture elongate-ovate, with thin
outer lip and thick columella. One large col-
umellar fold on upper half of columella dorsal

to two smaller folds (Fig. 19C). Columellar
folds perpendicular to the columella axis.
Largest columellar fold with four well-devel-
oped ridges composed of overlapping imbri-
cate plates (Fig. 19D, E); ridges partially worn
or highly eroded (resorbed?) in the whorls
above the penultimate whorl (= whorl preced-
ing body whorl). The two smaller columellar
folds smooth. Six to eight well-developed pal-
atal teeth present deep inside the aperture of
the body whorl (Fig. 19C). Rudimentary pal-
atal teeth in various stages of ontogeny
present immediately inside the outer lip in
some. Protoconch smooth, sinistrally het-
erostrophic, oriented 120% to teleoconch,
submerged 40-45% in first adult whorl, with
earliest portion of protoconch partially ex-
posed (Fig. 19F, H). Operculum light brown,
elongate-ovate, notched to fit largest col-
umellar fold (Fig. 191).

Head-foot (Fig. 204): Light yellow, with
prominent clumps of subepithelial, numerous
white cells posterior to and between eyes, on
tentacles, mentum, and foot, particularly just
anterior to operculum. Foot blunt posteriorly,
widening anteriorly, narrowing, then widening
again at slightly bifid propodium. Propodium
with rounded antero-lateral edges. Attach-
ment thread absent. Tentacles triangular,
connate, medially notched, laterally folded;
tentacular pads absent. Eyes black, subepi-
thelial on median side of tentacles. Mentum
retuse, with medial longitudinal groove (Fig.
11D). Columellar muscle divided into three
sections. Middle and longest portion, at-
tached to columella within the penultimate
whorl, tapering anteriorly and joining four-
way junction of alimentary tract. Visceral
mass composed of grey, black, brown, or
dark red-brown digestive tissue cells and
opaque reproductive organs. Visceral mass
partially covered in densely packed small
white cells.

Alimentary tract (Fig. 21A): When retracted,
introvert-proboscis extending posteriorly
from its medial aperture on the anterior apex
of mentum to enter cephalic hemocoel. Intro-
vert entering cephalic hemocoel, forming a
tight coil, and joining the buccal sac. Buccal
sac joining buccal pump, which is divided
into laterally flattened posterior section (bp2),
which is 10 times length of anterior section
(bp1). Anterior esophagus originating on ven-
tral surface of alimentary tract at juncture of
bp1-bp2. Anterior esophagus stretching pos-
teriorly, joining posterior esophagus and
paired salivary glands to form four-way junc-

498 WISE

tion. Posterior esophagus narrow at four-way
junction, widening posteriorly, extending into
visceral mass and joining stomach. Highly
folded, tightly packed salivary gland ducts at-
tached to outside of anterior esophagus.
Ducts extending from salivary glands to an-
terior esophagus-buccal pump juncture, then
straightening and entering stylet bulb. Sali-
vary glands attached distally to alimentary
tract at anterior end of posterior esophagus
by connective tissue or muscle. There are
two pairs of retractor muscles. The first pair
long, extending anteriorly from that portion of
muscular enclosure even with the buccal
sucker, attaching to the proboscis. The sec-
ond pair originating further anterior on the
proboscis, where first pair terminates and
muscular sleeve ends, continuing anteriorly
approximately 1 mm and attaching to the
proboscis.

Pallial cavity (Fig. 1B): Mantle and mantle
organs typical for the Pyramidellinae. Mantle
edge smooth. Ventral and dorsal ciliated
strips joining on mantle roof at posterior end
of mantle cavity. Very large, pigmented man-
tle organ, composed of transparent and
opaque cells, extending posteriorly, narrow-
ing and terminating at convergence of ventral
and dorsal ciliated strips. Pigmented mantle
organ secreting a very viscid, clear to opaque
substance. Large, yellow gland beneath ven-
tral ciliated strip extending posteriorly from
anterior edge of ventral strip to strip's termi-
nus. Gland composed primarily of large cells
filled with a yellow exudate and a few cells
containing red and white contents; bright yel-
low exudate released when the snail dis-
turbed (Table 5). Osphradium subtriangular,
with numerous elliptical white cells, mostly
concentrated on extreme left side of mantle
roof, narrowing as it extends laterally, termi-
nating just posterior to right mantle edge (Fig.
6B). Small, white gland cells originating at the
convergence of dorsal and ventral strips and
extending anteriorly atop the middle of the
gill to the anterior mantle edge. Gill com-
posed of a series of grooves and ridges per-
pendicular to and enclosed by opposing cil-
iated strips (Fig. 4B). Individual gill filaments
highly folded (Fig. 4C). Medial section of gill
surface with scattered tufts of long cilia, bor-
dered laterally by densely packed shorter
cilia.

Reproductive system: Typical of pyra-
midellids discussed herein (Fig. 5B). Penis in
cavity outside and ventral to nerve ring (Fig.
22A). Protrusile penis with sides that fold in-

ward to form a narrow groove that extends
length of organ. Grooved and deltoid tip of
penis ciliated. Posterior portion of penis
composed of bulbous halves containing
brown glandular cells. Muscle fibers and con-
nective tissue attached to posterior of penis
function as retractors and anchor penis to its
enclosure. Penis exiting the body through
medial opening ventral to mentum.

Nervous system: As in other pyramidellids
(Fig. 6A), with one exception: the osphradial
nerve bifurcating to innervate both right and
left portions of osphradium (Fig. 6B).

Ecology and Distribution: Pyramidella sulcata
occurs intertidally to subtidally throughout the
Indo-Pacific (Cernohorsky, 1972). It remains
within the sand during the day and at night is
epifaunal (this study). Host unknown.

Pyramidella crenulata (Holmes, 1859)

Pyramidella crenulata Holmes, 1859: 88, pl.
13, figs. 14, 14a. (Holotype: AMNH
099185; type locality: Pleistocene; South
Carolina.

Pyramidella (Longchaeus) crenulata
(Holmes); Perry & Schwengel, 1955: 118,
pl. 23, fig. 154; Andrews, 1977: 127, un-
numbered fig.; Abbott, 1974: 291, fig.
3462.

Obeliscus arenosa Conrad; Tuomey &
Holmes, 1857: pl. 26, fig. 17 (not of Con-
rad, 1843: 309).

Description: Shell (Fig. 23): Polished and
elongate-conical, 12-14 mm in length, com-
posed of 10-12 adult whorls. Whorls flat to
slightly convex, with moderately deep crenu-
lated sutures (Fig. 23B). Two color forms: (1)
solid white and (2) brown with white spots.
Body whorl 40% of shell length. Protoconch
and upper adult whorls eroded and pitted (Fig.
23A, E). Aperture elongate-ovate, with thick
columella and thin outer lip. One large col-
umellar fold on upper half of columella, dorsal
to two smaller folds, all folds perpendicular to
columella axis (Fig. 23C). Large columellar
fold with a single ridge composed of disjunct
and/or imbricate plates and bordered by an
outer notched edge (Fig. 23D). Smaller col-
umellar folds smooth. Generally with 3-4 fully
developed palatal teeth within aperture of
body whorl (Fig. 23C). Rudimentary palatals
are usually present just inside outer lip. Pro-
toconch smooth, sinistrally heterostrophic,
oriented 120° to teleoconch, submerged 40-

PYRAMIDELLID MORPHOLOGY AND PHYLOGENY 499

45% in first adult whorl, with earliest portion
of protoconch partially exposed (Fig. 23E, G).
Operculum light brown, elongate-ovate,
notched to accommodate largest columellar
fold (Fig. 23H, |).

Head-foot (Fig. 20B): White with prominent
white cells dispersed throughout but partic-
ularly ventral to and surrounding antero-dor-
sal edge of operculum. Foot blunt posteriorly,
wide across midfoot, narrowing at anterior
end. Propodium slightly bifid, with rounded
antero-lateral edges. Attachment thread ab-
sent. Tentacles triangular, connate, medially
notched, laterally folded tentacles; tentacular
pads absent. Eyes black subepithelial, spher-
ical, on median side of tentacles. Mentum re-
tuse mentum with medial longitudinal groove
(Fig. 11D). Columella muscle as in P. sulcata.
Visceral mass with dark, brown-red digestive
cells and opaque to translucent reproductive
organs. Visceral mass coils partially covered
by numerous densely packed white cells.

Alimentary tract (Fig. 21A): When retracted,
introvert-proboscis extending posteriorly
from its medial aperture on the anterior men-
tum tip to enter cephalic hemocoel. Introvert
entering cephalic hemocoel twisted into sin-
gle tight coil, joining buccal sac. Buccal
pump as in P. sulcata. Short anterior esoph-
agus uniting with buccal pump at juncture of
bp-bp2, extending posteriorly, joining poste-
rior esophagus and salivary glands, forming a
four-way junction. Posterior esophagus nar-
row at four-way junction, widening posteri-
orly prior to entering the visceral mass and
joining stomach. Salivary gland ducts affixed
to exterior of anterior esophagus. These
highly folded ducts extending anteriorly from
anterior esophagus-buccal pump junction,
entering stylet bulb. Salivary glands attached
distally to alimentary tract at anterior portion
of posterior esophagus. Retractor muscles
as in P. sulcata.

Pallial cavity (Fig. 1B): Mantle configuration
typical for the Pyramidellinae. Mantle edge
smooth. Ventral and dorsal ciliated strips
joining on mantle roof at posterior end of
mantle cavity. Very large, elongate pig-
mented mantle organ of clear to translucent
cells with a few scattered white cells, releas-
ing a small amount of opaque substance
flecked with white. Large, yellow gland be-
neath ventral ciliated strip extending length of
the strip. Gland composed primarily of large
cells filled with yellow exudate and a few cells
with white and red contents; viscid, bright
yellow exudate released when snail disturbed

(Table 5). Osphradium as in P. sulcata. Rows
of small white and brown cells at the conver-
gence of dorsal and ventral ciliated strips, ex-
tending anteriorly atop middle of gill, termi-
nating at anterior edge of mantle floor. Gill as
in P. sulcata.

Reproductive system: Typical of pyra-
midellids in this study (Fig. 5B). Penial com-
plex as in P. sulcata (Fig. 22A).

Nervous system: As in other taxa within the
subfamily Pyramidellinae (Fig. 6A, B).

Ecology and Distribution: Pyramidella crenu-
lata occurs intertidally to subtidally in sand
and mud from North Carolina to Texas and
West Indies (Abbott, 1974). As with other
members of the subfamily, it is probably epi-
faunal at night. Host unknown.

Pyramidella mitralis A. Adams, 1854

Pyramidella mitralis A. Adams, 1854: 814, pl.
172, fig. 9; (Holotype: BMNH 19862799;
type locality: St. Estevan, North llocos,
Isle of Luzon, Philippine Islands); 1855:
177; Sowerby, 1865, pl. 3, species 20.

Pyramidella propingua A. Adams, 1854: 814;
pl 172,119. 8; 18557177;

Pyramidella variegata A. Adams, 1854: 814,
pl. 172, fig. 10; 1855: 178.

Pyramidella (Otopleura) mitralis A. Adams;
Tryon, 1886: 305, pl. 73, figs. 94, 97, 2,
3.

Aphalista mitralis (A. Adams); Laseron, 1959:
187, figs. 4-6.

Otopleura mitralis (A. Adams); Cernohorsky,
1972: 201, pl. 57, fig. 6-6С; Kay, 1979:
412, fig. 133C.

Synonymic Remarks: Pyramidella propinqua
A. Adams and P. variegata A. Adams are
placed in synonymy on the authority of Tryon
(1886: 305) and Cernohorsky (1972: 201).

Description: Shell (Fig. 24): Thick, mitriform,
polished, 12-15 mm in length, with 9-10 adult
whorls. Shell elongate-ovate to elongate-nar-
row, variable in color. Elongate-ovate forms
with convex whorls, few to many prominent
axial ribs, and often colored with brown flam-
mules (Fig. 24A). Narrow-elongate forms of-
ten with numerous weak axial ribs (Fig. 24B),
sides of whorls convex to straight and banded
or plain. In all forms, body whorl 50% of shell
length. Adult whorls finely perforate (Fig. 24A).
Protoconch and upper adult whorls often
eroded and pitted (Fig. 24E, H). Aperture elon-
gate-ovate, with thick outer lip. One large col-

500 WISE

umellar fold on upper half of upper columella,
dorsal to two smaller folds (Fig. 24A, C), all
folds perpendicular to the columella axis.
Smaller folds smooth, larger fold with 3-4
ridges constructed of a series of overlapping
or imbricate plates (Fig. 24D). Plates unidi-
rectional, oriented opposite to coiling direc-
tion. Six to seven unequal palatal teeth deep
inside outer lip, with largest denticle in middle
of row. Rudimentary palatals usually present
just inside outer lip. Protoconch smooth,
heterostrophic, oriented 120° to teleoconch,
40-45% submerged in first adult whorl, with
earliest portion of protoconch partially sub-
merged (Fig. 24E, H). Operculum tan, elon-
gate-ovate, notched to accommodate largest
columeliar fold (Fig. 241).

Head-foot (Fig. 20C): White, with aggre-
gates of large numerous white cells between
and posterior to eyes, on tentacles, mentum,
and foot, particularly concentrated just ante-
rior of operculum. Anterior end of foot wide
anteriorly, with slight medial indentation and
acute antero-lateral projections. Foot nar-
rowing posterior to propodium, widening,
then tapering again to blunt apex. Tentacles
triangular, connate, medially notched, later-
ally folded; tentacular pads absent. Eyes
black, subepithelial, spherical, on median
side of tentacles. Mentum retuse, with medial
longitudinal groove (Fig. 11D). Columellar
and retractor muscles as in P. sulcata. Vis-
ceral mass containing reddish-dark brown
digestive tissue cells, translucent to opaque
reproductive structures. Visceral coils par-
tially covered by densely packed white cells.

Alimentary tract (Fig. 21A): Introvert-pro-
boscis arrangement and aperture, buccal
sac, buccal pump and remainder of alimen-
tary tract as in P. sulcata. Retractor muscle
arrangement as in P. sulcata.

Pallial cavity (Fig. 1B): Mantle and mantle
organs typical for the Pyramidellinae. Mantle
edge coarsely crenulate. Ventral and dorsal
ciliated strips joining on mantle roof at pos-
terior end of mantle cavity. Very large, pig-
mented mantle organ containing variably
sized opaque to clear cells with a few periph-
eral yellow and red cells, narrowing posteri-
orly, terminating at convergence of dorsal
and ventral ciliated strips. Organ releasing a
small amount of opaque substance flecked
with white. Large, yellow gland beneath ven-
tral ciliated strip, extending length of strip,
containing primarily large yellow cells and a
small number of cells filled with red or white
contents. Bright yellow exudate secreted

when snail disturbed (Table 5). Osphradium
as in P. sulcata. Thin line of white glandular
cells extending anteriorly from juncture of cil-
iated strips atop gill, terminating at anterior
edge of mantle roof. Gill as in P. sulcata.

Reproductive system: Typical of pyra-
midellids in this study (Fig. 5A). Penial com-
plex like that of P. sulcata (Fig. 22A).

Nervous system: Typical of the Pyramidel-
linae (Figs. 6A8B).

Ecology and Distribution: Pyramidella mitralis
occurs intertidally to subtidally on sand flats
throughout the Indo-Pacific (Cernohorsky,
1972). This snail remains buried during the
day and is epifaunal at night (this study). Stud-
ies in Mozambique showed that this species
occurs with the enteropneust Ptychodera
flava, which may serve as its host (MacNae 8
Kalt, 1958).

Subfamily Turbonillinae Simroth, 1907
Genus Turbonilla Risso, 1826

Turbonilla Risso, 1826: 224. Type species:
Turbonilla typica Dall & Bartsch, 1903
(new name for Turbonilla plicatula Risso,
1826, non Turbo plicatula Brocchi,
1814), by subsequent designation of Dall
8 Bartsch, in Arnold, 1903:269. See Re-
marks below.

Diagnosis: Shell white and lanceolate, 9 mm
in length, with 10-12 adult whorls. Whorls
slightly convex to straight. Each whorl with
prominent axial ribs, extending whorl length,
except on body whorl, where axial ribs ter-
minate prior to base. Intervening spaces
present between ribs. Body whorl 20% of
shell length. Umbilicus absent. Protoconch
smooth, sinistrally heterostrophic, perpen-
dicular to teleoconch, partially submerged
in first adult whorl. Aperture squarish, with
straight outer lip and slightly flared base. Col-
umellar folds absent. Operculum tan, lentic-
ular, with subcentric nucleus. Head-foot
white. Short foot, anteriorly truncated, with
lateral projections, tapered posterior. Tenta-
cles elongate, subtriangular, connate, ventro-
laterally folded; tentacular pads absent. Eyes
black, subepithelial on median side of tenta-
cles. Mentum incised, with shallow, longitu-
dinal groove. Introvert opening medial on an-
terior mentum tip. Buccal sac composed of
sucker, sheathed stylet, mouth/stylet aper-
ture and stylet bulb. Penis extending through
nerve ring beside proboscis. Anterior portion

PYRAMIDELLID MORPHOLOGY AND PHYLOGENY 501

of penis with several lateral rows of minute
cuticular hooks.

Remarks: Risso (1826) introduced Turbonilla
without designating a type species. He in-
cluded four species in the new genus: Tur-
bonilla costulata Risso, 1826; Turbo gracilis
Brocchi, 1814; and Turbo plicatula Risso,
1826; and Turbonilla humboldti Risso, 1826.
Almost all authors show the type species of
Turbonilla either as Turbo lacteus Linnaeus,
1758, or as Turbo elegantissimus Montagu,
1803, often with the later in the synonymy of
former. However, neither are originally т-
cluded species, and are therefore, not eligible
for type designation. Powell (1979: 256)
shows the type species as 7. striata Montagu,
by subsequent designation of Gray (1847).
This is not correct because Gray (1847: 160)
listed 7. elegantissima as the type species. A
search of the literature has revealed no source
for “T. striata Montagu,” and this name 15
evidently a lapsus calami.

The earliest apparent valid type designa-
tion is that of Dall & Bartsch (in Arnold, 1903),
in which they propose the new name Turbo-
nilla typica as a replacement name for Т. pli-
catula Risso, 1826, not Turbo (= Turbonilla)
plicatula Brocchi. Although there is no inter-
nal evidence, other than the specific name,
that Risso was simply transferring Brocchi's
species to his new genus, many authors have
considered this to be the case, with Bronn
(1843: 1328) apparently the first to do so. Ar-
naud (1978: 129), for Risso's Т. plicatula,
stated: “C'est Turbo plicatulus Brocchi,
1814, annexé par Risso!”.

Turbonilla hemphilli Bush, 1899

Turbonilla hemphilli Bush, 1899: 169, pl. 8,
fig. 3. (Holotype: ANSP 79013; type lo-
cality: Sarasota Bay, Florida); Andrews,
1971: 132; Abbott, 1974: 302-303, fig.
3682.

Turbonilla unilirata Bush, 1899: 165, pl. 8, fig.
6. (Holotype: ANSP 79010; type locality:
St. Thomas, West Indies).

Turbonilla penistoni Bush, 1899: 165-166, pl.
8, fig. 14. (Holotype: ANSP 70024; type
locality: Bermuda).

Turbonilla heilprini Bush, 1899: 167-168, pl.
8, fig. 13. (Holotype: ANSP 79009; type
locality: Bermuda).

Turbonilla abrupta Bush, 1899: 168, pl. 8, fig.
4. (Holotype: ANSP 79012; type locality:
St. Thomas, West Indies).

Synonymic Remarks: Holotypes of the above
species were examined and on the basis of
shell morphology determined to be conspe-
cific with T. hemphilli. Because these names
were all proposed in the same work, T. hemp-
hilli is here selected as senior synonym under
the Principle of the First Reviser (1.С.7.М. Ar-
ticle 24).

Description: Shell (Fig. 25): Thick, dull white,
acutely lanceolate, 7-8 mm in length, com-
posed of 12-13 adult whorls with sides
straight to slightly convex (Fig. 25A). Whorls
with 17-18 axial ribs. Except for body whorl,
axial ribs the length of each whorl, with an
elongate rectangular depression between
each rib (Fig. 25B). Axial ribs on body whorl
terminating at one-half of whorl’s length.
Body whorl 20% of shell length. Adult whorls
etched by numerous fine prosocline growth
lines. Aperture subquadrate, with base of
outer lip slightly flared. Columellar folds ab-
sent. Protoconch smooth, sinistrally hetero-
strophic, oriented 90° to teleoconch axis,
submerged 5-10% in first adult whorl, with
earliest portion of protoconch exposed (Fig.
25C-E). Operculum brown, lenticular, with
subcentric nucleus (Fig. 25F, G).

Head-foot (Fig. 26A): Opaque, with a large
number of white cells scattered throughout.
Propodium wide, with shallow medial inden-
tation. Posteriorly the foot narrowing sharply
to a blunt tip. Attachment thread absent.
Tentacles elongate, subtriangular, connate,
ventro-laterally folded; tentacular pads ab-
sent. Eyes black, subepithelial, round to kid-
ney-shaped, on median side of tentacles.
Mentum incised, with rounded antero-lateral
edges and shallow longitudinal groove (Fig.
11E). Highly coiled visceral mass of grey and
green digestive cells and opaque reproduc-
tive organs.

Alimentary tract (Fig. 21B): When retracted,
introvert-proboscis extending — posteriorly
from its aperture located anterio-medially on
mentum tip to enter cephalic hemocoel. In-
trovert joining buccal sac, which joins a very
short buccal pump divided into anterior (bp1)
and posterior sections (bp2); bp2 four times
length of bp1. Anterior esophagus originating
on ventral surface of alimentary tract at bp1-
bp2 juncture, joining posterior esophagus
and paired salivary glands to form a four-way
junction. Posterior esophagus extending into
visceral mass and joining stomach. Salivary
gland ducts extending anteriorly within walls
of anterior esophagus. At convergence of an-

502 WISE

terior esophagus and buccal pump, ducts
exiting and entering stylet bulb. Salivary
glands attached distally to alimentary tract at
anterior portion of posterior esophagus.

Pallial cavity (Fig. 1C): Mantle and mantle
organs characteristic of subfamily Turbonilli-
nae. Mantle edge etched by fine lines. Mantle
floor and roof darkly pigmented. Ventral and
dorsal ciliated strips joining posteriorly on
mantle roof of mantle cavity. Large, rectan-
gular, oblong pigmented mantle organ com-
posed primarily of clear cells surrounding a
narrow oblong region of bright yellow cells.
Gland beneath ventral ciliated strip extending
posteriorly from mantle edge to convergence
of ciliated strips. Anterior quarter of this gland
composed of large cells containing a thick
yellow exudate, whereas remainder com-
posed of cells filled with a white substance.
Both ventral ciliated strip gland and pig-
mented mantle organ secreting a viscous yel-
low exudate when snail disturbed (Table 5).
Gill absent.

Reproductive system: Typical of pyra-
midellids herein (Fig. 5B). Penis sharing nerve
ring with proboscis (Fig. 22B). Penis anterior,
attenuate, with a deep medial groove and
several lateral rows of minute cuticular hooks
(Fig. 27A), widening posteriorly, becoming
elongate and bulbous. Sperm sac perpendic-
ular to long axis of penis.

Nervous system: Typical of known pyra-
midellids (Fig. 6A).

Ecology and Distribution: Turbonilla hemphilli
occurs subtidally to intertidally in grass/mud
flats from Bermuda to Texas and south to St.
Thomas, West Indies (Abbott, 1974). Host un-
known.

Genus Houbricka new genus

Type species: Turbonilla incisa Bush, 1899,
here designated.

Diagnosis: Lanceolate, white to orange, with
brown band encircling lower portion of each
whorl, 6 mm in length, composed of 7-8
slightly convex whorls. Each whorl with nu-
merous strong axial ribs separated by 6-7
incised spiral shallow grooves between and
perpendicular to intercostal spaces; sculptur-
ing replaced on lower one-third of body whorl
by fine spiral lines. Body whorl 30% of shell
length. Protoconch smooth, sinistrally het-
erostrophic, oriented 90° to teleoconch, sub-
merged 5-10% in adult whorl. Aperture elon-

gate-ovate, with base of outer lip weakly
flared. Columellar folds absent. Operculum
brown, lenticular, with subcentric nucleus.
Head-foot white to reddish pink, with large
white cells dispersed throughout. Anteriorly,
foot flared slightly, not bifurcate. Foot nar-
rowing posterior to propodium, then widening
and ending in blunt apex. Attachment thread
absent. Tentacles elongate, subtriangular,
connate, ventro-laterally folded; tentacular
pads absent. Eyes black, subepithelial, ovate,
on median side of tentacles. Mentum incised,
with rounded lateral edges and shallow lon-
gitudinal groove. Introvert extending posteri-
orly from its medial aperture at anterior men-
tum tip to enter cephalic hemocoel. Introvert
forming an S-shaped loop and continuing
posteriorly to join buccal sac, which is com-
posed of sucker, sheathed stylet, mouth/
stylet aperture, and stylet bulb. Buccal sac
joining а buccal pump, which 1$ divided into
anterior (bp1) and posterior sections (bp2);
laterally flattened bp2 seven times length of
bp1. Anterior esophagus originating at junc-
ture of bp1-bp2, extending posteriorly to join
posterior esophagus and paired salivary
glands, forming a four-way junction. Salivary
gland ducts highly folded, attached to exterior
of anterior esophagus. Ducts extend from an-
terior esophagus to just posterior of buccal
pump-buccal sac juncture, where they detach
and enter stylet bulb. Tuberculate, inflated
posterior esophagus extending posteriorly to
enter visceral mass to join stomach. Salivary
glands attached distally to alimentary tract at
anterior portion of posterior esophagus. Re-
tracted, folded scoop-shape penis within cav-
ity that opens medial and ventral to mentum
outside nerve ring.

Remarks: The new genus introduced here
has been named to honor the late Dr. Richard
S. Houbrick for his very significant contribu-
tions to the science of malacology.

The justification for naming a new genus is
based on anatomical differences between
Houbricka incisa and our current understand-
ing of the genus Turbonilla. Regardless of the
nomenclatural problems outlined above, the
anatomies of certain species within the ge-
nus Turbonilla are well known (e.g., Turbonilla
elegantissima, Т. jeffreysii). When T. hemphilli
is compared to these taxa, this species, and
not H incisa, are very similar. Moreover, the
only taxa that are conchologically similar to
Houbricka incisa, are subgenera (sensu Dall 8
Bartsch, 1909), and their anatomy is un-

PYRAMIDELLID MORPHOLOGY AND PHYLOGENY 503

known. Therefore, until the type species of
these subgenera are examined in greater de-
tail, these names should not be used.

Houbricka incisa (Bush, 1899)

Turbonilla incisa Bush, 1899: 156-157, pl. 8,
fig. 12. (Holotype: ANSP 62800; type lo-
cality: West Florida); Abbott, 1974: 306,
fig. 3781.

Remarks: The holotype is a poor specimen,
which is badly worn, with both aperture and
protoconch broken.

Description: Shell (Fig. 28): Lanceolate, white
to orange, with brown band encircling lower
portion of each whorl, 6 mm in length, com-
posed of 7-8 slightly convex whorls (Fig. 284).
Each whorl with numerous strong axial ribs
separated by 6-7 incised spiral shallow
grooves between and perpendicular to inter-
costal spaces (Fig. 28B), this sculpture re-
placed on lower one-third of body whorl by
fine spiral lines. Body whorl 30% of shell
length. Aperture elongate-ovate, with base of
outer lip weakly flared. Columellar folds ab-
sent. In a few specimens, slight swellings
present on columella deep inside the body
whorl. Protoconch smooth, sinistrally het-
erostrophic, oriented 90° to teleoconch, sub-
merged 5-10% in first adult whorl, with ear-
liest portion of protoconch exposed (Fig. 28C,
E). Operculum brown, lenticular, with subcen-
tric nucleus (Fig. 28F, G).

Head-foot (Fig. 26B): White to reddish pink
(presence of hemoglobin?), with large white
cells dispersed throughout. Anteriorly, foot
flared slightly, not bifurcate. Foot narrowing
posterior to propodium, then widening and
ending in blunt apex. Attachment thread ab-
sent. Tentacles elongate, subtriangular, con-
nate, ventrolaterally folded; tentacular pads
absent. Eyes black, subepithelial, ovate, on
median side of tentacles. Mentum incised,
with rounded lateral edges and shallow lon-
gitudinal groove (Fig. 11F). Visceral mass of
grey, brown, orange or red digestive tissue
cells and opaque to translucent reproductive
organs.

Alimentary tract (Fig. 21C): Retracted intro-
vert-proboscis extending posteriorly from its
medial aperture at anterior mentum tip to en-
{ег cephalic hemocoel. Introvert forming
S-shaped loop, continuing posteriorly to join
buccal sac. Buccal sac joining buccal pump,

which is divided into anterior (bp1) and pos-
terior sections (bp2); laterally flattened bp2
seven times length of bp1. Anterior esopha-
gus originating at bp1-bp2 juncture, extend-
ing posteriorly to join posterior esophagus
and paired salivary glands, forming a four-
way junction. Salivary gland ducts highly
folded and attached to exterior of anterior
esophagus. Ducts extending from anterior
esophagus to just posterior of buccal pump-
buccal sac juncture, where they detach and
enter stylet bulb. Tuberculate, inflated poste-
rior esophagus extending posteriorly to enter
visceral mass and join the stomach. Salivary
glands attached distally to alimentary tract at
anterior portion of posterior esophagus.

Pallial cavity (Fig. 1A, C): Mantle and man-
tle organs only generally like those of the
subfamily Turbonillinae. Mantle edge crenu-
late. Dorsal surface of mantle roof with scat-
tered flecks of black pigment. A line of
subepithelial black cells extending posteri-
orly across the left side of mantle. Ventral and
dorsal strips converge posteriorly on roof of
mantle cavity. Small, oblong pigmented man-
tle organ (similar to the pigmented mantle or-
gan present in the Odostominae) containing
clear cells, cells filled with yellow exudate,
and a few scattered cells containing a red
substance, this organ secreting a thick,
bright yellow exudate when snail disturbed
(Table 5). Gland beneath ventral ciliated strip,
composed of large, black cells within a trans-
parent matrix, extending the length of the
strip. Gill absent.

Reproductive system: Typical of pyra-
midellids in this study (Fig. 5B). Penis outside
and ventral to nerve ring (Fig. 22C). Re-
tracted, folded penis within pocket that
opens medial and ventral to mentum. Paired
retractor muscles anchor penis to pocket
floor. Tapered, dorsoventrally flattened ante-
rior penis tip, scoop-shape with wide, medial
groove (Fig. 27В).

Nervous system: Typical of known pyra-
midellids (Fig. 6A).

Ecology and Distribution: Houbricka incisa
occurs intertidally to subtidally on both the
east and west coasts of southern Florida to
Texas (Abbott, 1972; this study). Host un-
known.

Genus Tathrella Laseron, 1959

Tathrella Laseron, 1959: 218. Type-species:
Tathrella iredalei Laseron, 1959, by orig-
inal designation.

504 WISE

Diagnosis: Shell white, transparent elongate,
conical, 6 mm in length, with 7-8 adult whorls.
Convex whorls with numerous small axial ribs.
Body whorl 40% of shell length. Umbilicus
absent. Protoconch smooth, sinistrally het-
erostrophic, oriented 95° to teleoconch, par-
tially submerged in first adult whorl. Aperture
ovate, with base of outer lip slightly flared.
Columellar folds absent. Head-foot white with
medial black stripe extending from anterior
end of mentum onto mantle floor. Anterior
portion of foot with medial indentation, flared
lateral projections. Foot narrowing posteriorly
to a blunt tip. Pedal gland producing an at-
tachment thread. Tentacles cylindrical, slen-
der, not connate; tentacular pads absent.
Eyes subepithelial, on median side of tenta-
cles. Mentum incised, with rounded, lateral
projections and longitudinal groove. Introvert-
proboscis aperture medial on anterior men-
tum apex. Introvert joining buccal sac, which
is composed of sucker, sheathed stylet,
mouth-stylet aperture, and stylet bulb. Buccal
sac joining buccal pump, which is composed
of two sections (bp1 and bp2). Anterior
esophagus originating at bp1-bp2 juncture,
extending posteriorly and joining posterior
esophagus and paired salivary glands. Sali-
vary gland ducts attached to exterior of an-
terior esophagus. Ducts extending from an-
terior esophagus-buccal pump juncture to
enter stylet bulb. Penis with sperm sac, shar-
ing nerve ring with proboscis.

Remarks: The monotypic genus, originally
described from the shell of a single specimen
was collected in 20 fms. (not 11 fms., as
stated by Laseron, 1959) off Port Curtis,
Queensland, Australia.

Tathrella iredalei Laseron, 1959

Tathrella iredalei Larson, 1959: 218, fig. 101.
(Holotype: AMS 105285, type locality:
Port Curtis, Gladestone, Queensland,
Australia).

Pyrgiscus sp. Cumming, 1988.

Turbonilla sp. Cumming, 1993.

Description: Shell (Fig. 29): Thin, chalky
white, translucent, 5-6 mm in length, com-
posed of 7-9 convex whorls. Body whorl 40%
of shell length (Fig. 29A). Adult whorls with a
number of slender axial ribs and microscopic
orthocline growth lines. Aperture elongate-
ovate, with outer lip flared at base (Fig. 29B).

Columella thick, without columellar folds. In
some specimens, 1-4 swellings present deep
within body whorl on columella (Fig. 290).
Protoconch smooth, sinistrally hetero-
strophic, oriented 95° to teleoconch axis,
submerged 10-15% in first adult whorl, with
earliest portion of protoconch partially ex-
posed (Fig. 29D, F). Operculum brown, len-
ticular, with subcentric nucleus (Figs. 29G, H).

Head-foot (Fig. 26C): White with prominent
black stripe extending from anterior mentum
tip across the head and, in some individuals,
onto mantle floor. Propodium with moderate
medial indentation and flared lateral projec-
tions. Foot narrowing posterior to propo-
dium, then widening and terminating in blunt
tip. Black lines of pigment of varying length
and definition typically on dorso-lateral sur-
face of foot. Opening to pedal gland a slit on
postero-ventral surface of foot extending one
fourth of foot's length anteriorly from poste-
rior end. Pedal gland producing an attach-
ment thread. Tentacles cylindrical, long, not
connate; tentacular pads absent. Eyes black,
subepithelial, on median side of tentacles.
Mentum incised, long, with knob-like antero-
lateral projections (Fig. 11G). Short, corpulent
visceral coil of light orange, gray, or creamy
white digestive tissue cells and opaque re-
productive organs.

Alimentary tract (Fig. 21D): When ге-
tracted, introvert/proboscis extending poste-
riorly from its medial aperture on anterior
mentum tip to enter cephalic hemocoel and
join buccal sac. Buccal sac connecting to a
moderately short buccal pump. Buccal pump
divided into anterior (bp1) and posterior sec-
tions (bp2); bp2 five times length of bp1.
Short anterior esophagus joining buccal
pump just posterior of buccal sac-buccal
pump juncture. Anterior esophagus continu-
ing posteriorly to join posterior esophagus
and salivary glands to form a four-way junc-
tion. Long, highly coiled posterior esophagus
extending to enter visceral mass and termi-
nate at stomach. Salivary gland ducts at-
tached to outer surface of anterior esopha-
gus. At convergence of anterior esophagus
and buccal pump, salivary ducts extend an-
teriorly to enter stylet bulb. Salivary glands
attached distally to alimentary tract at ante-
rior portion of posterior esophagus.

Pallial cavity (Figs. 1A, C): Mantle and man-
tle organs generally as described for the Tur-
bonillinae. Mantle edge finely scalloped. Ven-
tral and dorsal ciliated strips join posteriorly
on mantle roof of mantle cavity; strips bi-

PYRAMIDELLID MORPHOLOGY AND PHYLOGENY 505

sected by a thin line of brown, glandular cells
that extend the length of mantle cavity.
Small, oblong pigmented mantle organ (sim-
ilar to same organ in Odostominae) com-
posed primarily of cells containing a yellow
exudate and a few cells filled with white con-
tents. Gland, beneath ventral ciliated strip ex-
tending 50-60% of strip’s length. Gland filled
with cells containing a bright yellow exudate.
Snail exuding copious amounts of exudate
from ventral ciliated strip gland and a lesser
amount from pigmented mantle organ when
disturbed (Table 5). Gill absent.

Reproductive system: Typical of pyra-
midellids examined this study (Fig. 5B). Penis
sharing nerve ring with proboscis and when
protracted, extending through a medial
opening ventral to mentum (Fig. 22D). Poste-
riorly, penis continuing into cephalic hemo-
coel, looping once, and attaching ventrally
to columellar muscle. Anterior of penis medi-
ally grooved and ciliated. A slender duct
leading distally to bulbous sperm sac present
perpendicular to anterior portion of penis
sheath.

Nervous system: Characteristic of known
pyramidellids (Fig. 6A).

Ecology and Distribution: Tathrella iredalei,
which parasitizes various Tridacna species,
has inadvertently been sent to mariculture fa-
cilities throughout the Indo-Pacific (e.g., to
Australia, Philippines, Guam, and the So-
lomon Islands) in shipments of Tridacna. As a
result, this snail has been introduced into ar-
eas where it does not naturally occur, making
its original geographic distribution difficult to
determine.

ACKNOWLEDGEMENTS

This research is dedicated to my co-adviser
Dr. Richard S. Houbrick, who died in August
1993. | am particularly grateful to my other
co-advisor, Dr. Diana Lipscomb, who willingly
took on the responsibilities involved in seeing
this work completed. | also wish to thank my
committee members Dr. Jaren Horsley and
Dr. David Atkins, and the outside readers Dr.
Randall Packard, Dr. John Burns and Dr. Rob-
ert Hershler for their help in this endeavor.

| gratefully acknowledge financial support
from Sigma X, Lerner-Grey Fund, National
Capitol Shell Club, Delaware Museum,

George Washington University, Smithsonian
Institution's Predoctoral Fellowship and Re-
search Award at Smithsonian Marine Station
at Linkport.

Much of this work was accomplished at the
Smithsonian Marine Station, Ft. Pierce, Flor-
ida, and this paper is Smithsonian Marine Sta-
tion Contribution no. 380. | am indebted to
Julie Piraino, Sherry Reed, Joan Kaminski,
Woody Lee, Hugh Reichardt, and Dr. Mary
Rice for all their time and help.

At the National Museum of Natural History
Smithsonian Institution, | thank Victor Kranz
for photographic assistance; Suzanne Bran-
den, Walt Brown, and Peter Viola Scanning
Electron Microscopy Laboratory, for all their
insightful lessons and assistance; Molly Ryan
for her help with the POS machine and advice
in producing line drawings; Cathy Price,
Cheryl Bright, and especially Beth Fricano, for
their help with specimen preparation and his-
tology; Dr. Gerry Harasewych for his time and
valuable aid; Ron Larson, Smithsonian Librar-
ies, for his assistance in the library, and Marty
Joynt for being “Marty”.

| want to thank Dave Hopper, Alex Kerr, Dr.
José Leal, Dr. Paula Mikkelsen, Dr. Jim
Mclean, Frank Te, and Kati, Robert, and Rick
Wise for invaluable field assistance. | also
want to thank Bern Holthuis, Dr. Gustav
Paulay, Brad Peterson, Kati Wise, and Robert
Wise for collecting and sending living speci-
mens. The University of Guam Marine Station,
Kewalo Marine Laboratory and Pacific Bio-
medical Research Center, University of Ha-
waii, and the Los Angeles County Museum of
Natural History provided laboratory space. |
thank Clif Coney and Drs. Jim Mclean, Lou
Elridge, Mike Hadfield, and Drs. Petra and
Rudiger Bieler for their generous hospitality.

| thank Dr. Paula Mikkelsen for all her help
in Florida, Dick Petit and Dr. Eugene Coan for
the much needed nomenclatural assistance,
Dr. José Leal for his darkroom expertise and
nomenclatural help, Mr. Wybou of Ft. Pierce,
Florida for translating several articles, Dr. Sil-
vard Kool and Dr. Winston Ponder AMNH for
sending museum specimens and Rick Wise
for all the thought provoking discussions and
software help.

The critical comments of Dr. Winston Pon-
der, Mr. Richard Petit, Dr. Eugene V. Coan,
and an anonymous reviewer greatly improved
this manuscript.

Last, but not least, | want to gratefully ac-
knowledge the fantastic support | received
from the entire Wise family.

506 WISE

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Revised Ms. accepted 30 August 1995

MALACOLOGIA, 1996, 37(1-2): 513-536

INDEX

Taxa in bold are new; page numbers in
bold are pages where new taxa are

introduced or described. Page numbers in

italics indicate illustrations of taxa.

abbreviatus, Pythia 228, 230
abrupta, Turbonilla 501
Achatina pellucida 187, 189
acinoides, Melampus 314
Acochlidiacea 404
Acroloxus 171
Acteocina 376, 377, 379, 381, 382,
384, 385, 389, 390, 395, 397, 399,
402, 404, 405, 409, 412, 413, 415-
417, 419-421, 424, 436
Acteocina atrata 379, 436
bidentata 436
canaliculata 379, 393, 436
candei 436
Acteocinidae 376, 421
Acteon 311, 376, 377, 379, 381-383,
384, 385, 387, 390, 394-396, 399,
402, 404, 407, 409, 410, 415-422,
432
candens 432
denticulatus 194
finlayi 432
heteroclita 187
pelecais 432
tornatilis 388, 403, 407, 432
traskii 432
Acteonidae 376,421
acuminata, Alexia 196, 200, 203
acutidentata, Daecalochila 91
acutidentata, Daedalochila (Upsilodon)
91, 94, 96
adamnis, Linisa 90
Adelopneumona 193
Ademnestia arachis 402
adriatica, Auricula myosotis var. 194
aequalis, Melampus 248
aequalis, Monica 226
aequalis, Ovatella 193, 209, 210, 226,
230 332
aequalis, Pythia 230
Aequipecten qibbus 459
afra, Helix 213
afra, Pedipes 212,217
Akera 376, 381-383, 384, 385, 390-
393, 395, 396, 399, 400, 409, 410,
412, 416, 419-421, 440-441
bayeri 440
bullata 385, 392, 395, 440
Akeridae 376, 381, 421
alabamensis, Triodopsis (Triodopsis) 86
alba, Cylichna 435
alba, Microtralia 236
albicostulata, Linisa 90

albolabris, Helix 78

albolabris, Neohelix 80, 104
albolabris, Neohelix (Neohelix) 80, 95
albus, Gyraulus 19,64

albus, Melampus flavus 293

albus, Melampus flavus var. 291

Alexia 177, 191-194

acuminata 196, 200, 203
algerica 195, 197
armoricana 196, 200, 202
bermudensis 195, 197, 202
biasoletina 195
biasolettiana 195
bicolor 196
bidentata 196
borealis 195
ciliata 195
cossoni 196, 200
denticulata 192,195
exilis 196, 200
globulus 196, 200
hiriartí 195
letourneuxi 196, 200, 202
loweana 195
micheli 195, 197
micheli var. triplicata 197, 197
myosotis 195, 267
myosotis bermudensis 196
myosotis marylandica 196, 200, 202
myosotis myosotis 196
myosotis var. varicosa 196
obsoleta 195
oranica 196, 202
parva 196, 200
payraudeaui 195
pechaudi 196, 200
pulchella 196, 200
ringicula 196, 200, 202
setifer 195, 196, 197
setifer var. tenuis 195, 197
setifera 195
setigera 195
subflava 196, 202
terrestris 196, 200
vespertina 196
Alexia (Auricula) myosotis var. hiriarti
195, 197
Alexia (Kochia) denticulata 193
oranica 192, 193, 196, 200
Alexia (Myosotella) myosotis 196
algerica, Alexia 195, 197
algerica var. quadriplicata, Alexia 195,
197
aliciae, Euchemotrema monodon 79
alleni, Neohelix 80
alleni, Neohelix (Solemorbis) 85
Allochroa 314
bronni 175

513

514

Allogastropoda 310, 376, 444
Allogona 84, 87
profunda 71, 101, 104
Allogona (Allogona) 83, 87
profunda 87, 94, 99
Allogona (Dysmedoma) 83, 84, 87
lombardii 87
ptychophora 87
townsendiana 87, 94
Allogonini 84, 87
alternatus, Melampus coffeus var. 263,
265
altispira, Stenotrema altispira 79
altispira, Stenotrema (Stenotrema) 89
altissima, Ashmunella 87
altivagus, Mesodon (Akromesodon) 93
amanda, Cernuella 350, 352-356, 357
amanda, Helix 349, 350, 356
Amanthina 454
Amanthinidae 446, 454, 481-487
Amphibolidae 311
ampla, Helix 90
ampla, Praticolella (Praticolella) 90, 95
ampulla, Bulla 392, 436
Anaspidea 379, 412, 416, 418-420,
422, 424
Ancillus 193
Ancilus 193
andrewsae, Mesodon 104
andrewsae, Mesodon (Akromesodon) 93
Androgyna 311
angasi, Philine 397, 434
angellum, Stenotrema (Stenotrema) 89
angulata, Ashmunella 93, 96
anilis, Helix (Polygyra) 89
anilis, Linisa 90, 94
animasensis, Ashmunella 87
Anisus rotundatus 62
anteridon, Triodopsis (Triodopsis) 86
antillarum, Haminoea 388, 392, 395,
398, 400, 419, 420, 437
aperta, Philine 377, 389, 390, 393, 397,
399, 408, 420, 434, 435
Aphalista 496
mitralis 499
Aplustridae 376, 386, 421
Aplysia 376-378, 381, 382, 384, 386,
391-395, 399, 400, 404, 409, 410,
412,416, 418, 419, 421, 441
brasiliana 392, 441
californica 395, 441
dacylomela 441
depilans 441
fasciata 441
grandis 441
juliana 441
parvula 441
puncatata 441
Aplysiidae 418, 421
Aplysiomorpha 453, 454
Apodosis 174,211, 237
novimundi 237, 238, 241, 242

INDEX

Apogastropoda 171

Appalachina 83,84, 92, 93, 106
chilhoweensis 93
sayana 71, 94, 99, 101, 104
sayanus 93

appressa, Helix 92

appressa, Patera 104

appressa, Patera (Patera) 92, 95

approximans, Inflectarius (Inflectarius) 92

arachis, Ademnestia 402

Arbobuccinum argus 23

Archaepulmonata 311, 418

Archerelix 88 [see Stenotrema
(Archerelix)]

archeri, Еитопейх 72, 92, 104

Architectibranchia 416, 454

Architectonicidae 376, 377, 404, 421

Architectonicoidea 417, 444

Arctica islandica 371

Arctidae 364

arenosa, Obeliscus 498

arenosa, Xeropicta 6,9

argus, Arbobuccinum 23

ariadne, Daedalochila 91

ariadne, Daedalochila (Daedalochila) 91

Aricula gundlachi 263

Armiger crista 64
rotundatus 64

armigera, Vespericola 86

armoricana, Alexia 196, 200, 202

Asamiorbis 78 [see Neohelix
(Asamiorbis)]

Ascobulla 376, 383, 384, 390, 395,
398-400, 402, 407-410, 412, 416,
419, 439-440

Ascobulla ulla 385, 388, 405, 439

Ascobullidae 376

Ashmunella 83, 84, 87, 106, 107
altissima 87
angulata 93, 96
animasensis 87
ashmuni 87
bequaerti 87
binneyi 87
carlbadensis 87
chiricahuana 88
cockerelli 88
danielsi 88
edithae 88
ези ог 88
ferrissi 88
harrisi 88
hawleyi 88
hebardi 88
intricata 88
jamesensis 88
juanezensis 88
kochi 88
lenticula 88
lepidoderma 88
levettei 88
macromphala 88

mearnsi 88
mendax 88
meridionalis 88
mogollonensis 88
montivaga 88
mudgei 88
mudgi 72
organensis 88
pasonis 88
pilsbryana 88
proxima 88
pseudodonta 88
rhyssa 88, 94, 96
rhyssa miorhyssa 87
rileyensis 88
ruidosana 88
salinasensis 88
sprouli 88
tegillum 88
tetrodon 88
thomsoniana 88
tularosana 88
varicifera 88
watleyi 88
Ashmunellini 84, 87
ashmuni, Ashmunella 87
aspera, Patella 33-40
Astraea gibberosa 491
undosa 491
atrata, Acteocina 379, 436
Auicula bicolor 202
aulacomphala, Linisa 90
auricella, Auricula 183, 184

auricella, Auricula (Auricella) 179

Auricella 179
auricula, Bulimus 179, 183
auricula, Ellobium 183

Auricula 164, 174, 177, 178, 193, 231,

236, 247, 248, 302
auricella 183, 184
biasolettiana 194

bicolor 195, 197

bicolor var. subarmata 195
bidenta 263

bidentata 263

biplicata 245, 249, 258
botteriana 195, 202
ciliata 195, 197, 197, 203
cingulata 276

coniformis 249

conoidalis 249

cornea 262, 266, 269
denticulata 194

denticulata var. borealis 194

dominicense 178, 180
dominicensis 180, 185
floridana 285

globulus 287
heteroclita 187
infrequens 202

jaumei 263, 268, 269
kuesteri 275

INDEX

kutschigiana 194
microstoma 194
midae 178
monile 290
multivolvis 276, 279, 308
myosotis 192, 194, 201
myosotis var. adriatica 194
myosotis var. elongata 194
mysotis 194
nitens 303
obliqua 263
oliva 276, 279
olivula 249, 258
ovula 303
pellucens 180, 184
pusilla 303
reflexilabris 194, 197, 203
ringens 433
rugeli 285
sayi 202, 203
striata 205
tenella 194, 202
veneta 194
vespertina 195, 197
watsoni 195, 197, 202
watsoni scrobiculata 195, 197, 202
Auricula (Alexia) denticulata 195
meridionalis 195, 197, 203
myosotis 196
Auricula (Auricella) auricella 179
Auricula (Auricula) myosotis 194
Auricula (Auriculastrum) pellucens 180
Auricula (Auriculina) gangetica 179
Аипсиа (Conovulus) coniformis 249
ovula 303
pusillus 303
triplicatus 308
Auricula (Microtralia) minuscula 231,
232723532
Auriculacea 173, 174
Auriculaceae 173
Auriculadae 173
Auriculae 173
Auriculae'inae 173
Auriculastra 170, 184, 314
elongata 184
папа 232, 236
subula 175, 184
Auriculastrum 184
pellucens 180
Аипсшаз ит (Microtralia) minusculum
232
auriculata, Daedalochila (Daedalochila)
91,94
auriculata, Helix 91
auriculata, Polygyra 91
Auriculata 173
Auriculea 176
Auriculiadae 173
Auriculidae 173, 190
Auriculidea 173
Auriculina 173

515

516

Auriculina 179
Auriculinae 176, 177, 186
Auriculinella 168, 176, 177, 186, 228,
235, 312, 314
bidentata 164, 332
Auriculinella (Leucophytia) bidentata 175,
176; 202267
Auriculodes 178-179, 312
Auriculoidea 173
Auriculus 178
judae 178
pellucens 180
auriformis, Daedalochila (Daedalochila) 91
auris-felis, Bulimus 256
aurisfelis, Cassidula 192, 256, 257
aurisfelis, Cassidula (Cassidula) 175
aurisjudae, Ellobium (Ellobium) 170, 178,
179, 185, 192
aurismidae, Bulla 178
aurismidae, Ellobium (Ellobium) 175,177,
178 1, ЛОР. WSS, IA, Sey
australis, Ophicardelus 175, 225
Autonoe 178, 179
riparia 178, 180
Autonoella 179
riparia 179
avara, Daedalochila (Daedalochila) 91

B. fruticum 9

babylonia, Chemnitzia 491

babylonia, Cingulina 491

babylonia, Liamorpha 491

babylonia, Odostomia 445, 446, 450,
457, 459, 467, 463, 464, 466, 481-
487, 491-492

babylonica, Odostomia (Chrysallida) 491

babylonica, Odostomia (Cingulina) 491

bakeri, Praticolella (Filapex) 90

barbadense, Ellobium 257

barbatum, Stenotrema 79

barbatum, Stenotrema (Toxotrema) 89

barbigera, Helix 88

barbigerum, Stenotrema 79

barbigerum, Stenotrema (Archerelix) 88,
93, 106

Basommatophora 172, 311

Bathyomphalus contortus 64, 65

bayeri, Akera 440

Беаий, Cyclostremiscus 405, 408, 416

beauii, Cylindrobulla 439

behri, Linisa 90, 94, 96

Bellamya unicolor 50

bequaerti, Ashmunella 87

berlandieriana, Praticolella 81

berlandieriana, Praticolella (Praticolella)
90

bermudae, Volvatella 398-400, 440

bermudensis, Alexia 195, 197, 202

bermudensis, Alexia myosotis 196

bermudensis, Ovatella myostis 197

bermudensis, Phytia 196

bermudensis, Phytia myosotis var. 196

INDEX

biasoletina, Alexia 195
biasolettiana, Alexia 195
biasolettiana, Auricula 194
bicolor, Alexia 196
bicolor, Auicula 202
bicolor, Auricula 195, 197
bicruris, Linisa 90
bidenta, Auricula 263
bidentata, Acteocina 436
bidentata, Alexia 196
bidentata, Auricula 263
bidentata, Auriculinella 164, 332
bidentata, Auriculinella (Leucophytia)
175; 176, 202, 267
bidentatus, Melampus 203, 262, 266,
329
bidentatus, Melampus bidentatus 263
bidentatus, Melampus (Leuconia) 202,
267
bidentatus, Melampus (Melampus) 164,
165, 168, 170, 1731762202203;
246, 248, 249, 257, 258, 262-275,
264-270, 284, 285, 295, 300, 308,
314
bidentus, Melampus 263
binneyana, Patera (Vesperpatera) 92, 95
binneyana, Polygyra 92
binneyi, Ashmunella 87
Biomphalaria 19
camerunensis 14
glabrata 14
pfeifferi 18
biplicata, Auricula 245, 249, 258
biplicatus, Melampus 250
biplicatus, Melampus (Conovulus) 249
bishopi, Melampus coffeus var. 263, 265
bisuturalis, Boonea 488
Bithynia graeca 50
tentaculata 19, 50, 65
blandianum, Stenotrema 79
blandianum, Stenotrema (Pilsbrelix) 89
Blannaria 186
pellucida 187
Blaumeria 186
heteroclita 187
Blauneria 168, 173, 176, 177, 186, 191,
210, 228, 230, 2364312
cubensis 187
elliptiformis 186
gracilis 186
heteroclita 164, 170, 172, 175, 176,
186-191, 188, 189, 236, 330, 332
pellucida 187
boholensis, Melampus 164, 258
Boonea 449, 551, 488, 490
bisuturalis 488
cincta 445, 446, 450, 457, 459, 461,
489-491
impressa 488
seminuda 445, 446, 450, 457, 459,
460, 461, 463-465, 481-490
borealis, Alexia 195

INDEX

borealis, Auricula denticulata var. 194
borealis, Melampus 194, 203, 268
borealis, Melampus bidentatus var.
borealis, Phytia myosotis 196
botteriana, Auricula 195, 202
Bradybaena 84
similaris 95
Branchiopulmonata 311
brasiliana, Aplysia 392, 441
brevicula, Littorina 130
brevipila, Stenotrema 79
brevipila, Stenotrema (Stenotrema) 89
brevispira, Marinula 226
bronni, Allochroa 175
Brooksorbis 85 [see Triodopsis
(Brooksorbis)]
buccinea, Ringicula 377, 387, 388, 405,
432
bulimoides, Cleopatra 50
bulimoides, Melampus 277
Bulimus 247
auricula 179, 183
auris-felis 256
coniformis 249, 257
monile 290, 293
ovulus 303, 308
pedipes 213
Bulinus 19
truncatus 17
bulla, Melampus 276
bulla, Melampus (Melampus) 276
Bulla 379, 381-383, 384, 385-387, 393,
395, 399, 400, 402, 404, 405, 409,
410, 412, 416, 418-422, 424, 433,
436-437
ampulla 392,436
aurismidae 178
coffea 249, 256
gouldiana 405, 420, 436
solida 436
striata 387,388, 390, 409, 420, 436
bullaeoides, Detracia 275-277
bullaeoides, Melampus (Detracia) 277
bullaoides, Conovolus 277
bullaoides, Detracia 277
bullaoides, Melampus 275, 277
bullaoides, Melampus (Detracia) 169,
173, 175, 275, 276-284, 279-282,
295, 300, 308; 332
bullaoides, Tornatella 276
bullaoides, Voluta 276
Bullapex 204
Bullaria 418
bullata, Akera 385,392, 395, 440
Bullidae 421
Bulloidea 376, 412, 416, 417, 424
bulloides, Detracia 277
bulloides, Melampus (Detracia) 277
Bullomorpha 453, 454
burchi, Triodopsis 85
burchi, Triodopsis (Pilsbryorbis) 85
burlesoni, Daedalochila 81

263

DUT

burlesoni, Daedalochila (Upsilodon) 91
Bursa 23
crenulata 31
Bursatella 379
Bursidae 23
Busycon 419

caddoense, Stenotrema 79
caddoense, Stenotrema (Stenotrema) 89
Caenogastropoda 376
caerulea, Patella 33-40
caffeus, Melampus 250
californica, Aplysia 395, 441
caloosaensis, Polygyra 89
calvescens, Stenotrema (Stenotrema) 89
calyculata, Smaragdinella 437
Calyptogena 363-372
magnifica 363, 364, 365, 366-371,
37105372
pacifica 365, 369, 369-370, 371, 372
phaseoliformis 365, 367, 368-370,
371,312
Camaenidae 95, 106
camerunensis, Biomphalaria 14
canaliculata, Acteocina 379, 393, 436
Canariella 158
discobolus 159
eutropis 159
gomerae 159
hispidula 159
leprosa 159
р/апапа 159
candei, Acteocina 436
candens, Acteon 432
candida, Praticolella (Praticolella) 90
cantralli, Linisa 90
Caracolla limbata 350, 352, 353, 355,
356
Carditidae 363, 364
carinata, Leptoxis 50
carinatus, Planorbis 66
carlbadensis, Ashmunella 87
carolinensis, Xolotrema (Xolotrema) 85
cartusiana, M. 9
caruanae, Cernuella 350, 357
Carychiidae 173
Carychiinae 174, 775, 313, 315
Carychiopsis 314
Carychium 176, 177, 193, 209, 312,
314
minimum 175
personatum 194, 197, 201, 202
tridentatum 175, 193, 209, 313, 331
Cassidaria rugosa 23, 31
Cassidula 170, 176, 177, 191, 192;
205, 256, 302, 312-315
aurisfelis 192, 256, 257
labrella 193,209, 259
mustelina 332
Cassidula (Cassidula) aurisfelis 175
doliolum 175
Cassidulinae 174, 186, 191, 211, 232

518

Cassis tuberosa 23
catascopium, Lymnaea 62, 64, 65
cepa, Cepolis 95
Cepaea hortensis 111
normalis 111-122, 145
Cephalaspidea 375-442
Cepolis 72, 84, 96
cepa 95
cereolus, Polygyra 81,89
Cernuella 349, 350, 353, 356-359
amanda 350, 352-356, 357
caruanae 350, 357
rugosa 350
virgata 6, 9
Cernuellopsis 356, 359
ghisotti 359
ceylanicum, Ellobium 184
Chemnitzia babylonia 491
seminuda 488
chilhoweensis, Appalachina 93
chilhoweensis, Mesodon 104, 105
chinense, Ellobium (Ellobium) 179
chiricahuana, Ashmunella 88
chisosensis, Daedalochila 81
chisosensis, Daedalochila (Upsilodon) 91
Chrysallida 450, 451, 457, 490
cincta 445, 450, 457, 481-487, 489,
490
torita 490
chrysoma, Retusa 382, 438
ciliata, Alexia 195
ciliata, Auricula 195, 197, 197, 203
cincta, Boonea 445, 446, 450, 457, 459,
461, 489-491
cincta, Chrysallida 445, 450, 457, 481-
487, 489, 490
cincta, Odostomia (Chrysallida) 489
cincta, Volvatella 440
cingulata, Auricula 276
cingulata, Melampus (Tifata) 277
cingulata, Tralia 275, 277
cingulatus, Melampus 276
cingulatus, Melampus (Tralia) 277
Cingulina babylonia 491
judithae 491
claibornensis, Triodopsis (Shelfordorbis)

clarki, Detracia 297, 299

clarki, Melampus 297

clarki, Patera (Patera) 92

clarkii, Melampus 297

Clathrella 454

clausa, Laemodonta 206
clausa, Plecotrema 209
Clauselia 193

clausus, Mesodon (Mesodon) 93
clavus, Scaphander 433
clenchi, Patera (Vesperpatera) 92
Cleopatra bulimoides 50
cockerelli, Ashmunella 88
coffea, Bulla 249, 256

coffea, Conovulus 249

INDEX

coffea, Melampus 249
coffea, Melampus (Melampus) 250
coffea, Voluta 249, 256, 257
coffee, Conovulus 249
coffeus, Melampus 247, 249, 258, 329,
331
coffeus, Melampus coffeus 250
coffeus, Melampus (Conovulus) 249
coffeus, Melampus (Melampus) 164,
168, 770, 170, 172, 173; 07348237
176, 245, 247, 249-262, 251-256,
265, 266, 268, 290, 29472957300;
3087332
cohuttense, Stenotrema 79
cohuttensis, Polygyra 88
cohuttensis, Stenotrema (Cohutta) 88, 95
columbiana, Polygyra 86
columbiana, Vespericola 95
columbianus, Vespericola 86
complanata, Triodopsis (Pilsbryorbis) 85
concinna, Marinula 226
coniformis, Auricula 249
coniformis, Auricula (Conovulus) 249
coniformis, Bulimus 249, 257
coniformis, Conovulus 249
coniformis, Melampus 247, 249
coniformis, Pedipes 249
coniformis, Ringicula 377, 387, 388,
405, 432
conoidalis, Auricula 249
Conovolus 247
bullaoides 277
Conovula 247
Conovulae 247
Conovulidae 173
Conovulum 247
Conovulus 193, 213, 247, 248
Conovulus coffea 249
coffee 249
coniformis 249
denticulatus 195
monile 290
pellucens 180
Conovulus (Alexia) denticulata 195
contectus, Viviparus 41-52
contortus, Bathyomphalus 64, 65
contrerasi, Odostomia (Chrysallida) 490
convexa, Stenotrema 88
cookiana, Sagda 95
copei, Triodopsis 86
cornea, Auricula 262, 266, 269
cornea, Cremnobates 225
corneliae, Giffordius 89
corneus, Melampus 263
corneus, Melampus bidentatus 263
corneus, Melampus (Melampus) 263
corneus, Planorbarius 14, 53-68
Cornirostridae 377
coronata, Runcina 401
coronatus, Melampus 290, 293
coronulus, Melampus 291
cossoni, Alexia 196, 200

costata, Tralia (Persa) 175

costulata, Turbonilla 501

couloni, Linisa 90

cragini, Triodopsis 86

cragini, Triodopsis (Haroldorbis) 95

Creedonia 169, 172, 173, 175, 176,
190, 210-213, 225-226, 236, 238,
240, 312-314
succinea 173, 175, 202, 203, 225-

INDEX 519

Cylindrobulla 376, 377, 381, 383, 384,
384, 385, 386, 387, 390, 395, 398-
400, 407, 402, 403, 406, 407-410,
412, 416, 419, 420, 439
beauli 439

Cylindrobullidae 376

Cylindrotis 170, 186, 191, 314
quadrasi 175

Сутанит 23

231, 227,229, 230, 236, 330-332

Cremnobates 225, 312
cornea 225
parva 225, 226
solida 225

crenulata, Bursa 31

crenulata, Pyramidella 445, 446, 450,
458, 464, 472, 474, 475, 481-487,
498-499

dacylomela, Aplysia 441
Daedalochila 81, 84, 89-91, 106

acutidentata 91

arladne 91

burlesoni 81

chisosensis 81

hippocrepis 81

leporina 81

crenulata, Pyramidella (Longchaeus) 498 multiplicata 81

Crepidula 381
fornicata 445, 459
crista, Armiger 64
crosseana, Melampus (Sayella) 495

plicata 91

uvulifera 81
Daedalochila (Acutidens) 91
Daedalochila (Daedalocheila) 83

crosseana, Petitella 445, 446, 450, 457- Daedalochila (Daedalochila) 91

459, 464, 465, 469, 470, 481-487,
495-496
crosseana, Sayella 495
Cryptomastix 84, 86, 87
Cryptomastix (Bupigona) 83,84, 87
hendersoni 94
Сгурютазих (Cryptomastix) 83, 87
devia 87
mullani 87, 94
sanburni 87
Cryptomastix (Micranepsia) 83, 84, 87
germana 87, 94
cubense, Plecotrema 206
cubensis, Blauneria 187
cubensis, Laemodonta 173, 193, 205-
210, 207-209, 236, 241, 330, 332
cubensis, Laemodonta (Bullapex) 204,
206
cubensis, Odostomia 187, 190
cubensis, Odostomia (Tornatellina) 187
cubensis, Oleacina (Stobilus) 187
cubensis, Physella 333-348
cubensis, Plecotrema 206
cubensis, Tornatellina 187, 189
Cyclostremellinae 444
Cyclostremiscus Беаий 405, 408, 416

arladne 91
auriculata 91,94
auriformis 91
avara 91
delecta 91
hausmani 91
oppilata 91
peninsulae 92
postelliana 92
subclausa 92
uvulifera 72, 92, 107
Daedalochila (Upsilodon) 81, 83, 84, 91
acutidentata 91, 94, 96
burlesoni 91
chisosensis 91
dalli 91
hippocrepis 72, 91, 93, 94, 96, 107
leporina 91
multiplicata 91
poeyi 91
sterni 91
dalli, Daedalochila (Upsilodon) 91
danielsi, Ashmunella 88
deceptrix, Odostomia (Chrysallida) 490
deceptum, Stenotrema 79
deceptum, Stenotrema (Pilsbrelix) 89

Cylichna 376, 377, 384, 385, 390, 395- delecta, Daedalochila (Daedalochila) 91

397, 402, 407-409, 412, 413, 416,
417, 424, 435-436
alba 435
cylindracea 390, 406, 407, 434
magna 435
verrilli 435
Cylichnatys 418
Cylichnidae 376, 402, 421
cylindracea, Cylichna 390, 406, 407,
434
cylindrellus, Scaphander 390

deltoidea, Millerelix (Prattelix) 91
denotata, Xolotrema 104

denotata, Xolotrema (Xolotrema) 85, 95
denticulata, Alexia 192, 195
denticulata, Alexia (Kochia) 193
denticulata, Auricula 194
denticulata, Auricula (Alexia) 195
denticulata, Conovulus (Alexia) 195
denticulata, Jaminia 192, 194
denticulata, Ovatella 202
denticulata, Philine 434

520

denticulata, Phytia 194, 201, 202
denticulata, Phytia myosotis 196
denticulata, Pythia 194
denticulata, Voluta 169, 193, 194, 797,
201
denticulatus, Acteon 194
denticulatus, Conovulus 195
denticulatus, Melampus 195
dentifera, Helix 78
dentifera, Neohelix 106
dentifera, Neohelix (Asamiorbis) 94
depilans, Aplysia 441
depilatum, Stenotrema altispira 79
depilatum, Stenotrema (Stenotrema)
altispira 89
Detracia 169, 173, 248, 275-276, 295,
3027 312
bullaeoides 275-277
bullaoides 277
bulloides 277
clarki 297, 299
floridana 284, 285
parana 287, 289
roquesana 277, 279, 281
devia, Cryptomastix (Cryptomastix) 87
didyma, Odostomia 445, 446, 450, 459,
461, 463, 464, 467, 481-487, 492-
493
didyma, Odostomia (Cyclodostomia) 492
discobolus, Canariella 159
discoidea, Triodopsis (Vagvolgyrorbis) 86
dissecta, Linisa 90
Distorsio perdistorta 23
doerfeuilliana, Millerelix 81, 91
dolabratus, Trochus 496
doliolum, Cassidula (Cassidula) 175
dominicense, Auricula 178, 180
dominicense, Ellobium 330
dominicense, Ellobium (Auriculodes) 170,
172,176, 179, 180-186, 181-184,
332
dominicensis, Auricula 180, 185
Doridella steinbergae 420
downieanus, Inflectarius (Hubrichtius) 92,
94
duryi, Helisoma 14,17
dysoni, Linisa 90

eburnea, Wingenella 497

edentatus, Inflectarius (Inflectarius) 92
edgarianum, Stenotrema 79
edgarianum, Stenotrema (Archerelix) 88
edithae, Ashmunella 88

edulis, Mytilus 124, 129, 130
edvardsi, Stenotrema 79

edvardsi, Stenotrema (Archerelix) 88
eiseni, Tegula 491

eiseni, Tequla 445

elegans, Haminoea 398, 419, 437
elegantissima, Turbo 501
elegantissima, Turbonilla 450, 451, 502
Eleutherobranchia 418

INDEX

elevatus, Mesodon (Aphalogona) 93, 94
elliptiformis, Blauneria 186
Ellobiidae 163-332
Ellobiinae 168-170, 172, 174, 175, 176,
177, 186, 191, 211, 231 28522740)
30247313
Ellobium 164, 170, 173, 174, 176-186,
231, 236, 247, 3027312 731478115
auricula 183
barbadense 257
ceylanicum 184
dominicense 330
inflammatum 257
midae 178
pellucens 180
Ellobium (Auriculodes) dominicense 170,
172, 176, 179, 180-186, 181-184,
332
дапдейсит 175
gaziense 179, 181, 184
pellucens 183
stagnale 179
Ellobium (Ellobium) aurisjudae 170, 178,
179,185, 192
aurismidae 175,177,178, 181, 182,
185, 192332
chinense 179
subnodosum 179
elodes, Lymnaea 62, 65, 345
elongata, Auricula myosotis var.
elongata, Auriculastra 184
elongatus, Pedipes 226, 227, 228
Elysia 379, 402
timida 409
viridis 400
Ensiphorus 275
longidens 276
Enterodonta 205
Eobania vermiculata 2, 9
Ercolania 402
lozanoi 400
esuritor, Ashmunella 88
Euchemotrema 79, 83, 84, 88, 93
fasciatum 88
fraternum 79, 88
hubrichti 88
leai 88, 94
monodon 79, 88
monodon aliciae 79
occidaneum 88
wichitorum 88
euglypta, Linisa 90
Eusiphorus 275
euthales, Vespericola 86
Euthyneura 377, 394, 404, 408, 418,
444
eutropis, Canariella 159
eutropis, Helix 159
exiguus, Pseudomelampus
exilis, Alexia 196, 200
exodon, Polygyra stenotrema 89
exodon, Stenotrema 79

194

175, 302, 332

INDEX

exodon, Stenotrema (Pilsbrelix) 89, 93

falklandica, Philine 377, 389, 390, 397,
399, 434, 435
fallax, Triodopsis 71, 86
fallax, Triodopsis (Triodopsis) 86, 101
Fargoa 451
fasciata, Aplysia 441
fasciatum, Euchemotrema 88
fasciatus, Melampus 275, 308
fasciatus, Persicula barbadensis 256
fatigiata, Millerelix (Prattelix) 91
fausti, Oncomelania hupensis 149, 152,
153
ferrissi, Ashmunella 88
ferrissi, Inflectarius 104-106
ferrissi, Inflectarius (Inflectarius) 92, 94
ferruginea, Runcina 402
ficula, Volvatella 399, 440
figulina, Helix 1
filholi, Marinula 226, 227, 229, 331
filmargo, Helicella (Helicopsis) 1
filmargo, Martha 1
finlayi, Acteon 432
firmini, Monica 226
firminii, Ovatella 164, 175, 193, 226
flava, Voluta 293, 294
flavescens, Praticolella (Praticolella) 90
flavus, Melampus 290, 291, 293-295
florida, Stenotrema 79
florida, Stenotrema (Stenotrema) 89
floridana, Auricula 285
floridana, Detracia 284, 285
floridana, Tralia 275,284
floridana, Tralia (Tifata) 284
floridanus, Melampus 265, 284, 285
floridanus, Melampus (Detracia) 164,
IO; 173,203, 246, 275, 276, 281,
282, 284-289, 287, 290, 300, 314
floridanus, Melampus (Tralia) 284
floridianus, Melampus 284
fontalis, Physa 64
fornicata, Crepidula 445,459
fosteri, Polygyra appressa 85
fosteri, Xolotrema 71, 97, 103
fosteri, Xolotrema (Wilcoxorbis) 85, 95,
97
fragilis, Volvatella 400, 440
fraternum, Euchemotrema 79, 88
fraudulenta, Triodopsis (Shelfordorbis) 85
frumentum, Sarnia 175, 232
frumentum, Talia (Sarnia) 302
fruticum, B. 9
fulciden, Triodopsis (Macmillanorbis) 86
Fumonelix 84, 92, 101, 104, 105
archeri 72, 92, 104
jonesiana 92
jonesianus 72, 104
orestes 72, 92, 104
weatherbyi 92
wheatleyi 92, 94
Fumonelix (Fumonelix) 83

521

fusca, Melampus 291, 294

galea, Топпа 23-32, 25, 26
gangetica, Auricula (Auriculina) 179
gangeticum, Ellobium (Auriculodes)
Gastropteron 384
gattoi, Helix 357
gaziense, Ellobium (Auriculodes)
181, 184
Gegania 377, 379, 384, 384, 385, 386,
390, 391, 394, 395, 398-400, 408-
410, 413, 415-417, 419, 422, 442
valkyrie 377, 378, 390, 396, 397,
408, 417, 422, 442
georgianus, Viviparus 50
Geovula 178
germana, Cryptomastix (Micranespia) 87,
4

175

179;

germana, Helix 87

ghisottii, Cernuellopsis 359

gibba, Philine 377, 382, 389, 390, 420,
434

gibberosa, Astraea 491

Giffordius 81, 83, 84, 89, 96, 104, 105
corneliae 89
pinchoti 81, 89, 94, 104, 105

glabrata, Biomphalaria 14

globulosus, Pedipes 214, 215

globulsus, Pedipes 214

globulus, Alexia 196, 200

globulus, Auricula 287

globulus, Melampus (Detracia) 275

globulus, Pedipes 214, 216, 217

globulus, Tralia (Tifata) 275

gomerae, Canariella 159

gomerae, Helix (Gonostoma) 159

gouldiana, Bulla 405, 420, 436

gracılis, Blauneria 186

gracilis, Melampus 194

gracilis, Millerelix 91

gracilis, Monica 226

gracilis, Turbo 501

graeca, Bithynia 50

grandis, Aplysia 441

granifer, Melampus (Signia) 175

griseola, Praticolella (Praticolella) 90

guangxiensis, Oncomelania hupensis 139,
145, 152, 153

gundlachi, Апсиа 263

gundlachi, Melampus 246, 263, 268

gundlachi, Melampus coffeus 263

gundlachi, Melampus coffeus var.

Gymnophora 418

Gyraulus albus 19, 64

gyrina, Physa 66, 345

gyrina, Physella 345, 346

263

Haliotis 459

Наттоеа 376, 377, 381, 382, 384,
385, 386, 390, 391, 393-395, 398-
400, 402, 405, 409, 410, 412, 416,
418-420, 424, 433, 437

522

antillarum 388, 392, 395, 398, 400,
419, 420, 437

elegans 398, 419, 437

hydatis 398, 402, 420, 437

musetta 437

natalensis 387

navicula 398

solitaria 387,419, 420, 437

succinea 419, 420, 437

virescens 398, 437

zelandiae 419, 437
Haminoeidae 376, 417, 420, 421
hapla, Vespericola 86
harrisi, Ashmunella 88
hausmani, Daedalochila (Daedalochila) 91
hawleyi, Ashmunella 88
hebardi, Ashmunella 88
heilprini, Turbonilla 501
Heliacus variegatus 416
Helicella 349, 358

pappi 1-11
Helicella (Helicopsis) filmargo 1
Helicella (Xerothracia) pappi 1-11
Helicidae 353
helicina, Pythia 193
Helicopsis 1
Helisoma duryi 14, 17

trivolvis 64, 65
Helix afra 213

albolabris 78

amanda 349, 350, 356

ampla 90

appressa 92

auriculata 91

barbigera 88

dentifera 78

eutropis 159

figulina 1

gattoi 357

germana 87

hippocrepis 91

hirsuta 89

inflecta 92

jejuna 90

limbata 350, 356

loricata 87

lucorum 2,9

mobiliana 90

monodon 88

mooreana 91

pennsylvanicus 92

profunda 87

pustula 90

rozeti 356

scarabeus 193

stenotrema 88

thyroidus 93

townsendiana 87

tridentata 85

usticensis 349

variabilis 359

wheatleyi 92

INDEX

Helix (Ciliella) lanosa 159

Helix (Gonostoma) дотегае 159
hispidula subhispidula 159

Helix (Polygyra) anilis 89

Helminthoglypta 84
tudiculata 95

Helminthoglyptidae 95

hemphilli, Sayella 445, 446, 450, 457-
459, 464, 465, 468, 469, 481-487,
494-495

hemphilli, Turbonilla 445, 446, 450, 458,
464, 473, 474, 477, 478, 481-487,
501-502

hemphillir, Leuconia 493, 494

hemphillii, Melampus (Sayella) 494

hendersoni, Cryptomastix (Bupigona) 94

hendersoni, Polygyra mullani 87

hendersoni, Polygyra (Bupiogona) 87

henriettae, Triodopsis (Haroldorbis) 72,
86

hertleini, Linisa 90

Heterobranchia 376, 416, 421, 443-511

heteroclita, Acteon 187

heteroclita, Auricula 187

heteroclita, Blaumeria 187

heteroclita, Blauneria 164, 170, 172,
175, 176, 186-191, 11887 189236
330, 332

heteroclita, Voluta 186

Heterostropha 451

hindsii, Linisa 90

hipolitensis, Odostomia (Chrysallida) 489

hippocrepis, Daedalochila 81

hippocrepis, Daedalochila (Upsilodon) 72,
91, 93, 94, 96, 107

hippocrepis, Helix 91

hiriarti, Alexia 195

hiriarti, Alexia (Auricula) myosotis var.
195197

hirsuta, Helix 89

hirsuta, Plecotrema 209

hirsutum, Stenotrema 79

hirsutum, Stenotrema (Toxotrema) 89,
93, 95

hirsutus, Hochbergellus 86, 93

hispidula, Canariella 159

Hochbergellus 83,84, 86
hirsutus 86, 93

Holothuria sanctori 23
tubulosa 23

hombergi, Tritonia 395

hopetonensis, Triodopsis (Triodopsis) 86

hortensis, Cepaea 111

Houbricka 458, 502-503
incisa 445, 446, 450, 458, 464, 473,

474, 478, 479, 481-487, 502, 503

hubrichti, Euchemotrema 88

humboldti, Turbonilla 501

humilis, Lymnaea 65

hupensis, Oncomelania 133-156

hupensis, Oncomelania hupensis 134,
139, 140, 145, 152-154

hupensis, Oncomelania hupensis (Таиз И
form) 738, 153

Hydatina 376-378, 381-383, 384, 385,

386, 388-390, 395, 398, 399, 402,
409, 410, 415-417, 419, 420, 422,
433
physis 389, 396, 398, 433
velum 389, 398, 418, 433
vesicaria 389, 433
Hydatinidae 386
hydatis, Наттоеа 398, 402, 420, 437
Hydrobia 144
ulvae 381
Hydrobiidae 381
Hygromia 356, 359
Hygromiidae 353
Hygromiinae 358

idiogenes, Linisa 90
implicata, Millerelix 91
impressa, Boonea 488
incisa, Houbricka 445, 446, 450, 458,
464, 473, 474, 478, 479, 481-487,
502, 503
incisa, Turbonilla 458, 502, 503
indianorum, Patera (Vesperpatera) 92
inflammatum, Ellobium 257
inflecta, Helix 92
Inflectarius 84, 92, 107
ferrissi 104-106
inflectus 104
Inflectarius (Hubrichtius) 83, 84, 92
downieanus 92, 94
kalmianus 92
Inflectarius (Inflectarius) 83, 92
approximans 92
edentatus 92
ferrissi 92, 94
inflectus 92, 94, 101
magazinensis 92
гиде! 72, 92
smithi 92
subpalliatus 92
verus 92
Inflectarius (Summinflectarius) 101
inflectus, Inflectarius 104
inflectus, Inflectarius (Inflectarius) 92,
94, 101
infrequens, Auricula 202
instabilis, Retusa 393
insularis, Pedipes 219, 220, 223
insularis, Rangitotoa 231, 232, 236
integra, Physa 66
intricata, Ashmunella 88
iredalei, Tathrella 445, 446, 450, 454,
458, 464, 473, 478, 480, 481-487,
503-505
islandica, Arctica 371

jacksoni, Millerelix (Prattelix) 91
jamesensis, Ashmunella 88

INDEX 523

Jaminia 192, 193
denticulata 192, 194
quinquedens 194
seminuda 488
japonica, Julia 400
japonica, Philine 434
jaumei, Auricula 263, 268, 269
jaumei, Melampus 263
jeffreysi, Turbonilla 502
Jeffreysii, Turbonilla 450
jejuna, Helix 90
jejuna, Praticolella (Filapex) 90, 95
jonesiana, Fumonelix 92
jonesianus, Fumonelix 72, 104
juanezensis, Ashmunella 88
judae, Auriculus 178
judithae, Cingulina 491
judithae, Odostomia (Miralda) 491
judithae, Pyramidelloides 491
Julia japonica 400
juliana, Aplysia 441
Juliidae 421
junior, Tralia (Alexia) myosotis forma 196
juxtidens, Polygyra tridentata 86
juxtidens, Triodopsis 86
juxtidens, Triodopsis (Vagvolgyrorbis) 86,
95

kalmianus, Inflectarius (Hubrichtius) 92
kalmianus, Mesodon 92

karokorum, Vespericola 86

Kelliellidae 364

kiowaensis, Patera (Vesperpatera) 92
kochi, Ashmunella 88

Kochia 192

kuesteri, Auricula 275

kutschigiana, Auricula 194

labrella, Cassidula 193, 209, 259

labrosum, Stenotrema 79

labrosum, Stenotrema (Toxotrema) 89

lacteus, Turbo 501

Laemodonta 164, 173, 176, 177, 179,
190, 192, 204-206, 211, 236, 312,
313
clausa 206
cubensis 173, 193, 205-210, 207-

209,236, 241, 330, 332

molinifera 209
octanfracta 175,206, 207, 209
punctigera 205
striata 204, 205, 209

Laemodonta (Bullapex) cubensis 204,
206

laevior, Patera 71, 97, 103

laevior, Patera (Patera) 92, 97

laguncula, Volvatella 440

Laimodon 205

Laimodonta 179, 204, 205

lanosa, Helix (Ciliella) 159

lapillus, Nucella 407

lawae, Praticolella 81

524

lawae, Praticolella (Filapex) 90
leai, Euchemotrema 88, 94
leatherwoodi, Patera (Vesperpatera) 92
lens, Lindolhomia 1
lenticula, Ashmunella 88
lepidoderma, Ashmunella 88
leporina, Daedalochila 81
leporina, Daedalochila (Upsilodon) 91
leprosa, Canariella 159
Leptoxis carinata 50
letourneuxi, Alexia 196, 200, 202
Leuconia 228
hemphilli 493, 494
occidentalis 231, 232
succinea 226
Eeuconopsis 173, 124, 176, 211, 212,
237238, 2437 2443122313
manningi 170, 172, 238, 241-243,
242, 243, ЗЛА, 330
novimundi 170, 172, 238-241, 239,
240, 243, 330, 332
obsoleta 175, 237, 240, 244
rapanulensis 238, 241, 243, 244
Leucopepla 235
occidentalis 232
levettei, Ashmunella 88
Liamorpha babylonia 491
lignarius, Scaphander 389, 401, 409,
433
limax, Tamanovalva 400
limbata, Caracolla 350, 352, 353, 355,
356
limbata, Helix 350, 356
Limnaea 174
Lindolhomia lens 1
lineatus, Melampus 248, 263, 266, 267
lineatus, Melampus bidentatus 263, 270
lineatus, Melampus bidentatus var. 262,
265
lineatus, Melampus (Melampus) 262
Linisa 81, 83, 84, 89 90, 107
adamnis 90
albicostulata 90
anilis 90, 94
aulacomphala 90
behri 90, 94, 96
bicruris 90
cantralli 90
couloni 90
dissecta 90
dysoni 90
euglypta 90
hertleini 90
hindsii 90
idiogenes 90
matermontana 90
nelsoni 90
palgioglossa 90
pergrandis 90
polita 90
ponsonbyi 90
richardsoni 90

INDEX

suprazonata 90
tamaulipasensis 81, 90
texasiana 81, 90
ventrosula 90
yucatanea 90
Lirator 205
lithica, Millerelix 91
Littorina 123-132, 399, 404, 417
brevicula 130
neritoides 123-132
obtusata 130
punctata 123-132
saxatilis 123-132; 125
strigta 130
Littorina (Melaraphe) neritoides 123-132;
128
punctata 123-132; 125
Littorinacea 310
livida, Odostomia (Syrnola) 494
livida, Sayella 494
Lobosculum 81, 83, 84, 89, 90, 93, 96,
107
pustula 81, 90, 94
pustuloides 90
lombardii, Allogona (Dysmedoma) 87
longidens, Ensiphorus 276
longidens, Melampus (Ensiforus) 275
loricata, Helix 87
loricata, Trilobopsis 87, 95
loweana, Alexia 195
lozanoi, Ercolania 400
lucorum, Helix 2,9
luteus, Melampus 269, 308
[утпаеа catascopium 62, 64, 65
elodes 62, 65, 345
humilis 65
palustris 64, 65
регедга 13-21, 62, 64-66
stagnalis 13-21, 50
lynchnuchus, Pleurodonte 95

M. cartusiana 9

Macmillanorbis 85-86 [see Triodopsis
(Macmillanorbis)]

macromphala, Ashmunella 88

maculosa, Топпа 23-32, 29-31

Maelampus 247

magazinensis, Inflectarius (Inflectarius)
92

magna, Cylichna 435

magnifica, Calyptogena 363, 364, 365,
366-371, 3117872

magnifumosum, Stenotrema 79

magnifumosum, Stenotrema (Stenotrema)
89

major, Neohelix 70, 71

malleatus, Viviparus 50

mandroni, Маппийа 226, 331

manningi, Leuconopsis 170, 172, 238,
241-243, 242, 243, 314, 330

maoria, Opimilda 416

marinensis, Vespericola 86

INDEX

Marinula 164, 176, 184, 211, 213, 217,
225, 228,230; 238312 315
brevispira 226
concinna 226
filholi 226, 227, 229, 331
mandroni 226, 331
parva 227, 331
patula 225
pepita 175, 226, 227, 229, 308, 331
succinea 226
tristanensis 229, 331
velaini 226, 331
xanthostoma 217, 225, 226, 230, 331

Marinula (Cremnobates) xanthostoma
332

Marinula (Marinula) tristanensis 332

Maripythia 225

Marsyas 178

martensiana, Polygyra 90

martensiana, Polygyra (Eduardus) 90

martensiana, Praticolella (Eduardus) 95

Martha filmargo 1

marylandica, Alexia myosotis 196, 200,
202

marylandica, Phytia myosotis 196

matermontana, Linisa 90

Mathildidae 376, 377, 421

maxillatum, Stenotrema 79

maxillatum, Stenotrema (Stenotrema) 89,
95, 107

maxillifer, Stenotrema (Stenotrema) 72

mearnsi, Ashmunella 88

megasoma, Vespericola 86

Melampa 247
monile 290

Melampidae 173, 174

Melampinae 169, 170, 172, 174, 175,
177,211, 231, 243-247, 258, 302;
313, 315

Melampodinae 243, 246

Melampoides 176, 299, 315

Melampus 164, 169, 176, 179, 213,
231, 246-249, 270, 291, 297, 299,
312,314, 494
acinoides 314
aequalis 248

` bidentatus 203, 262, 266, 329
bidentatus bidentatus 263
bidentatus corneus 263
bidentatus lineatus 263,270
bidentatus redfieldi 263, 270
bidentatus var. borealis 263
bidentatus var. lineatus 262, 265
bidentus 263
biplicatus 250
boholensis 164, 258
borealis 194, 203, 268
bulimoides 277
bulla 276
bullaoides 275, 277
caffeus 250
cingulatus 276

525

clarki 297
clarki 297
coffea 249
coffea var. microspira 244, 250
coffeus 247,249, 258, 329, 331
coffeus coffeus 250
coffeus gundlachi 263
coffeus var. alternatus 263, 265
coffeus var. bishopi 263, 265
coffeus var. gundlachi 263
coffeus var. redfieldi 263
coffeus var. verticalis 263, 265
coniformis 247, 249
corneus 263
coronatus 290, 293
coronulus 291
denticulatus 195
fasciatus 275, 308
flavus 290, 291, 293-295
flavus albus 293
flavus purpureus 293
flavus var. albus 291
flavus var. purpureus 291
floridanus 265, 284, 285
floridianus 284
fusca 291, 294
gracilis 194
gundlachi 246, 263, 268
jaumei 263
lineatus 248, 263, 266, 267
luteus 269, 308
monile 275,290, 291, 330
monilis 291
morrisoni 330
myosotis 195
nitens 303
nucleolus 270, 308
obliquus 262, 266-268, 270
oblongus 277, 279, 281
olivula 250, 258
ovulum 303
pallescentis 269
paranus 330
poeyi 277
pusillus 303
redfieldi 263, 268
riparius 180
spiralis 263, 269
torosa 291, 294
turritus 195, 203
Melampus (Autonoe) riparius 180
Melampus (Autonoella) riparius 180
Melampus (Conovulus) biplicatus 249
coffeus 249
Melampus (Detracia) 315
bullaeoides 277
bullaoides 169, 173, 175, 275, 276-
284, 279-282, 295, 300, 308, 332
bulloides 277
floridanus 164, 170, 173, 203, 246,
275, 276, 281, 282, 284-289, 287,
290, 300, 314

526

globulus 275
топйе 173, 241, 247, 249, 250, 257,
267, 275,276, 290-297, 293, 294,

308

morrisoni 173, 275, 276, 295, 297-
302, 299-301

paranus 164, 170, 173, 176, 276,
289-290, 290

Melampus (Ensiforus) longidens 275

Melampus (Leuconia) bidentatus 202,
267

Melampus (Lirator) multisulcatus 205

Melampus (Melampus) bidentatus 164,
1165, 168, 170, 173; 176; 202.203;
246, 248, 249, 257, 258, 262-275,
264-270, 284, 285, 295, 300, 308,

314
bulla 276
coffea 250

Gorfeus 164, 168, IO 17051725
17,3, 775, 175, 176.245; 24 o Cae eje
262, 251-256, 265, 266, 268, 290,
2942957, 3007 S087 332

corneus 263

lineatus 262

monile 290

obliquus 263

Melampus (Micromelampus) nucleolus

175

Melampus (Microtralia) minusculus 232
Melampus (Pira) monile 291
monilis 291
Melampus (Sayella) crosseana 495
hemphillii 494
Melampus (Signia) granifer 175
Melampus (Tifata) cingulata 277
Melampus (Tralia) cingulatus 277

floridanus 284

olivula 250

pusillus 303

Melania 174

Melanoides tuberculata 50

Melanopsis 174

Melaraphe 130

mendax, Ashmunella 88

meridionalis, Ashmunella 88

meridionalis, Auricula (Alexia) 195, 197,
203

Mesodon 84, 92, 93, 707

andrewsae 104

chilhoweensis 104, 105

kalmianus 92

normalis 71, 104

zaletus 104

Mesodon (Akromesodon) 83, 84, 93,

104, 105

altivagus 93

andrewsae 93

normalis 93, 94

Mesodon (Aphalogona) 83, 84, 93, 107

elevatus 93, 94

mitchellianus 93

INDEX

zaletus 93
Mesodon (Mesodon) 83, 93
clausus 93
sanus 93
thyroidus 93, 94
trossulus 93
Mesodontini 84, 89, 92, 93
Metaruncina setoensis 402
micheli, Alexia 195, 197
Micromelampus 270
microspira, Melampus coffea var. 244,
250
microstoma, Auricula 194
Microtralia 169, 172, 174, 176, 190,
210-212, 228, 230-232, 236302}
312
alba 236
occidentalis 172, 175, 197, 232-237,
232, 234-236, 330, 332
Microxeromagna 358
midae, Аипсийа 178
midae, Ellobium 178
Millerelix 81, 84, 89, 91
doerfeuilliana 81, 91
gracilis 91
implicata 91
lithica 91
mooreana 81, 91
plicata 81
tholus 91
Millerelix (Millerelix) 83, 91, 96
mooreana 94
tamaulipasensis 72
Millerelix (Prattelix) 81, 83, 84, 91, 93,
96
deltoidea 91
fatigiata 91
jackson! 91
peregrina 91
plicata 72, 80, 91, 94
simpson 91
troostiana 91
minima, Oncomelania 133, 147
minimum, Carychium 175
minuscula, Auricula (Microtralia) 231,
2329232
minuscula, Tralia 232
minuscula, Tralia (Alexia) 232, 232, 236
minusculum, Auriculastrum (Microtralia)
232
minusculus, Melampus (Microtralia) 232
minuta, Voluta 257,294, 295
miorhyssa, Ashmunella rhyssa 87
miorhyssa, Polygyra 87
mirabilis, Pedipes 170, 172, 175, 213-
218, 215-217, 219) 221, 223295)
314: 330.332
mirabilis, Turbo 213
mirandus, Pedipes 219, 223
mitchellianus, Mesodon (Aphalogona) 93
mitralis, Aphalista 499
mitralis, Pyramidella 445, 446, 450, 451,

INDEX

452, 458, 464, 472-474, 476, 481-
487, 496, 499-500
mitralis, Pyramidella (Otopleura) 499
mobiliana, Helix 90
mobiliana, Praticolella 81
mobiliana, Praticolella (Farragutia) 90, 95
mogollonensis, Ashmunella 88
molinifera, Laemodonta 209
Monacha 9
Monica aequalis 226
firminii 226
gracilis 226
monile, Auricula 290
monile, Bulimus 290, 293
monile, Conovulus 290
monile, Melampa 290
monile, Melampus 275,290, 291, 330
monile, Melampus (Detracia) 173, 241,
247, 249, 250, 257, 267, 275, 276,
290-297, 293, 294, 308
monile, Melampus (Melampus) 290
monile, Melampus (Pira) 291
monile, Pira 291
monile, Voluta 294
monilis, Melampus 291
monilis, Melampus (Pira) 291
monilis, Obeliscus 497
monodon, Euchemotrema 79, 88
monodon, Helix 88
montivaga, Ashmunella 88
mooreana, Helix 91
mooreana, Millerelix 81, 91
mooreana, Millerelix (Millerelix) 94
morosum, Stenotrema (Stenotrema) 89
morrisoni, Melampus 330
morrisoni, Melampus (Detracia) 173, 275,
276, 295, 297-302, 299-307
mudgei, Ashmunella 88
mudgii, Ashmunella 72
mullani, Cryptomastix (Cryptomastix) 87,
94
multilineata, Webbhelix 95
multiplicata, Daedalochila 81
multiplicata, Daedalochila (Upsilodon) 91
multisulcatus, Melampus (Lirator) 205
multivolvis, Auricula 276, 279, 308
musetta, Haminoea 437
mustelina, Cassidula 332
Myosotella 173, 175, 176, 191, 192,
192-194, 201, 240, 312, 314
myosotis 164, 169, 172, 175, 176,
193, 194-204, 197, 200, 201, 228,
230. 232. 207, 2658 270, 302.314,
330, 332
payraudeaui 192, 194
myosotis, Alexia 195, 267
myosotis, Alexia myosotis 196
myosotis, Alexia (Myosotella) 196
myosotis, Auricula 192, 194, 201
myosotis, Auricula (Alexia) 196
myosotis, Auricula (Auricula) 194
myosotis, Melampus 195

527

myosotis, Myosotella 164, 169, 172,
175, 176, 193, 194-204, 197, 200,
201,228, 230,232, 207,268. 270;
302.314, 330.332

myosotis, Ovatella 196, 202

myosotis, Ovatella (Alexia) 197

myosotis, Ovatella (Myosotella) 197

myosotis, Phytia 196, 201

myosotis, Phytia myosotis 196

myosotis, Pythia 194

mysotis, Auricula 194

Mytilus edulis 124, 129, 130

nana, Auriculastra 232, 236
nantahalae, Patera clarki 72
natalensis, Haminoea 387
Natica 381
Naticidae 381
naticoides, Pedipes 219, 220, 221], 223
navicula, Haminoea 398
Nealexia 192, 193
neglecta, Triodopsis (Vagvolgyrorbis) 86
nelsoni, Linisa 90
Neohelix 78, 84
albolabris 80, 104
alleni 80
dentifera 106
major 70, 71
solemi 80
Neohelix (Asamiorbis) 78, 83, 84
dentifera 94
Neohelix (Neohelix) 78, 83
albolabris 80, 95
Neohelix (Solemorbis) 78, 80, 85; 83, 84
alleni 85
solemi 85, 95
Neohelix (Wilcoxorbis) 83
Nerítina 193
Nerítino 193
neritoides, Littorina 123-132
neritoides, Littorina (Melaraphe)
128
nitens, Auricula 303
nitens, Melampus 303
nitida, Ringicula 377,387, 388, 396,
405, 415, 432
normalis, Cepaea 111-122, 145
normalis, Mesodon 71, 104
normalis, Mesodon (Akromesodon) 93,
94
normalis, Polygyra andrewsae 93
norrisi, Norrisia 459, 491
Norrisia norrisi 459, 491
Notaspidea 418, 421
novimundi, Apodosis 237, 238, 241, 242
novimundi, Leuconopsis 170, 172, 238-
241, 239, 240, 243, 330, 332
Nucella lapillus 407
nucleolus, Melampus 270, 308
nucleolus, Melampus (Micromelampus)
175
Nudibranchia 404, 418, 421

123-132;

528

Obeliscus 496
arenosa 498
monilis 497
sulcatus 497
teres 497
tessellatus 497
obliqua, Auricula 263
obliquus, Melampus 262, 266-268, 270
obliquus, Melampus (Melampus) 263
oblongus, Melampus 277, 279, 281
obsoleta, Alexia 195
obsoleta, Leuconopsis 175,237, 240,
244
obsoleta, Triodopsis (Triodopsis) 86
obstricta, Xolotrema (Xolotrema) 85
obtusa, Retusa 382, 384, 391, 393,
396, 398, 399, 420, 438
obtusata, Littorina 130
obvia, Xerolenta 1
ocampi, Praticola 90
occidaneum, Euchemotrema 88
occidentalis, Leuconia 231, 232
occidentalis, Leucopepla 232
occidentalis, Microtralia 172, 175, 197,
232-237, 232, 234.23673307332
occidentalis, Xolotrema (Wilcoxorbis) 85
octanfracta, Laemodonta 175,206, 207,
209
octanfracta, Pedipes 205
Odostomia 449, 451, 488
babylonia 445, 446, 450, 457, 459,
461, 463, 464, 466, 481-487, 491-
492
cubensis 187,190
didyma 445, 446, 450, 459, 467,
463, 464, 467, 481-487, 492-493
promesces 490
santorium 490
seminuda 488
unidentata 451
Odostomia (Chrysallida) babylonica 491
cincta 489
contrerasi 490
deceptrix 490
hipolitensis 489
promeces 489
pulcherrima 489
pulcia 489
santorium 489
sapia 490
seminuda 488
toyatani 488
vicola 489
vincta 489
willisi 488
Odostomia (Cingulina) babylonica 491
Odostomia (Cyclodostomia) didyma 492
Odostomia (Miralda) judithae 491
Odostomia (Syrnola) livida 494
Odostomia (Tornatellina) cubensis 187
Odostominae 444, 448, 450, 452, 457,
463, 488

INDEX

Oleacina 189
Oleacina (Stobilus) cubensis 187
oliva, Auricula 276, 279
Oliva 381
olivula, Auricula 249, 258
olivula, Melampus 250, 258
olivula, Melampus (Tralia) 250
olneyae, Polygyra mullani 87
Omalogyroidea 417
Oncomelania 144, 145, 147, 154
hupensis 133-156
hupensis fausti 149, 152, 153
hupensis guangxiensis 139, 145, 152,

1153

hupensis hupensis 134, 139, 140,
145, 152-154

hupensis hupensis (fausti form) 138,
153

hupensis quadrasi 145
hupensis robertsoni 134, 138, 149,
(532154
hupensis tangi 134, 138, 149, 153,
154
minima 133, 147
operculata, Retusa 382, 438
Ophicardelus 176, 191, 192, 314
australis 175, 225
Opimilda maoria 416
Opisthobranchia 310, 311, 376, 396,
416-418, 443
oppilata, Daedalochila (Daedalochila) 91
oranica, Alexia 196, 202
oranica, Alexia (Kochia) 192, 193, 196,
200
orestes, Fumonelix 72,92, 104
organensis, Ashmunella 88
orientalis, Philine 397, 434, 435
orius, Vespericola 86
Otina otis 171
otis, Otina 171
Otopleura mitralis 445, 499
ovalis, Pedipes 170, 172, 213-215, 217-
225, 220-223, 228, 236, 2477330
ovalis, Pedipes mirabilis forma 219
Ovatella 175, 176, 191-193, 211, 226,
240, 312-314
aequalis 193, 209, 210, 226, 230,
332
denticulata 202
firminii 164, 175, 193, 226
myosotis 196, 202
myostis bermudensis 197
Ovatella (Alexia) myosotis 197
Ovatella (Myosotella) myosotis 197
ovula, Auricula 303
ovula, Auricula (Conovulus) 303
ovula, Tralla 236,303, 330/1332
ovula, Tralia (Tralia) 174, 175, 247, 295,
302, 303-310, 304-308, 314
ovulum, Melampus 303
ovulum, Pythia 303
ovulus, Bulimus 303, 308

INDEX

ovulus, Pedipes 217

pachyloma, Praticolella (Praticolella) 90
pacifica, Calyptogena 365, 369, 369-
370, 371; 372
palgioglossa, Linisa 90
pallescentis, Melampus 269
paludosa, Polygyra 89
palustris, Lymnaea 64, 65
palustris, Triodopsis (Triodopsis) 86
panselena, Patera (Patera) 92
pappi, Helicella 1-11
pappi, Helicella (Xerothracia) 1-11
parana, Detracia 287, 289
paranus, Melampus 330
paranus, Melampus (Detracia) 164, 170,
173, 176, 276, 289-290, 290
parva, Alexia 196, 200
parva, Cremnobates 225, 226
parva, Marinula 227, 331
parvula, Aplysia 441
pasonis, Ashmunella 88
Patella aspera 33-40
caerulea 33-40
ulyssioponensis 34
Patellidae 33-40
Patera 84, 101
appressa 104
clarki nantahalae 72
laevior 71, 97, 103
Patera (Patera) 83, 92
appressa 92, 95
clarki 92
laevior 92, 97
panselena 92
perigrapta 92
sargentiana 92
Patera (Ragsdaleorbis) 83, 84, 92
pennsylvanica 95
pennsylvanicus 92
Patera (Vesperpatera) 83, 84, 92, 707
binneyana 92, 95
clenchi 92
indianorum 92
kiowaensis 92
leatherwoodi 92
roemeri 92
patula, Marinula 225
patulus, Pythia 230
payraudeaui, Alexia 195
payraudeaui, Myosotella 192, 194
pechaudi, Alexia 196, 200
Pedipedinae 170, 172, 174, 175, 211-
21292593302.313>315
pedipes, Bulimus 213
pedipes, Pedipes 175, 212, 213, 215,
219,217
Pedipes 164, 169, 173, 176, 186, 210-
213, 225, 226, 230; 236, 238, 240,
312,313
afra 212, 217
coniformis 249

529

elongatus 226, 227, 228
globulosus 214, 215
globulsus 214
globulus 214, 216, 217
insularis 219, 220, 223
mirabilis 170, 172, 175, 213-218,
215-217, 219, 221, 223, 295, 314,
330, 332
mirabilis forma ovalis 219
mirandus 219, 223
naticoides 219, 220, 221, 223
octanfracta 205
ovalis 170, 172, 213-215, 217-225,
220-223, 228, 236, 241, 330
ovulus 217
pédipes, 175, 212, 218,215, 275.
217
quadridens 213, 214, 219
tridens 214,218, 219, 220, 221, 223
pelecais, Acteon 432
pellucens, Аипсийа 180, 184
pellucens, Auricula (Auriculastrum) 180
pellucens, Auriculastrum 180
pellucens, Auriculus 180
pellucens, Conovulus 180
pellucens, Ellobium 180
pellucens, Ellobium (Auriculodes) 183
pellucida, Achatina 187, 189
pellucida, Blanneria 187
pellucida, Blauneria 187
pendula, Triodopsis (Vagvolgyrorbis) 86
peninsulae, Daedalochila (Daedalochila)
92
penistens, Trilobopsis 87
penistoni, Turbonilla 501
pennsylvanica, Patera (Ragsdaleorbis) 95
pennsylvanicus, Helix 92
pennsylvanicus, Patera (Ragsdaleorbis) 92
Pentaganglionata 394, 404, 408
pepita, Marinula 175, 226, 227, 229,
308, 331
perdistorta, Distorsio 23
perdix, Tonna 23
peregra, Lymnaea 13-21, 62, 64-66
peregrina, Millerelix (Prattelix) 91
pergrandis, Linisa 90
perigrapta, Patera (Patera) 92
Persa 302
Persicula barbadensis fasciatus 256
persimilis, Volvulella 402
personatum, Carychium 194, 197, 201,
202
Petitella 495
crosseana 445, 446, 450, 457-459,
464, 465, 469, 470, 481-487, 495-
496
pfeifferi, Biomphalaria 18
Phanerophthalmus 393
Phaneropneumana 193
Phaneropneumona 193
phaseoliformis, Calyptogena 365, 367,
368-370, 371, 312

530

Phestilla sibogae 420

Philine 376, 377, 381, 382, 384, 385,

386, 388-390, 395, 397, 399, 400,
402, 404, 408-410, 412, 413, 415-
422, 424, 434-435

angasi 397, 434

aperta 377, 389, 390, 393, 397, 399,

408, 420, 434, 435
denticulata 434

falklandica 377, 389, 390, 397, 399,

434, 435

gibba 377, 382, 389, 390, 420, 434

japonica 434
orientalis 397, 434, 435
quadripatia 434
Philinidae 421
Philinoglossa praelonga 418
Philinoglossidae 421
Philinoidea 390, 412, 413, 416, 417,
419, 424
Phitia 192
Physa fontalis 64
gyrina 66, 345
integra 66
Physella cubensis 333-348
gyrina 345, 346
virgata 345
physis, Hydatina 389, 396, 398, 433
Phythya 192
Phytia 192, 193, 201
Phytia bermudensis 196
denticulata 194, 201, 202
myosotis 196, 201
myosotis borealis 196
myosotis denticulata 196
myosotis marylandica 196
myosotis myosotis 196
myosotis var. bermudensis 196
scarabeus 209
picea, Triodopsis (Shelfordorbis) 85
pilosus, Vespericola 86

Pilsbrelix 89 [see Stenotrema (Pilsbrelix)]

pilsbryana, Ashmunella 88

pilsbryi, Stenotrema 79

pilsbryi, Stenotrema (Archerelix) 88

Pilsbryorbis 85 [see Triodopsis
(Pilsbryorbis)]

pilula, Stenotrema 79

pilula, Stenotrema (Stenotrema) 89

pinchoti, Giffordius 81, 89, 94, 104, 105

pinicola, Vespericola 87
Pira 275, 302
monile 291
piscinalis, Valvata 50, 62
planaria, Canariella 159
Planorbarius corneus 14, 53-68
planorbis, Planorbis 19, 53-68
Planorbis carinatus 66
planorbis 19, 53-68
platysayoides, Polygyra 85
platysayoides, Triodopsis 72, 85, 104,
105

INDEX

platysayoides, Triodopsis (Brooksorbis)
85,95
Plecotrema 204, 205
clausa 209
cubense 206
cubensis 206
hirsuta 209
typica 204
Plectrotrema 164, 177
Pleurodonte 84, 96
lynchnuchus 95
Plicacea 213
plicata, Daedalochila 91
plicata, Millerelix 81
plicata, Millerelix (Prattelix) 72, 80, 91,
94
plicata, Polygyra 91
plicata, Pythia 332
plicata, Pythia (Pythia) 175
plicatula, Turbo 500, 501
plicatula, Turbonilla 500, 501
poeyi, Daedalochila (Upsilodon) 91
poeyi, Melampus 277
polita, Linisa 90
Polloneriella 358
Polygyra 81, 83, 84, 89, 93, 96, 105
andrewsae normalis 93
appressa fosteri 85
auriculata 91
binneyana 92
caloosaensis 89
cereolus 81, 89
cohuttensis 88
columbiana 86
martensiana 90
miorhyssa 87
mullani hendersoni 87
mullani olneyae 87
paludosa 89
platysayoides 85
plicata 91
septemvolva 89, 95
septemvolva volvoxis 81
stenotrema exodon 89
tridentata 73
tridentata juxtidens 86
tridentata tennesseensis 85
troostiana 73
Polygyra (Bupiogona) hendersoni 87
Polygyra (Daedalochila) texasiana 89
Polygyra (Eduardus) martensiana 90
Polygyra (Ermyodon) 90
Polygyra (Linisia) 90
Polygyra (Monophysis) 90
Polygyra (Solidens) 90
Polygyridae 69-110
Polygyrinae 86
Polygyrinai 87, 88
Polygyrini 87, 82, 84, 89, 90, 104, 106
Pomatiopsis 133
ponsonbyi, Linisa 90

INDEX

postelliana, Daedalochila (Daedalochila)
92
praelonga, Philinoglossa 418
Praticola ocampi 90
Praticolella 84, 89, 90, 96, 106
berlandieriana 81
lawae 81
mobiliana 81
Praticolella (Eduardus) 83, 84, 90
martensiana 95
Praticolella (Farragutia) 81, 83, 84, 90
mobiliana 90, 95
Praticolella (Filapex) 81, 83, 84, 90
bakeri 90
jejuna 90, 95
lawae 90
Praticolella (Praticolella) 81, 83, 90
ampla 90, 95
berlandieriana 90
candida 90
flavescens 90
griseola 90
pachyloma 90
strebeliana 90
taeniata 90
trimatris 90
pratii, Pyramidella 497
Prattelix 91 [see Millerelix (Prattelix)]
pressleyi, Vespericola 87
pricena, Wingenella 497
profunda, Allogona 71, 101, 104
profunda, Allogona (Allogona) 87, 94, 99
profunda, Helix 87
promeces, Odostomia (Chrysallida) 489
promesces, Odostomia 490
propinqua, Pyramidella 499
Prosobranchia 443, 444
proxima, Ashmunella 88
pseudodonta, Ashmunella 88
Pseudomelampus 169, 174, 211, 212,
232, 302. 912-314
exiguus 175, 302, 332
ptychophora, Allogona (Dysmedoma) 87
pulchella, Alexia 196, 200
pulchella, Tralia 275
- pulcherrima, Odostomia (Chrysallida) 489
pulcia, Odostomia (Chrysallida) 489
Pulmonata 311, 376, 443, 444
puncatata, Aplysia 441
punctata, Littorina 123-132
punctata, Littorina (Melaraphe) 123-132;
125
punctigera, Laemodonta 205
punctostriatus, Scaphander 389, 433
purpureus, Melampus flavus 293
purpureus, Melampus flavus var.
pusilla, Аипсиа 303
pusilla, Tralia 302,303
pusilla, Voluta 303,308
pusillus, Auricula (Conovulus) 303
pusillus, Melampus 303
pusillus, Melampus (Tralia) 303

291

531

pustula, Helix 90
pustula, Lobosculum 87,90, 94
pustuloides, Lobosculum 90
Pyramidella 449, 453, 496
crenulata 445, 446, 450, 458, 464,
472, 474, 475, 481-487, 498-499
mitralis 445, 446, 450, 451, 452,
458, 464, 472-474, 476, 481-487,
496, 499-500
pratii 497
propinqua 499
sulcata 445, 446, 449, 450, 452,
458, 464, 471-474, 481-487, 497-
500
teres 497
tessellatus 497
variegata 499
Pyramidella (Longchaeus) crenulata 498
Pyramidella (Otopleura) mitralis 499
Pyramidellidae 169, 376, 377, 404, 418,
420, 421, 443-511
Pyramidellinae 444, 448-450, 452, 455,
458, 460, 472, 496, 500
Pyramidelloidea 310, 417, 444, 446, 447
Pyramidelloides judithae 491
Pyramidellus 496
Pyrgiscus 504
pyriformis, Volvatella 440
Pythia 164, 170, 173, 176, 191-193,
209, 247, 312-314
abbreviatus 228, 230
aequalis 230
denticulata 194
helicina 193
myosotis 194
ovulum 303
patulus 230
plicata 332
scarabeus 193
triplicata 303
Pythia (Pythia) plicata 175
scarabaeus 175
Pythia (Trigonopythia) trigona 175
Pythiinae 169, 170, 172, 174, 175, 186,
191192, 20572117212 240/2599,
313,315

gibbus, Aequipecten 459

quadrasi, Cylindrotis 175

quadrasi, Oncomelania hupensis 145
quadridens, Pedipes 213, 214, 219
quadripatia, Philine 434

quinquedens, Jaminia 194

Ranellidae 23

Rangitotoa 211, 231, 246
insularis 231, 232, 236

rapanuiensis, Leuconopsis 238, 241,
243, 244

redfieldi, Melampus 263, 268

redfieldi, Melampus bidentatus 263, 270

redfieldi, Melampus coffeus var. 263

532

reflexa, Voluta 194, 197, 202
reflexilabris, Auricula 194, 197, 203
Retusa 376, 377, 381, 384, 386, 391,
395, 396, 399, 402, 409, 412, 413,
415-419, 421, 422, 424, 438-439
chrysoma 382, 438
instabilis 393
obtusa 382,384, 391, 393, 396, 398,
399, 420, 438
operculata 382,438
semisulcata 377,398, 399, 438
truncatula 384, 393, 396, 398-400,
420, 438
Retusidae 376, 402, 417, 421
rhyssa, Ashmunella 88, 94, 96
Rhytophorus 176, 315
richardsoni, Linisa 90
rileyensis, Ashmunella 88
ringens, Auricula 433
ringens, Tralia (Alexia) myosotis var. 196
ringens, Voluta 194, 197, 202
ringicula, Alexia 196, 200, 202
Ringicula 312, 332, 376, 377, 379; 381-
383, 384, 387-389, 393, 395, 402,
405, 407-410, 412, 415, 416, 422,
432-433
buccinea 377,387, 388, 405, 432
coniformis 377,387, 388, 405, 432
nitida 377,387, 388, 396, 405, 415,
432
Ringiculidae 376, 421
riparia, Autonoe 178, 180
riparia, Autonoella 179
riparius, Melampus 180
riparius, Melampus (Autonoe) 180
riparius, Melampus (Autonoella) 180
Rissoacea 310
Rissoellidae 377
Rissoelloidea 417
robertsoni, Oncomelania hupensis 134,
138, 149, 153, 154
roemeri, Patera (Vesperpatera) 92
roperi, Trilobopsis 72, 87
roquesana, Detracia 277, 279, 281
rotundatus, Anisus 62
rotundatus, Armiger 64
rozeti, Helix 356
rugeli, Auricula 285
rugeli, Inflectarius (Inflectarius) 72, 92
rugosa, Cassidaria 23, 31
rugosa, Cernuella 350
rugosa, Triodopsis 85
rugosa, Triodopsis tridentata 85
rugosa, Triodopsis (Macmillanorbis) 86,
95
ruidosana, Ashmunella 88
Runcina 391
coronata 401
ferruginea 402
Runcinidae 391, 402

Sacoglossa 379, 381, 412, 416, 418,

INDEX

419, 421, 424
Sagda 84, 96
cookiana 95
Sagdidae 95
salinasensis, Ashmunella 88
sanburni, Cryptomastix (Cryptomastix) 87
sanctori, Holothuria 23
santorium, Odostomia 490
santorium, Odostomia (Chrysallida) 489
sanus, Mesodon (Mesodon) 93
sapia, Odostomia (Chrysallida) 490
sargentiana, Patera (Patera) 92
Sarnia 174, 232, 302
frumentum 175, 232
saxatilis, Littorina 123-132; 125
sayana, Appalachina 71, 94, 99, 101,
104

sayanus, Appalachina 93
Sayella 191, 493-494
crosseana 495
hemphilli 445, 446, 450, 457-459,
464, 465, 468, 469, 481-487, 494-
495
livida 494
Sayellinae 493
sayi, Auricula 202, 203
Scaphander 376,377,379, 381, 382,
384, 385, 386, 388-390, 397 397
399, 400, 402, 404, 405, 408-410,
415, 416, 418, 419, 424, 433-434
clavus 433
cylindrellus 390
lignarius 389,401, 409, 433
punctostriatus 389, 433
watsoni 389, 390, 434
Scaphandridae 376
scarabaeus, Pythia (Pythia)
scarabeus, Helix 193
scarabeus, Pythia 193
Scarabinae 191
Scarabus 191, 247
scrobiculata, Auricula watsoni 195, 197,
202
sculpta, Tralia ovula 303, 304, 308
seminuda, Boonea 445, 446, 450, 457,
459, 460, 461, 463-465, 481-490
seminuda, Chemnitzia 488
seminuda, Jaminia 488
seminuda, Odostomia 488
seminuda, Odostomia (Chrysallida) 488
semisulcata, Retusa 377, 398, 399, 438
septemvolva, Polygyra 89, 95
setifer, Alexia 195, 196, 197
setifera, Alexia 195
setigera, Alexia 195
setoensis, Metaruncina 402
shasta, Vespericola 87
sibogae, Phestilla 420
sierrana, Vespericola 87
Signia 302
similaris, Bradybaena 95
simile, Stenotrema (Toxotrema) 89

175

INDEX

simpson, Millerelix (Prattelix) 91
Sinistrobranchia 444
Siphonariidae 311
Smaragdinella 376,377, 382, 384, 391,
393, 395, 400, 402, 405, 408-410,
412, 416, 418, 424, 437-438
calyculata 437
smithi, Inflectarius (Inflectarius) 92
solemi, Neohelix 80
solemi, Neohelix (Solemorbis) 85, 95
Solemorbis 78, 80, 85 [see Neohelix
(Solemorbis)]
soleneri, Triodopsis (Triodopsis) 86
solida, Bulla 436
solida, Cremnobates 225
solitaria, Haminoea 387, 419, 420, 437
spelaeum, Zospeum 175
Sphincterochila zonata 9
spinosum, Stenotrema 79
spinosum, Stenotrema (Stenotrema) 89,
95
spiralis, Melampus 263, 269
sprouli, Ashmunella 88
stagnale, Ellobium (Auriculodes)
stagnalis, [утпаеа 13-21, 50
steinbergae, Doridella 420
stenotrema, Helix 88
stenotrema, Stenotrema 79
stenotrema, Stenotrema (Stenotrema) 89,
95
Stenotrema 84, 88, 106, 107
altıspira altispira 79
altispira depilatum 79
barbatum 79
barbigerum 79
blandianum 79
brevipila 79
caddoense 79
cohuttense 79
convexa 88
deceptum 79
edgarianum 79
edvardsi 79
exodon 79
exodon turbinella 79
‘florida 79
hirsutum 79
labrosum 79
magnifumosum 79
maxillatum 79
pilsbryi 79
pilula 79
spinosum 79
stenotrema 79
turbinella 79
unciferum 79
Stenotrema (Archerelix) 79, 83, 84, 88
barbigerum 88, 93, 106
edgarianum 88
edvardsi 88
pilsbryi 88
Stenotrema (Cohutta) 79, 83, 84, 88, 93

179

533

cohuttensis 88, 95
Stenotrema (Coracollatus) 89
Stenotrema (Maxillifer) 89
Stenotrema (Pilsbrelix) 79, 83, 84, 88,
89
blandianum 89
deceptum 89
exodon 89, 93
turbinella 89
uncifera 107
Stenotrema (Stenosoma) 89
Stenotrema (Stenotrema) 79, 83, 88, 89
altispira 89
altispira depilatum 89
angellum 89
brevipila 89
caddoense 89
calvescens 89
florida 89
magnifumosum 89
maxillatum 89, 95, 107
maxillifer 72
morosum 89
pilula 89
spinosum 89, 95
stenotrema 89, 95
unciferum 72,89
waldense 89
Stenotrema (Тохойета) 79, 83, 84, 88,
89
barbatum 89
hirsutum 89, 93, 95
labrosum 89
simile 89
Stenotremini 79, 82, 84, 88, 106
sterni, Daedalochila (Upsilodon) 91
Stolidoma 186, 315
strebeliana, Praticolella (Praticolella) 90
striata, Auricula 205
striata, Bulla 387, 388, 390, 409, 420,
436
striata, Laemodonta 204, 205, 209
striata, Turbonilla 501
strigta, Littorina 130
Stylommatophora 172
subarmata, Auricula bicolor var. 195
subclausa, Daedalochila (Daedalochila) 92
subflava, Alexia 196, 202
subhispidula, Helix (Gonostoma) hispidula
159
subnodosum, Ellobium (Ellobium) 179
subpalliatus, Inflectarius (Inflectarius) 92
subula, Auriculastra 175, 184
succinea, Creedonia 173, 175, 202, 203,
225-231, 227, 2297 280,236, 330-
332
succinea, Haminoea 419, 420, 437
succinea, Leuconia 226
succinea, Marinula 226
sulcata, Pyramidella 445, 446, 449, 450,
452, 458, 464, 471-474, 481-487,
497-500

534

sulcatus, Obeliscus 497
suprazonata, Linisa 90

taeniata, Praticolella (Praticolella) 90
Tamanovalva limax 400
tamaulipasensis, Linisa 81, 90
tamaulipasensis, Millerelix (Millerelix) 72
tangi, Oncomelania hupensis 134, 138,
149, 153, 154
Tathrella 503-504
iredalei 445, 446, 450, 454, 458,
464, 473, 478, 480, 481-487, 503-
505
Tectibranchiata 376
tegillum, Ashmunella 88
Tegula eiseni 491
tehamana, Trilobopsis 87
tenella, Auricula 194, 202
tennesseensis, Polygyra tridentata 85
tennesseensis, Triodopsis (Pilsbryorbis)
85, 95
tentaculata, Bithynia 19, 50, 65
tenuis, Alexia setifer var. 195, 197
Tequla eiseni 445
teres, Obeliscus 497
teres, Pyramidella 497
terrestris, Alexia 196, 200
tessellatus, Obeliscus 497
tessellatus, Pyramidella 497
tetrodon, Ashmunella 88
texasiana, Linisa 81, 90
texasiana, Polygyra (Daedalochila) 89
tholus, Millerelix 91
thomsoniana, Ashmunella 88
thyroidus, Helix 93
thyroidus, Mesodon (Mesodon) 93, 94
Thysanophora 84
Thysanophoridae 95
Tifata 275, 302
timida, Elysia 409
Tonna galea 23-32, 25, 26
maculosa 23-32, 29-31
perdix 23
torita, Chrysallida 490
Tornatella 213
bullaoides 276
cubensis 187, 189
tornatilis, Acteon 388, 403, 407, 432
Tornatina 405
torosa, Melampus 291, 294
townsendiana, Allogona (Dysmedoma)
87,94
townsendiana, Helix 87
toyatani, Odostomia (Chrysallida) 488
trachypepla, Trilobopsis 87
Tralia 173,174, 176, 246, 302-303,
312,313
cingulata 275, 277
floridana 275, 284
minuscula 232
ovula 236, 303, 330, 332
ovula sculpta 303, 304, 308

INDEX

pulchella 275
pusilla 302, 303
venezuelana 303, 304, 308, 309
vetula 304
Tralia (Alexia) minuscula 232, 232, 236
myosotis forma junior 196
myosotis var. ringens 196
Tralia (Persa) costata 175
Tralia (Sarnia) frumentum 302
Tralia (Tifata) floridana 284
globulus 275
Tralia (Tralia) ovula 174, 175, 247, 295,
302, 303-310, 304-308, 314
vetula 309
Tralica 302
Traliopsis 315
traskii, Acteon 432
Tricula 147
Tridacna 445, 505
tridens, Pedipes 214, 218, 219, 220,
2217223
tridentata, Helix 85
tridentata, Polygyra 73
tridentata, Triodopsis 70, 74, 86, 103,
104, 106, 107
tridentata, Triodopsis (Triodopsis) 86, 95
tridentatum, Carychium 175, 193, 209,
313: 331
trigona, Ру га (Trigonopythia) 175
Trilobopsis 83, 84, 87
loricata 87, 95
penistens 87
roperi 72,87
tehamana 87
trachypepla 87
trimatris, Praticolella (Praticolella) 90
Triodopsinae 78
Triodopsini 79, 84, 106
Triodopsis 84, 85, 87, 101, 107
burchi 85
copei 86
cragini 86
fallax 71, 86
fraudulenta vulgata 85
juxtidens 86
platysayoides 72, 85, 104, 105
гидоза 85
tridentata 70, 74, 86, 103, 104, 106,
107
tridentata rugosa 85
vulgata 85, 104
Triodopsis (Brooksorbis) 83,84, 85
platysayoides 85, 95
Triodopsis (Haroldorbis) 83, 84, 86, 101
cragini 95
henriettae 72, 86
vultuosa 86
Triodopsis (Macmillanorbis) 83, 84, 85-
86
fulciden 86
rugosa 86, 95

INDEX

Triodopsis (Pilsbryorbis) 83, 84, 85, 101
burchi 85
complanata 85
tennesseensis 85, 95
Triodopsis (Shelfordorbis) 83, 84
claibornensis 85
fraudulenta 85
picea 85
vulgata 85, 95
Triodopsis (Triodopsis) 83, 85, 86
alabamensis 86
anteridon 86
fallax 86, 101
hopetonensis 86
obsoleta 86
palustris 86
soleneri 86
tridentata 86, 95
vannostrandi 86
Triodopsis (Vagvolgyrorbis) 83-85, 86
discoidea 86
juxtidens 86, 95
neglecta 86
pendula 86
triplicata, Alexia micheli var.
triplicata, Pythia 303
triplicata, Voluta 303, 304, 308
triplicatus, Auricula (Conovulus) 308
tristanensis, Marinula 229, 331
tristanensis, Marinula (Marinula) 332
Tritonia hombergi 395
trivolvis, Helisoma 64, 65
Trochoidea 349, 358
Trochus dolabratus 496
troostiana, Millerelix (Prattelix) 91
troostiana, Polygyra 73
trossulus, Mesodon (Mesodon) 93
Truncatella 144
truncatula, Retusa 384, 393, 396, 398-
400, 420, 438
truncatus, Bulinus 17
tuberculata, Melanoides 50
tuberosa, Cassis 23
tubulosa, Holothuria 23
tudiculata, Helminthoglypta 95
tularosana, Ashmunella 88
turbinella, Stenotrema 79
turbinella, Stenotrema exodon 79
turbinella, Stenotrema (Pilsbrelix) 89
Turbo elegantissima 501
gracilis 501
lacteus 501
mirabilis 213
plicatula 500, 501
Turbonilla 459, 500-502, 504
abrupta 501
costulata 501
elegantissima 450, 451, 502
heïlprini 501
hemphilli 445, 446, 450, 458, 464,
473, 474, 477, 478, 481-487, 501-
502

197, 197

535

humboldti 501

incisa 458, 502, 503

jeffreysi 502

Jeffreysi 450

penistoni 501

plicatu'a 500, 501

striata 501

typica 500

unilirata 501
Turbonillinae 444, 448, 450, 478, 500
turritus, Melampus 195, 203
typica, Plecotrema 204
typica, Turbonilla 500

ulla, Ascobulla 385, 388, 405, 439

ulvae, Hydrobia 381

ulyssioponensis, Patella 34

Umbraculacea 387

uncifera, Stenotrema (Pilsbrelix)

unciferum, Stenotrema 79

unciferum, Stenotrema (Stenotrema) 72,
89

undosa, Astraea 491

unicolor, Bellamya 50

unidentata, Odostomia 451

unilirata, Turbonilla 501

Ur-Basommatophora 311

Urbasommatophora 311

usticensis, Helix 349

uvulifera, Daedalochila 81

uvulifera, Daedalochila (Daedalochila) 72,
92,107

107

Vagvolgyrorbis 86 [see Triodopsis
(Vagvolgyrorbis)]
valkyrie, Gegania 377,378, 390, 396,
397, 408, 417, 422, 442
Valvata 400
piscinalis 50, 62
Valvatidae 377,418
vannostrandi, Triodopsis (Triodopsis) 86
variabilis, Helix 359
varicifera, Ashmunella 88
varicosa, Alexia myosotis var.
variegata, Pyramidella 499
variegatus, Heliacus 416
velaini, Marinula 226, 331
velum, Hydatına 389, 398, 418, 433
Veneridae 364
veneta, Auricula 194
venezuelana, Tralia 303, 304, 308, 309
ventricosa, Volvatella 398, 440
ventrosula, Linisa 90
vermiculata, Eobania 2,9
verrillii, Cylichna 435
verticalis, Melampus coffeus var.
265
verus, Inflectarius (Inflectarius) 92
vesicaria, Hydatina 389, 433
Vesicomya 363
Vesicomyidae 363

196

263,

536

Vespericola 83, 84, 86, 93, 106
armigera 86
columbiana 95
columbianus 86
euthales 86
hapla 86
karokorum 86
marinensis 86
megasoma 86
orius 86
pilosus 86
pinicola 87
pressleyi 87
shasta 87
sierrana 87
Vespericolini 84, 86
vespertina, Alexia 196
vespertina, Auricula 195, 197
vetula, Tralia 304
vetula, Tralia (Tralia) 309
vicola, Odostomia (Chrysallida) 489
vigorouxi, Volvatella 398
vincta, Odostomia (Chrysallida) 489
virescens, Haminoea 398, 437
virgata, Cernuella 6, 9
virgata, Physella 345
viridis, Elysia 400
Vitrinellidae 405
viviparus, Viviparus 50
Viviparus contectus 41-52
georgianus 50
malleatus 50
viviparus 50
Voluta bullaoides 276
coffea 249, 256, 257
denticulata 169, 193, 194, 797, 201
flava 293, 294
heteroclita 186
minuta 257, 294, 295
monile 294
pusilla 303, 308
reflexa 194, 197, 202
ringens 194, 197, 202
triplicata 303, 304, 308
Volvatella 376, 377, 382, 383, 384,
395, 398-400, 407-410, 412, 416,
419, 440
bermudae 398-400, 440
cincta 440
ficula 399, 440
fragilis 400, 440
laguncula 440
pyriformis 440
ventricosa 398, 440
vigorouxi 398
volvoxis, Polygyra septemvolva 81
Volvulella persimilis 402
vulgata, Triodopsis 85, 104
vulgata, Triodopsis fraudulenta 85

vulgata, Triodopsis (Shelfordorbis) 85, 95

vultuosa, Triodopsis (Haroldorbis) 86

INDEX

waldense, Stenotrema (Stenotrema) 89
watleyi, Ashmunella 88
watsoni, Auricula 195, 197, 202
watsoni, Scaphander 389, 390, 434
weatherbyi, Fumonelix 92
Webbhelix 78, 83, 84

multilineata 95
wheatleyi, Fumonelix 92, 94
wheatleyi, Helix 92
wichitorum, Euchemotrema 88
willisi, Odostomia (Chrysallida) 488
Wingenella eburnea 497

pricena 497

Xanthonychidae 95

xanthostoma, Marinula 217, 225, 226,
230, 331

xanthostoma, Marinula (Cremnobates)
332

Xeroamanda 349-361

Xerocincta 357,359

Xerolenta obvia 1

Xeromagna 358

Xeromunda 358

Xerophila 358

Xeropicta arenosa 6, 9

Xeroplana 350

Xerosecta 358, 359

Xolotrema 78,84,85, 101
denotata 104
fosteri 71, 97, 103

Xolotrema (Wilcoxorbis) 83, 84, 101
fosteri 85, 95, 97
occidentalis 85

Xolotrema (Xolotrema) 83-85
carolinensis 85
denotata 85, 95
obstricta 85

yucatanea, Linisa 90

zaletus, Mesodon 104
zaletus, Mesodon (Aphalogona) 93
zelandiae, Haminoea 419, 437
zonata, Sphincterochila 9
Zospeum 176, 313

spelaeum 175

MALACOLOGIA

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VOL. 37 MALACOLOGIA

CONTENTS

J. D. ACUNA & M. A. MUÑOZ
А Taxonomic Application of Multivariate Mixture Analysis in Patellidae

KATERINE COSTIL & JACQUES DAGUZAN

Comparative Life Cycle and Growth of two Freshwater Gastropod Species,

Planorbarius Corneus (L.) and Planorbis Planorbis (L.) .......................
GEORGE M. DAVIS, ZHANG YI, GUO YUAN HUA 8 CHRISTINA SPOLSKY

Population Genetics and Systematic Status of Oncomelania Hupensis (Gas-

tropoda: Pomatiopsidae) Throughout China
M. LAZARIDOU-DIMITRIADOU

The Life Cycle, Demographic Analysis, Growth and Secondary Production of the

Snail Helicella (Xerothracia) Pappi (Schutt, 1962) (Gastropoda Pulmonata) in E.

Macedonia (SISSCe) ооо anse di ce diia AS O et
М. A. EDWARDS 4 М. J. THORNE

Response to Bouchet 8 Rocroi; ‘The Lottery of Bibliographical Databases:

RS TOC Wands Qo TRONO "asadas a ees
М. ELEUTHERIADIS & М. LAZARIDOU-DIMITRIADOU

The Life Cycle, Population Dynamics, Growth and Secondary Production of the

Snail Viviparus Contectus (Millet) (Gastropoda: Prosobranchia) in the Marshes of

the River Strymonas, Serres, Macedonia, Northern Greece ..................
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When Shells Do Not Tell: 145 Million Years of Evolution in North America's

Polygyrid Land Snails, with a Revision and Conservation Priorities ...........

A. HONEK

Geographic Distribution and Shell Colour and Banding Polymorphism in Mar-

ginal Populations of Cepaea Nemoralis (Gastropoda, Helicidae) ..............
MICHAEL J. KENNISH, ANTONIETO S. TAN, & RICHARD А. LUTZ

Shell Microstructure of Vesicomyid Clams from Various Hydrothermal Vent and

еее ЯК ДЕ = Пе о aan
GIUSEPPE MANGANELLI, LEONARDO FAVILLI & FOLCO GIUSTI

The Taxonomic Status of Xeroamanda Monterosato, 1892 (Pulmonata,

FAY COMMAS) na ra vas es Ma e
ANTONIO M. De FRIAS MARTINS

Anatomy and Systematics of the Western Atlantic Ellobiidae (Gastropoda:

аа
PAULA М. MIKKELSEN

The Evolutionary Relationships of Cephalaspidea S.L. (Gastropoda: Opistho-

branchia)-A:PhylogenettG AnalVSiS coros rosas 020. 0000 Rene a
HILARY PIGGOTT & GEORGES DUSSART

Egg-Laying and Associated Behavioural Responses of Lymnaea Peregra

(Müller) and Lymnaea Stagnalis (L.) to Calcium in their Environment ..........
LUIZ RICARDO LOPES DE SIMONE

Anatomical Study on Tonna Galea (Linné, 1758) and Tonna Maculosa (Dillwyn,

1817) (Mesogastropoda, Tonnoidea, Tonnidae) from Brazilian Region. ........
DONALD L. THOMAS 8 JAMES В. McCLINTOCK

Aspects of the Population Dynamics and Physiological Ecology of the Gastro-

pod Physella Cubensis (Pulmonata: Physidae) Living in a Warm-Temperate

steam and Ephemeral Pond. Habitat: sion cise азов on as nales

1996

33

53

133

157

41

69

111

363

349

163

375

13

23

333

В. VITTURI, A. LIBERTINI, М. PANOZZO & G. MEZZAPELLE
Karyotype Analysis and Genome Size in Three Mediterranean Species of Peri-
winkles (Prosobranchia: Mesogastropoda) ................................ 123
JOHN B. WISE
Morphology and Phylogenetic Relationships of Certain Pyramidellid Taxa
(Heterobranchia) cusco en ae ann an ale AR 443

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VOL. 37,N0.2 - - MALACOLOGIA | by
CONTENTS N . > май En E ¿oe
148 | NR iy
ANTONIO M. pe FRIAS MARTINS.
J Anatomy and Systematics of the Western Atlantic. Ellobiidae (Gastropoda:
у’ Pulmönatal ade do de aa ae clos eects RS DE ВВ at
DONALD Е. THOMAS, & JAMES В. McCLINTOCK . We x
и Aspects of the Population Dynamics and Physiological Ecology of the Gastro-
Pp pod Physella Cubensis (Pulmonata: Physidae) Не in а Warm- Temperate _
| Stream and Ephemeral Pond Habitat SO 2 to и.
GIUSEPPE MANGANELLI, LEONARDO FAVILLI & FOLCO GIUSTI = INES

| e: branchia): lg LR SM We PRE Pr PE corse cs ar _ 375
JOHN В. WISE | a 12
Morphology and Phylogenetic’ Relationships of Certain Pyramidellid Taxa pos

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