THE OPERCULA OF NERITOPSID GASTROPODS AND THEIR

PHYLOGENETIC IMPORTANCE
ANDRZEJ KAIM 1 AND PRZEMYSLAW SZTAJNER 2 =

1
Instytut Paleobiologii PAN, ul. Twarda 51/55, PL-00–818 Warszawa, Poland;
2
Zaklad Geologii i Paleogeografii, Instytut Nauk o Morzu, Uniwersytet Szczeciński, ul. Felczaka 3a, PL-71–412 Szczecin, Poland
=

(Received 27 May 2004; accepted 8 December)

ABSTRACT
The fossil record of neritopsid opercula and shells shows that the shell shape typical for Neritopsidae and
Neritidae appeared in the Triassic. The ancestors of Neritimorpha were most probably forms similar to
Naticopsis. The operculum of Recent Neritopsis is composed of two calcitic parts secreted from inside and
an aragonitic callus deposited from outside. Similar neritopsid opercula were already present in the
Late Triassic. The first opercula with asymmetrically situated muscle scars, possibly ancestral for
neritids, also appeared at that time.

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INTRODUCTION
The strongly convolute protoconch was believed by Bandel &
Frýda (1999) to be the most important shell character uniting
two extant families (Neritopsidae and Neritidae) of the subclass
Neritimorpha (¼Neritopsina). According to Bandel & Frýda
(1999), this is the neritimorph trait with the longest fossil
record. Other characters commonly used for high-level
taxonomic purposes are the microstructure of the shell and
operculum and the morphology of the latter (Ponder, 1998).
Although the operculum of Neritopsis radula has been known
since the nineteenth century (e.g. Crosse, 1875; Souverbie &
Montrouzier, 1875; Fischer, 1887), in recent papers the opercu-
lum has only been illustrated in a little-known paper by Jagt &
Janssen (1988). This contribution has apparently had only
limited public exposure (e.g. not known to Checa & Jiménez-
Jiménez, 1998) and even in the recent reviews (e.g. Ponder,
1998), the illustrations are copied from nineteenth century
monographs (e.g. Fischer, 1887). The aim of this paper is to
summarize the available data on the evolution of neritopsid
opercula and shells.
Although the Neritidae are common in various Recent,
especially shallow water environments (e.g. Bandel, 2001;
Sasaki, 2001), the Neritopsidae are a relic family represented
only by two marine species (Ponder, 1998) known to live in
submarine caves or deeply buried under coral rubble (Kase &
Hayami, 1992; Holthuis, 1995). The information from soft
parts of this crucial group of the Neritimorpha is still hardly
exploited (Holthuis, 1995; Kano, Chiba & Kase, 2002), but it
suggests that Neritopsidae are related to the shell-less, slug-like
Titiscaniidae (Kano et al., 2002). The latter family has even
been synonymized with Neritopsidae by Kano et al. (2002)
from the standpoint of molecular phylogenetics. The internal
coiling of the Neritopsis shell is an ancestral state, while the
larval development and radula are derived.
The fossil record of Neritopsis-like gastropods ranges back to
the Triassic (Knight et al., 1960; Zardini, 1978) but they were
most common during the Jurassic and Cretaceous (e.g.
Brösamlen, 1909; Kase & Maeda, 1980; Hägele, 1997). The
shell of Neritopsis (Fig. 4A– C) is composed of two wholly aragon-
itic layers (Suzuki, Togo & Uozumi, 1991). The inner shell wall
is not resorbed during ontogeny, so the columella is present. The
opercula of Neritopsidae were described by Bandel (1990) as cal-
citic but this has never been reported in detail. The neritopsid
Correspondence: A. Kaim; e-mail: kaim@twarda.pan.pl
operculum has a distinctive trapezoidal shape. Such opercula
have been known for a long time (e.g. Crosse, 1875; Souverbie
& Montrouzier, 1875; Fischer, 1887; Jagt & Janssen, 1988)
although they have usually been only poorly illustrated.
The shell of Neritidae is composed of three calcareous parts:
the outer part is calcitic, whereas the middle and inner ones
are aragonitic (Gainey & Wise, 1980; Suzuki et al., 1991). The
neritids dissolve the inner wall of their shell, which lacks the
columella as a result. The operculum is spirally coiled, semi-
circular and fully aragonitic (Suzuki et al., 1991; Sasaki, 2001).
It also bears a peg-like apophysis, a structure expanding the
surface of muscle attachment (Ponder, 1998; Sasaki, 2001).
Due to this modification, the muscle scars are asymmetrical.
The fossil record of undoubted neritid shells ranges back to the
Jurassic (Schmidt, 1905; Figs 4I, J), possibly even to the Triassic
(Kittl, 1892).
MATERIALS AND RESULTS
The neritopsid opercula described and illustrated in this paper
came from three different time levels. The silicified opercula of
Jurassic Neritopsis subvaricosa Brösamlen, 1909 and associated
inner/outer moulds of the shells were found in an abandoned
quarry of the early Oxfordian (Late Jurassic) micritic limestone
at Wrzosowa (Czestochowa region, Poland). The material is
represented by five‘ silicified opercula, 30 inner and two outer
shell moulds. No opercula were found preserved in place inside
the aperture, but the connection of the opercula to these shells
is based on the co-occurrence of only these two neritimorph
fossils at the locality. The material is housed at the Museum of
Szczecin University (MSzcz) and Institute of Paleobiology
PAS in Warsaw (ZPAL).
The opercula of unidentified neritopsid gastropods from the
Quadrata Zone, Early Campanian (Late Cretaceous) from the
Downend Quarry in Portsmouth, England has been deposited
by A. Gale at the Natural History Museum in London
(BMNH). In addition, some Jurassic neritopsid opercula
housed at that Museum has been examined. Details of these
additional opercula have not been provided due to the poor
state of their preservation.
To illustrate the juvenile whorls of the shell and operculum
morphology and microstructure of the extant Neritopsis radula,
material housed at the Naturhistoriska Riksmuseet in
Stockholm, Sweden has also been examined (SMNH).
To study their microstructure, the opercula were embedded in
epoxy resin, cut and polished. They were then etched in 1% HCl

Journal of Molluscan Studies (2005) 71: 211– 219. Advance Access Publication: 28 June 2005 doi:10.1093/mollus/eyi029
# The Author 2005. Published by Oxford University Studies on behalf of The Malacological Society of London, all rights reserved.
 A. KAIM AND P. SZTAJNER
for 7 min, sputtered with platinum and examined under a
Philips XL-20 scanning electron microscope.
For determination of the aragonitic and calcitic parts of the
operculum, we used a simple method of staining with Feigl sol-
ution (Leitmeier & Feigl, 1934). The operculum section was
covered by a drop of Feigl solution (a mixture of silver sulphate
and manganese sulphate with dilute sodium hydroxide sol-
ution). After a few minutes the aragonitic part turns grey,
while the calcite part remains unchanged.
Operculum of the Late Jurassic Neritopsis subvaricosa
The operculum from the early Oxfordian of Wrzosowa is con-
centric, bowl-like trapeziform with the narrower part oriented
adaxially in the shell peristome (Fig. 1A – C). Its inner surface
shows the abaxial semilunar zone and adaxial elevated projec-
tion. The central field of this projection is somewhat depressed
and ornamented with an indistinct radial striation and
concentric growth increments. In the centre of the inner part
of the operculum a nucleus is visible. The muscle scars are
located symmetrically on both sides of the inner surface of
the operculum.
The outer surface of the operculum has an elevated adaxial
margin and a conical depression at the place where the
nucleus is visible. The depression bears distinctive growth
lines. The abaxial part of the operculum is much thinner and
also bears growth lines. The shell of N. subvaricosa was described
and illustrated by Brösamlen (1909) and our material (Fig. 4E–
H) does not provide any new information about the shell.
The primary microstructure of the opercula from Wrzosowa
has been altered by silification. The aragonitic shells of
N. subvaricosa have been dissolved during diagenesis of the
rock, whereas the opercula have been substituted by silica.
This type of preservation is typical for Neritopsis-like gastropods
and even findings of well-preserved fossil opercula attached to
the apertures of internal moulds of shells have been reported
(see Brösamlen, 1909: pl. 19, fig. 18). This indirect evidence
suggests that the shell was aragonitic (like extant Neritopsis),
while the operculum was primarily calcitic.
Neritopsid operculum from the Early Jurassic of France
The opercula from Torcian (Early Jurassic) of May in
Normandy, France (Fig. 1G, H) are generally similar to the
Late Jurassic specimen from Wrzosowa described above. The
most important difference is that the outer surface of the
Torcian operculum is covered by a smooth layer similar to the
outer callus of the Recent Neritopsis radula (see below).
Neritopsid operculum from the Late Cretaceous of England
The operculum from Early Campanian chalk of the Downend
Quarry in Portsmouth is trapeziform, axially elongated
(Fig. 1I– K). As in the Jurassic specimen from Wrzosowa, the
inner surface also shows the abaxial semilunar zone and
adaxial projection, but it submerges under the abaxial part of
the operculum at one-third of the operculum width. The projec-
tion has one central, elongated elevation and two depressions on
both sides. Unlike its geologically older relative, the whole
internal surface of the projection is ornamented, in this speci-
men, with a distinct radial striation. Most important is the
finding that the muscle scars are located in shallow pockets,
symmetrically placed on both sides of the central projection.
Similarly to the Late Jurassic operculum, the outer surface is
uneven and bears distinctive growth lines. No shell correspond-
ing to this operculum is known.
In cross-section, the operculum appears to be composed of two
mineral parts. Staining with Feigl solution (Leitmeier & Feigl,
212
1934) suggests that both parts are composed of calcite. The
matrix is highly porous and in places the original microstructure
is altered by diagenesis (Fig. 2B). In the best preserved part of
the operculum the calcite is composed of needles typical of
fibrous prismatic structure (Fig. 2D). The two mineral parts
are separated by a thin organic layer (Fig. 2C).
Operculum of the Recent Neritopsis radula
The operculum is trapeziform and weakly elongated axially
(Fig. 1D – F). The inner surface shows the abaxial semilunar
zone and adaxial projection. The projection submerges under
the abaxial part of the operculum at half the operculum
width. The projection has a wide central depression ornamented
with thick radial ribs. Similarly to its Cretaceous predecessors,
the muscle scars are located in pockets symmetrically placed
on both sides of the central projection.
The outer surface of the operculum is covered with a callus
and growth lines are invisible apart from the abaxial-most
part of the operculum. The adaxial margin shows a shallow
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depression in its central part.
The operculum of Recent Neritopsis is composed of three
mineral parts. Staining with Feigl solution (Leitmeier & Feigl,
1934) suggests that internal parts of the operculum (cl1, cl2)
are composed of calcite, while the external callus (al) is aragon-
itic. The external aragonitic callus (al) and the calcitic part of
the operculum above the muscle attachment surface (cl2;
Fig. 2A) are composed of needles typical of fibrous prismatic
structure (Fig. 3C, G), while the calcitic tissue covering the
muscle attachment from below (cl1; Fig. 2A) consists of first-
order prisms composed of second-order elongated prisms, a
pattern typical for composite prismatic structures (Carter
et al., 1990). The secretion of the calcitic operculum starts with
a single unit (cl), which later becomes separated with an
organic (ol) layer marking the muscle attachment area orna-
mented with radial ribs and concentric increments (Fig. 3A).
After development of the muscle attachment, its migration
during the growth of the operculum leaves an organic intercala-
tion (ol) separating the adaxial (cl1) part of the operculum
(located below the intercalation) from the abaxial (cl2) one
(located above).
DISCUSSION
Most of the fossil neritopsid opercula we examined are fully cal-
citic, while the extant Neritopsis radula has an additional aragon-
itic callus secreted from outside. This difference is artificial, as
the outer layer composed of aragonite, the less stable crystallo-
graphic form of calcium carbonate, usually dissolves during dia-
genesis. The outer surface of the operculum after removal of the
aragonitic layer is uneven and shows the growth lines of the
underlying calcitic layer. The surface of the callus is smooth,
which indicates that it was completely covered by the secretory
soft tissue. Exceptionally, the callus may be preserved in strata
that contain unaltered aragonitic fossils. Such occurrences
from Carnian (Late Triassic) of St Cassian in Italy (Zardini,
1978) and from Toarcian (Early Jurassic) of May in France
(Fig. 1G, H) are known.
The protoconch of Neritopsis radula
In their review of Neritimorpha Bandel & Frýda (1999: pl. 1,
fig. 2) illustrated a protoconch from a shell identified as Neri-
topsis radula. This protoconch from Mauritius is similar to the
ones of Nerita and Neritina (e.g. Bandel & Frýda, 1999: pl. 1,
fig. 5), but not to the protoconch of N. radula from New
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Figure 1. Neritopsid fossil and Recent opercula. A–C. Operculum (MSzcz S.365) of Neritopsis subvaricosa Brösamlen, 1909 from the early Oxfordian,
Late Jurassic of Wrzosowa, Czestochowa region, Poland. A. SEM stereographs of the inner surface. B. SEM stereographs of the outer surface. C. Side
view. D–F. Operculum (SMNH ‘ 52419/1) of Neritopsis radula Linné, 1758 from off Mauritius, Indian Ocean. D. SEM stereographs of the inner surface.
E. SEM stereographs of the outer surface. F. Side view. G, H. Neritopsid opercula (BMNH PI MG 1111 and 1112 respectively) from the Toarcian
(Early Jurassic) of May, Normandy, France. G. Inner surface. H. Outer surface. I–K. Neritopsid operculum (BMNH PI MG 1115) from the
Early Campanian (Late Cretaceous) of Downend Quarry in Portsmouth, England. I. Side view. J. Inner surface. K. Outer surface.

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Figure 2. A. Operculum (SMNH 52419/1) of Recent Neritopsis radula Linné, 1758 from off Mauritius, Indian Ocean; longitudinal section. B–D.
Neritopsid operculum (BMNH PI MG 1114) from Early Campanian, Late Cretaceous of Downend Quarry in Portsmouth. B. Longitudinal
section, dissolved outer aragonitic layer marked with dashed line. C. First calcitic part (cl1) and second calcitic part (cl2) separated by organic
layer (ol). D. Needles of second calcitic layer (cl2).

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Figure 3. Operculum (SMNH 52419/1) microstructures of recent Neritopsis radula Linné, 1758 from off Mauritius, Indian Ocean. A. Initial part of the
operculum showing an initial inner calcitic part (cl) that bifurcates into two calcitic parts (cl1 and cl2) separated by organic layer (ol). B. The inner
calcitic part (cl) covered by outer aragonitic part (al). C. Aragonitic, fibrous-prismatic outer part (al). D. Border between aragonitic, fibrous-prismatic
outer part and first calcitic, composite-prismatic inner part (cl1). E. First calcitic, composite-prismatic inner part (cl1). F. First calcitic, composite-
prismatic inner part (cl1) and second calcitic fibrous-prismatic inner part (cl2) demarcated by organic layer (ol). G. Second calcitic fibrous-prismatic
inner part (cl2). H. Surface of organic layer (ol) with imprinted prism endings of first calcitic, composite-prismatic inner part (cl1). All figures, apart
from H, are in the same orientation as whole operculum section shown in Figure 2A.

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Figure 4. Shells of neritimorph gastropods. A. Neritopsis radula Linné, 1758 from off Mauritius, Indian Ocean; shell (SMNH 52419/2) with an oper-
culum in situ. B– D. Neritopsis radula Linné, 1758 from Banc de Touhou, off New Caledonia. B. Shell (SMNH 52419/3) in lateral view. C. Shell (SMNH
52419/4) in apical view. D. Close-up of protoconch. E–H. Shells (MSzcz S.367–8) of Neritopsis subvaricosa Brösamlen, 1909 from Oxfordian (Late Jur-
assic) of Wrzosowa, Czestochowa region, Poland. E. Rubber cast of the outer mould. F. Side view of inner mould. G. Rubber cast of outer mould
(MSzcz S.368). H. Apical ‘ view of inner mould (MSzcz S.367). I, J. Shell (ZPAL Ga.12/1) Lissochilus concinna (Roemer, 1836) from the late Oxfordian,
Late Jurassic of Kleby in Western Pomerania, Poland. I. Apertural view. J. Apical view. B– D courtesy of A. Warén.
=

Caledonia in the collection examined by ourselves (Fig. 4D) at
the Naturhistoriska Riksmuseet in Stockholm. The New Cale-
donia specimen is clearly non-planktotrophic with no initial
whorl visible (Fig. 4D). It closely resembles Eocene Neritopsis par-
isiensis Deshayes, 1864 illustrated by Bandel & Frýda (1999: pl.
1, fig. 4) and N. radula from Yonaguri Island, Okinawa, illus-
trated by Kano & Kase (2000). As specimens of N. radula from
off Mauritius and Java housed at the Naturhistoriska Riksmu-
seet in Stockholm are also apparently non-planktotrophic (A.
Warén, personal communication), it is possible that the gastro-
pod from Mauritius illustrated by Bandel & Frýda (1999) rep-
resents another species or even belongs to another genus (e.g.
Nerita ). Unfortunately Bandel & Frýda (1999) illustrated only
the apical part of the shell, making identification difficult.
Evolution of the neritopsid operculum
The concentric, trapeziform, neritopsid operculum is unique
among Gastropoda (Sasaki, 2001) at least since the Late Triassic
(Zardini, 1978). The opercula of all other Neritimorpha (apart
from the terrestrial Helicinidae) are composed of aragonite
(Suzuki et al., 1991). Although Helicinidae have calcitic oper-
cula without an apophysis, calcification occurs only on the
outer surface and the inner part is organic (Suzuki et al., 1991,
Sasaki, 2001).
The neritopsid operculum seems to have evolved from a tear-
drop-shaped, concentrically-coiled operculum with a single,
large muscle scar. Such opercula are reported to occur in the
Early Carboniferous (Mississippian) gastropod Naticopsis

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Figure 5. Possible relationship between fossil and extant Neritimorpha. A. Opercula in inner view. Operculum of Naticopsis genevievensis Meek &
Worthen, 1867 redrawn from Gordon & Yochelson (1982a); opercula form 1 and 2 redrawn from Linsley et al., (1989); naticopsid and neritopsid oper-
cula of Zardini, 1978, redrawn from Zardini (1978); operculum of Neritopsis subvaricosa Brösamlen, 1909, redrawn from the original (see Fig. 1A) from
Wrzosowa, Czestochowa region, Poland; neritopsid operculum from Downend Quarry redrawn from original (see Fig. 1J); operculum of Nerita albicilla
‘
Linné, 1758 simplified after Sasaki (2001); operculum of Neritopsis radula Linné, 1758 redrawn from original from off Mauritius (see Fig. 1D). B. Shells.
Shell of Naticopsis buttsi Gordon & Yochelson, 1982, redrawn from Gordon & Yochelson (1982b); shell of Trachydomia nodosa (Meek & Worthen, 1861),
redrawn from Knight et al. (1960); shell of Trachyspira delphinuloides Gemmelaro, 1889, redrawn from Knight et al., (1960); Lissochilus concinna (Roemer,
1836), redrawn from original (see Fig. 4I) from Kleby in Western Pomerania, Poland; shell of Neritopsis subvaricosa Brösamlen, 1909, redrawn from
=

Brösamlen (1909); shell of Nerita albicilla Linné, 1758,‘ simplified after Sasaki (2001); shell of Neritopsis radula Linné, 1758, redrawn from Ponder (1998).

(Gordon & Yochelson, 1982a, b). Similar opercula are also
known from the late Carboniferous (Pennsylvanian; Form 4 of
Linsley et al., 1989) of the United States through the Triassic
(Kittl, 1892). In the Pennsylvanian, opercula with a curved
muscle scar with incipient subdivision into two lobes (Form 1
of Linsley et al., 1989) and opercula with two distinctive lobes
of a single muscle scar (Form 2 of Linsley et al., 1989) appeared
for the first time. It is probable that, subsequently, the muscle
bifurcated into two separate muscles, as such scars are pro-
nounced in the Late Triassic opercula illustrated by Zardini
(1978) and the Jurassic opercula illustrated herein (Fig. 1A–
C, G, H). Possibly the opercula of form 1 belonged to
Trachydomia (Linsley et al., 1989), the genus postulated by
Batten (1984) to be ancestral for Neritopsis.
The muscle scars of Triassic and Jurassic neritopsid opercula
are two flat and large circular fields. In the Cretaceous the
fields have been drawn in under the thickening and expanding
inner calcitic layer of the operculum. As a result, sockets for
muscle attachment developed. In the extant Neritopsis the
sockets are deeper and narrower.
Relationship of Neritopsidae with Neritidae and the fossil record
The Neritidae have protoconchs with strongly adpressed whorls
with hardly visible suture and the initial whorl not emerging
over the protoconch surface (e.g. Bandel & Frýda, 1999;
Bandel, 2001; Warén & Bouchet, 2001), a form that is rare
among the Gastropoda (Bandel, 1992). It is highly unlikely
that such a protoconch with no internal coiling developed
twice independently. All the neritopsid protoconchs examined
during this study are non-planktotrophic (Fig. 4D). Bandel &
Frýda (1999) identified a neritid-like protoconch as belonging
to Neritopsis radula, but this identification is doubtful. In such a
situation it is difficult to interpret whether the common ancestor
of Neritopsis and Nerita already had this kind of larval

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 A. KAIM AND P. SZTAJNER
development. The earliest gastropods with a similar protoconch
are known from the Late Triassic (Bandel, 1992). The absence of
a calcitic wall on their teleoconch may suggest that they are
related to the Neritopsidae rather than the Neritidae. During
the Triassic, neritid gastropods, able to dissolve the internal
wall of the shell, appeared (Kittl, 1892; Cossmann, 1925).
Such gastropods are common in Jurassic shallow-water
limestone is (Schmidt, 1905; Dmoch, 1971; see Fig. 4I, J). The
opercula typical of Neritidae (aragonitic, paucispiral with
apophysis) remain unknown in the Jurassic and Cretaceous,
because aragonitic fossils are generally rare in ancient strata.
The exceptional gastropod collection of Zardini (1978) from
Carnian (Late Triassic) of the Italian Alps reveals a wide
range of opercular morphologies preserved with aragonitic
layers. Among others, there are also opercula with asymmetric
muscle scars (Fig. 5). Possibly, this operculum is an early
member of the lineage leading to the Neritidae.
Bandel & Heidelberger (2001) described a Devonian paucis-
piral operculum found in situ in the neritoid shell of Hessonia pili-
gera (Sandberger & Sandberger, 1856). Unfortunately, only the
external view of the operculum is available (Bandel & Heidel-
berger, 2001; Heidelberger, 2001). It is unknown whether the
operculum has an apophysis and whether the calcium carbonate
was secreted on the inner surface of the operculum, which is
typical for the Neritidae (Sasaki, 2001).
Disregarding Hessonia piligera, the relationship between Neri-
topsidae and Neritidae can be summarized as in Figure 5. The
ancestors of both groups are proposed to be Naticopsis-like, oper-
culate gastropods, possibly with a protoconch like those of early
caenogastropods (e.g. Dzik, 1994: fig. 30H, I). The uniquely
neritid type of protoconch possibly appeared in the latest
Palaeozoic or the Triassic. Both groups diverged from the
Naticopsis-Trachydomia stem in the Triassic. The Neritopsidae
developed a concentric, calcite-aragonitic operculum with two
separate, symmetrical muscle attachments and an aragonitic
shell without dissolution of the inner wall. The Neritidae devel-
oped a wholly aragonitic semilunar operculum with apophysis
and asymmetrical attachment scars, in addition to a shell with
an outer calcitic wall and the ability of resorption.
Bandel & Frýda (1999) subdivided Neritimorpha
(¼Neritopsina) into two new orders. The Cycloneritimorpha,
encompassing true neritimorphs with Nerita-type protoconchs
and Cyrtoneritimorpha, with openly coiled protoconchs,
rooted by Bandel & Frýda (1999) in the Ordovician. The
latter order is composed of two families: the platyceratid-like
Orthonychiidae and the poorly known Early Devonian monoge-
neric Vltaviellidae (cf. Frýda & Manda, 1997; Bandel & Frýda,
1999). Bandel & Frýda (1999: fig. 3) placed Platyceratoidea in
the Cycloneritimorpha, as the only Palaeozoic member of the
order. They separated platyceratids from the Orthonychiidae,
although these two contemporary groups have a unique, para-
sitic mode of life on pelmatozoans (see Baumiller, 2002, for
review). This grouping seems to be highly conjectural as no
well-preserved protoconch of Platyceras has been found and the
shells are not similar. Moreover, there is no indication that
any neritimorphs feed on crinoids. Although the taxonomic pos-
ition of the platyceratid gastropods lies outside the scope of this
paper, it has to be pointed out that the type of protoconch found
in Orthonychia by Bandel & Frýda (1999) suggests an euomphalid
origin for the platyceratids (Kaim, 2004). The true neritopsid-
like gastropods were probably derived from early gastropods
with planktotrophic development, which are commonly found
in the Palaeozoic fossil record (e.g. Dzik, 1978, 1994; Yoo,
1988, 1994).
Pseudorthonychia and related Triassic forms (e.g. Palaeonarica of
Bandel, 1992) can be included in the family Cortinellidae with
convolute protoconch and planispiral teleoconch (Bandel,
2000). This family is possibly intermediate between
218
Neritopsinidae and Neritidae in having the ability to resorb
the internal wall in the protoconch but lacking the calcitic
outer wall of the teleoconch (Bandel, 2000). The ancestors of
Neritopsis did not resorb their inner whorls as neritids do. The
members of the family Cortinellidae are known exclusively
from the Triassic.
CONCLUSIONS
The protoconch of Late Palaeozoic Naticopsis has never been
reported in detail, but in the Silurian it was similar to proto-
conchs of contemporary caenogastropods (Dzik, 1994). The ner-
itimorph mode of larval development possibly appeared in the
Permian or perhaps the Triassic, prior to or just after the diver-
gence of the Neritopsidae and Neritidae. Since that time, the
shell and operculum of Neritopsinidae have not changed sub-
stantially. The symmetrical operculum of Neritopsinidae is com-
posed of two internal calcitic parts and a single outer aragonitic
part, while the asymmetrical operculum of Neritidae is exclu-
sively aragonitic.
The evolutionary success of Neritidae may lie in the strength-
Downloaded from http://mollus.oxfordjournals.org/ by guest on December 5, 2013
ening of the apertural closure, manifested by enlargement of the
retractor attachment fields. This was apparently a modification
against predatory attacks, most probably by crabs. It could also
help prevent desiccation during low tide. Increased predation
pressure may have led to the gradual disappearance of the
conservative Neritopsidae from their normal environments,
and their shift to cryptic habitats.
ACKNOWLEDGEMENTS
We are grateful to J. Dzik for identification of the Jurassic
operculum from Wrzosowa and making us aware of its import-
ance. The examination of Neritopsis radula housed at the
Naturhistoriska Riksmuseet in Stockholm, Sweden was possible
due to the HighLat Programme (European Community –
Access to Research Infrastructure Action of the Improving
Human Potential Programme) to A. Kaim under the kind gui-
dance of A. Warén, who also supplied some pictures of Recent
Neritopsis radula. The examination of Jurassic and Cretaceous
Neritopsis opercula at the Natural History Museum in London
was possible due to the SysResource Programme (European
Community – Access to Research Infrastructure Action of the
Improving Human Potential Programme) to A. Kaim under
the kind guidance of J. Todd. We are deeply indebted to
J. Dzik and A. Warén for discussion and many helpful hints
during the preparation of this paper. We are also grateful to
T. Baumiller, J. Dzik and A. Warén who kindly reviewed this
paper. The paper greatly benefited from two anonymous
reviews. T. Wesolowska is acknowledged for preparing the
=
Feigl solution. The rubber casts were made by W. Siciński and
some of optic photographs were taken by G. Dziewińska and
M. Dziewiński. C. Kulicki and Z. Strak are acknowledged for
‘
their assistance in preparing the operculum section.
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