Paleontological Society

Historic and Current Considerations for Revision of Paleozoic Gastropod Classification

Author(s): Ellis L. Yochelson
Source: Journal of Paleontology, Vol. 58, No. 1 (Jan., 1984), pp. 259-269
Published by: SEPM Society for Sedimentary Geology
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 JOURNAL OF PALEONTOLOGY, V. 58, NO. 1, P. 259-269, 1 FIG., JANUARY 1984

HISTORICAND CURRENT CONSIDERATIONSFOR REVISION

OF PALEOZOICGASTROPODCLASSIFICATION
ELLISL. YOCHELSON
U. S. Geological Survey,Washington,D.C. 20560
ABSTRACT-The notion that a parallel-sided slit in the outer lip of a shell indicates the presence of two
gills has been a significantconceptin classificationof Paleozoicgastropods.However,a deep backward-
curvingaperturalnotch may also indicate two gills. The Ordoviciantaxa Lesueurillaand Lecanospira
have such a notch and thereforewere pleurotomariaceans,in at least a functionalsense. Historically
they were never judged to be Pleurotomariaceaand if they are transferred,the subordercontaining
this superfamilyshould be expandedand recast.Removal of these generafrom the Euomphalaceaand
Macluritacea,respectively,furtherweakensany assignmentof both of those superfamiliesto the same
suborder,a taxonomic arrangementwhich has been suspect for some time. The Euomphalaceamay
have been derived from Lecanospira-likepleurotomariaceangastropods.The Macluritaceahave been
reducedin content by other recent investigationsand are judged even more restrictedan offshoot of
the main stock of gastropodevolution than they were in past classifications.

INTRODUCTION fills in with shell material, while retaining its

REDISCOVERYOF a significant pleurotomari- same relative position behind the rest of the
acean gastropod feature in a shell form atyp- outer lip. The shell area where the slit has
ical of the pleurotomariaceans has interesting moved forward is a band, called simply
ramifications for higher level classification enough the "slitband."
within the Paleozoic Gastropoda. A revised In a shell that has the outer lip broken back,
classification formalizing the views discussed one cannot see the slit, but interruption of
below would be useful, but presentation of the growth lines marks its position on the side
the concept in a preliminary form will allow of the shell. In some fossil pleurotomari-
others to review the data critically, before any aceans, the growth increments added to the
major revisions are undertaken. To put the posterior of the slit may be coarser than the
new interpretation into perspective, some growth lines and thus more prominent. In
historical data are given to explain why the shape they are crescentic, duplicating the
present-day classification has evolved the way round end of the slit. In token of this, the
it has. This background provides some no- term "slitband" was replaced, by J. B. Knight,
tion of the amount of preconception that one in the literature with the more elegant word
can bring to a study, a point that applies to "selenizone," to denote the successive cres-
this work as well as to my critique of older centic lunulae that succesively fill in the pos-
concepts. terior end of the slit. It may be argued that a
selenizone is a special kind of a slitband, for
SLIT AND SELENIZONE IN GASTROPODS
none of the living pleurotomariids have ru-
Trochus and most other living conical gas- gose growth lines and many fossil pleuroto-
tropods have a simple straight outer lip on mariacean gastropods have such faint growth
the aperture. Living pleurotomariid gastro- lines the shell appears smooth. A selenizone,
pods also have a conical trochiform shell, but in this highly specialized sense of a structure
a key morphological feature in these gastro- bearing prominent lunulae, is more common
pods is the slit, a prominent emargination in in late Paleozoic gastropods than it is in mid-
the outer lip which has straight sides parallel dle Paleozoic ones, and it appears to be rel-
for some distance back from the general area atively rare among early Paleozoic forms. The
of the outer lip. Water, which has passed over relative prominence of growth lines among
the gills paired on either side of this shell various families of gastropods has never been
feature, and the contents of the anal tube are studied and it is not clear what meaning, if
discharged through the slit. Paired gills are any, is to be attached to more prominent
generally considered a primitive condition of growth lines.
gastropods. As growth increments are added The slitband may be slightly depressed be-
to the aperture, they also are added to the low the general level of the whorl, flush with
rounded end of the slit so that it continually the whorl surface, or raised as a welt; it may
Copyright ? 1984, The Society of Economic 259 0022-3360/84/0058-0259$03.00
Paleontologists and Mineralogists and
The Paleontological Society

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260 ELLIS L. YOCHELSON
be bordered by parallel spiral lirae or it may
not be set off. The slitband is not always easy
to observe when strong lunulae are absent,
particularly so on a gastropod having faint
growth lines on a flush nonbordered seleni-
zone.
Although the slitband is fairly well known
in many fossil genera, little detail is available
on the slit itself, for it is rare to find a fossil
gastropod with a complete unbroken aper-
ture. One generalization is that the slit is most
commonly near the mid-whorl; in those pleu-
rotomariaceans having an angulated whorl
profile, the selenizone is commonly on the
angulation. Another generalization is that
though the lip may be symmetrical imme-
diately adjacent to the slit, a short distance
away from the slit the lower part of the ap-
erture is inclined at a different angle from that
of the upper part. The lower margin of the
slit, itself, may be shorter than the upper mar-
gin, or far more rarely, the lower part of the
aperture may extend farther forward than the
upper part of the aperture. Of course, except
in bilaterally symmetrical shells, the Beller-
ophontacea, the slit-bearing aperture cannot
be truly symmetrical in an assymetrical shell,
but in a few pleurotomariaceans, it does ap-
proach a symmetrical condition. (Attached
mesogastropods Siliquaridae have a single gill
and a deep slit near the suture; none of the
Paleozoic pleurotomariaceans discussed are
in any other way similar to that family.)
A distinction has been made in classifica-
tion between those gastropods that have a
sinus in the outer lip and those that have a
slit, for the morphologic difference is easy to
express. In sinuate gastropods, the course of
an individual growth line, which reflects
the course of the growing apertural lip can be
traced without a break from the suture to the
base, whereas in a slit-bearing gastropod, the
course of the growth line is interrupted by the
slit, and the lines below the slit cannot be
matched with those above. So far as this fea-
ture is concerned, there is a continuous spec-
trum from those gastropods having a narrow
shallow sinus in the aperture, through those
with the sides of the sinus subparallel for a
short distance, to those in which the parallel
sides of the slit are distinct for a long distance.
Paleozoic gastropods that have a very wide
shallow reentrant in the lip, such as Sinuopea,
are not to be confused with those more nar-
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rowly sinuate forms, such as Keenia, even
though current classification associates them
closely and gives little consideration to the
relative width of the sinus in the curved outer
lip.
Information on the slit depth cannot be
obtained from the shape of growth lines. On
the upper and lower parts of the outer lip,
growth lines may sweep toward the slit in an
arc or they may be nearly straight for most
of their length. Whatever their earlier incli-
nation, they turn backward along the straight
margins of the slit and run virtually parallel
for some before joining at the curved base of
the slit. Some living pleurotomariids have a
slit half a whorl deep. In some Permian silic-
ified Euconospira, an exceedingly narrow slit
has been observed that may be even deeper.
Within limits, when compared to the width
of an individual whorl, the slit may be rela-
tively broad or quite narrow. The most sig-
nificant way to express slit depth appears to
be with reference to the circumference of the
final whorl. On the basis of limited data, a
third generalization is that a broad slit is rel-
atively short compared with the whorl cir-
cumference, and a narrow slit is deep. The
prime consideration for the organism could
have been whether this area of anal discharge
was narrow and deep or broad and shallow;
that the shallow reentrant was a deep sinus
or a shallow slit may have had no functional
significance.
Only a few pleurotomariids survive in
present-day seas, and specimens are rare; be-
cause they live in deep water in a low-energy
environment commonly the aperture, in-
cluding the slit, is unbroken. A century ago
living pleurotomariids were novel "living
fossils." Now that more is known, they are
best thought of as remnants of a more exten-
sive group. The basic morphology of the slit
is significant, but there clearly is less vari-
ability in the features today than is found in
the fossil record. To reason by comparison,
Nautilus provides much information of value
to the study of fossil cephalopods, but one
cannot translate information on its life habits
to Ordovician Orthoceras.
PSEUDOSELENIZONE AND NOTCH-KEEL
In his classic redescription of the type
species of Paleozoic Gastropoda, Knight
(1941, p. 26, 27) introduced two terms in his
 PALEOZOIC GASTROPOD REVISION 261

glossary that otherwise have been little used. class Prosobranchia, only a few forms--pleu-
The first was "pseudoselenizone": "a seleni- rotomariaceans- have paired aspidobranch
zone like band generated by a very narrow gills. Most of the living members of the order
sinus or one with a narrow tip or by some Mesogastropoda have a single gill of a dif-
other feature than the true anal slit or notch. ferent construction, a pectinobranch gill, and
Commonly its margins are not sharply commonly a siphon to bring water to the
defined as in a true slit." The second was mantle cavity. Neogastropoda have still more
"notch-keel": "a ridge, angulation, or carina specialized shell shapes and, accordingly,
generated by a shallow notch, or even a short, mantle cavities of a more modified construc-
internal channel in the outer lip where it rais- tion. The Opisthobranchia have a reduced
es the external shell features. "A notch itself shell or lack one entirely so that the gill may
was" a shallow, but narrow re-entrant in the be exposed. It is easy to interpret this pro-
outer lip. A notch may generate a notch-keel gression as a graded series from primitive to
or selenizone." advanced, as the gastropods strove, in an
I have no recollection of Dr. Knight ever evolutionary sense, to resolve the problems
discussing in detail the term "notch-keel." of sanitation of the anterior mantle cavity
My impression is that the term may have when only one gill and no slit was present.
been derived from Knight's conversations Knight (1952, p. 11-14), like earlier work-
with either E. O. Ulrich or Josiah Bridge, for ers, recognized the relationship of the living
Ulrich did use the term "notch" in his un- pleurotomariids to the Paleozoic fossils hav-
published manuscript on Ozarkian-age gas- ing a selenizone. He emphasized it, for in
tropods. Presumably some gastropods, such these fossils the presence of a slit was a means
as the Late Cambrian Sinuopea, have a of drawing inferences concerning the soft
U-shaped sinuate emargination in the outer parts, primarily the presence of paired gills.
lip, whereas others have a V-shaped emar- Knight also was interested in shell coiling and
gination comparable with the notch one might in torsion, that is, twisting of the soft parts.
cut into a tree with two strokes of an axe. In In his view, they were distinct events. He
a later paper, Knight (1952) introduced the judged that a slit was prima facie evidence
concept of the notch-keel as the site of anal of paired gills in an anterior mantle cavity
discharge and thus, in a functional sense, and therefore an indicator of torsion. Knight's
comparable with a slit; he did not elaborate systematic studies were primarily of Penn-
further. He never compared width of a slit sylvanian and Permian gastropods and in
with width of a pseudoselenizone, nor did he most of the pleurotomariaceans he studied,
ever state how a pseudoselenizone was gen- the selenizone was a prominent morphologic
erated if the area was not a site of anal dis- feature.
charges. The concept of notch-keel is not en- Knight had a strong antipathy to the group
tirely clear. Therefore, pseudoselenizone, like Amphigastropoda, the theoretical concept of
a true selenizone, has some width, whereas a Wenz (1940) and earlier writers, who regard-
notch-keel has effectively no width apart from ed both flat cap-shaped monoplacophorans
the width of the entire gastropod aperture and bilaterally symmetrical coiled bellero-
because the sides of a notch come together at phontiform shells as nontorted organisms. On
a point. Neither "notch" nor "notch-keel" the one hand, Knight recognized that the
appears in the glossary of the "Treatise" (Cox, multiple paired muscle scars of the Mono-
1960). placophora suggested lack of torsion; on the
other hand, he recognized that the slit in Bel-
THE WORK OF KNIGHT ON THE
lerophon was equally strong evidence of tor-
GASTROPOD SLIT RECONSIDERED sion. To resolve this problem, he expanded
After his 1941 book, Knight (1952) de- the Gastropoda to include two subclasses: 1,
voted years to a seminal paper on the origin the untorted Monoplacophora and Polypla-
and early evolution of Monoplacophora and cophora combined, which he collectively
Gastropoda. At the time, one of the major termed Isopleura; and 2, the torted Gastropo-
synthesizing themes concerning living Gas- da in a traditional sense, called by him An-
tropoda was involved with the mantle cavity. isopleura.
In the order Archaeogastropoda of the sub- To summarize his views on phylogeny, he

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262 ELLIS L. YOCHELSON
had the Gastropoda (Anisopleura) originate
with bilaterally symmetrical curved com-
pressed forms, such as Helcionella, which
lacked a slit but were presumed to have
undergone torsion. These forms would in turn
give rise to the bilaterally symmetrical, coiled,
slit-bearing bellerophontaceans. The next step
would lead to the asymmetrically coiled slit-
bearing pleurotomariaceans. All other sub-
sequent groups ofgastropods would have been
derived from the pleurotomariacean paired-
gill level of morphology.
The last two decades have seen consider-
able controversy concerning gastropods and
molluscan phylogeny. All views or even all
changes of opinion by an individual investi-
gator cannot be summarized easily. I main-
tain my argument of some years ago (Yoch-
elson, 1967) that the bellerophontiform shells
that have a slit are gastropods, but I likewise
still acknowledge that some coiled bilaterally
symmetrical fossils, like Cyrtonella, which
lack a slit may be forms that had not under-
gone torsion. In contrast to Knight, I still hold
my opinion that the pleurotomariaceans gave
rise to bellerophontaceans (Yochelson, 1967).
Finally, I think that the Early and Middle
Cambrian mollusk Helcionella and its allies
are not bellerophontaceans nor even gastro-
pods. Whatever the shade of opinion that
might have been expressed in the past decade,
I believe that everyone who has written on
the phylogeny of the early gastropods would
agree that Knight's sequence of gastropods
from a compressed bilaterally symmetrical to
slit-bearing and torted asymmetrical shells
can no longer be accepted as a general evo-
lutionary statement.
THE WORK OF KNIGHT ON GASTROPOD
COILING AND OPERCULA RECONSIDERED
Not only was Knight among the first to
recognize the significance of the slit for in-
terpretation of anatomy of early pleuroto-
mariaceans, he was also a pioneer in attempt-
ing to reconstruct the soft parts of fossil
gastropods from the coiling of the shell
(Knight, 1952). For bilaterally symmetrical
shells, he introduced the concept of isostro-
phic coiling, all other gastropods being asym-
metrically coiled or anisostrophic; the term
"anisostrophic" collectively covers orthos-
trophic or hyperstrophic shell shapes. When
a shell is held spire upward, aperture facing
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the observer, the typical living gastropod has
an orthostrophic coil toward the right, or dex-
tral. Living gastropods that are sinistral, coil-
ing to the left, are uncommon but fairly well
known. "The true sinistral gastropod is in all
respects a mirror image of a dextral gastro-
pod" (Knight, 1952, p. 9). A rare condition
among living gastropods is hyperstrophic
coiling: "... in which the shell is inverted
and what appears to be the spire is homol-
ogous with the base of orthostrophic forms .
... The shell resembles superficially a sinis-
tral shell but the soft parts are dextral"
(Knight, 1952, p. 9).
Little is known of fossil opercula, yet they
are important to the issue of hyperstrophy.
In living forms the operculum coils opposite
from the shell. Thus, an orthostrophic dextral
gastropod has an operculum which coils to
the left when viewed from the exterior.
Inasmuch as living marine gastropods have
an operculum, one presumes that fossil shells
also had this associated structure, but because
most opercula are not calcified, a poor record
of them exists (Yochelson, 1979a). It is quite
apart from the additional problem of asso-
ciating loose opercula with shells. To over-
generalize, opercula may fit at or just within
the aperture or may be drawn far into the
shell. The living pleurotomariids have a small
corneous operculum that is retracted beyond
the end of the slit, as would seem reasonable
if the purpose of the operculum is to close
the soft parts off from the environment.
A prominent gastropod in the Middle and
Late Ordovician is Maclurites, known from
both its shell and calcified operculum for more
than a century (Salter, 1859, p. 9). Maclurites
has been recognized as an atypical gastropod,
because the operculum coils to the left and
the shell in conventional orientation is also
sinistral. Knight interpreted this as a hyper-
strophically coiled gastropod, in part because
in living gastropods operculum and shell have
opposite directions of coiling. He went on to
elaborate upon the concept of hyperstrophy,
recognizing that in theory one could have both
dextral and sinistral hyperstrophic shells. He
then assembled 16 other genera of Paleozoic
gastropods that he assumed were hyper-
strophic rather than sinistral (Knight, 1952,
p. 36-40).
The assumption of hyperstrophy was based
on interpretation of shell shape and direction
 PALEOZOIC GASTROPOD REVISION
of coiling only, not on the coiling of shell and
operculum. From the standpoint of con-
chology, these 17 genera form a heteroge-
neous group. Most are conical to moderately
high spired. A few, namely Lecanospira, Le-
sueurilla, Macluritella, and Omphalocirrus
are flattened on one side and more or less
planorbiform in shape; in that respect, they
show some similarity to Maclurites. At the
time of his work, only the operculum of Ma-
clurites was known. I have since found an
operculum for Palliseria, a simple flat plate
with an interior muscle prong (Yochelson,
unpublished data) and Teiichispira Yochel-
son and Jones (1968) has been added to the
Macluritidae; this genus is known from both
shell and elongate operculum. None of the
three opercula are really coiled.
For a massive calcified operculum to close
the aperture, the shell should have a fairly
simple opening. The aperture of Maclurites
lacks any reentrant and has an outer lip that
is nearly vertical, when the shell is placed in
stable position with the flat surface down-
ward. Omphalocirrus also has these highly
generalized features of the aperture, but ex-
cept for flattening of one side other aspects
of the shell morphology are different; the cir-
cular multispiral calcified operculum is strik-
ingly different from the operculum of Maclu-
rites (Yochelson and Linsley, 1972). Although
Omphalocirrus was considered a macluritid
by Knight, it now is placed in the Euom-
phalacea (Yochelson, 1966; Linsley, 1978).
Knight suggested that because fossil uni-
valve mollusk shells were coiled asymmet-
rically, the soft parts had undergone torsion.
However, because certain shells, such as
Matherella, were presumably hyperstrophic
and because the earliest members of this hy-
perstrophic assemblage appeared in the fossil
record before the slit-bearing pleurotomari-
aceans, they were judged by him to be an
early offshoot of the bellerophontaceans.
Knight's proposals relative to hyperstrophic
coiling were embodied in the "Treatise"
(Knight, Batten and Yochelson, 1960) in the
classification of the Macluritacea and in com-
bining the Macluritacea and Euomphalacea
in the suborder Macluritina.
Omphalocirrus has been shown rather con-
vincingly not to be allied to Maclurites
(Yochelson, 1966; Linsley, 1978), but many
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263
of the other genera in Knight's grouping of
Macluritacea remain little studied.
Most persons are not aware that the con-
cept of Paleozoic hyperstrophic gastropods
ultimately stems from such a small amount
of data on shells and opercula. If opercula
could be found for a few of the other genera
which Knight considered hyperstrophic, some
of our current confusion might be clarified.
In particular, high-spired Matherella and its
allies are considered hyperstrophic by fiat,
rather than by fact. Further, in recent years,
the concept of hyperstrophic coiling has be-
come muddled. It can imply a major modi-
fication of the soft parts. It can also be used
in a geometric sense alone referring to the
shell. Not all hyperstrophically coiled shells,
as the term is variously used, must auto-
matically be assumed to have hyperstrophic
soft parts.
SPECULATIONS CONCERNING KNIGHT'S
CLASSIFICATION OF
PALEOZOIC GASTROPODS
Knight began his systematic work in the
early 1930's, studying Pennsylvanian gastro-
pods from a restricted stratigraphic interval
and geographic area in Missouri. He attempt-
ed to document the entire gastropod assem-
blage but was unable to complete work on
any of the slit-bearing forms because of un-
certainty regarding the use of generic con-
cepts then prevailing. This led him to deter-
mine the type species of all Paleozoic
gastropod genera which had been described
to that time and to describe the type specimen
of each in detail (Knight, 1941), published
just prior to entry of the United States into
the second world war.
Few people have ever studied or even to-
day are studying Paleozoic Gastropoda. Gas-
tropods may be the least studied group among
the abundant macrofossils in the Paleozoic.
Knight was the first to apply modern concepts
of functional morphology and evolution to
the Paleozoic gastropods. He elaborated on
the concept of the Monoplacophora and rec-
ognized their distinctiveness from the Gas-
tropoda. He was much taken with the evo-
lutionary question of torsion and coiling, and
because he was willing to speculate, attempt-
ed to fit the available data into a framework
of time of occurrences in the fossil record.
264 ELLIS L. YOCHELSON

Apart from Wenz, he was the first to attempt
a classification of Paleozoic gastropods with-
out directly referring them to extant families.
The German gastropod worker W. Wenz
began his classification of Gastropoda in the
1930's and completed it during the second
world war (Wenz, 1939-1944). The two had
met in Germany and had corresponded brief-
ly before the novel paper by Wenz (1940)
proposing Monoplacophora and combining
them with Bellerophontacea as Amphigas-
tropoda. In Knight's view, Wenz was wrong
in his concept of the Amphigastropoda.
The evolutionary scheme of Knight, based
on the slit, rationalized the interpretation of
Bellerophontacea as an extinct major group
closely allied to Pleurotomariacea. The coiled
shell marked the start of torsion within the
Mollusca, and the change from symmetrical
to asymmetrical coiling could be accounted
for in terms of evolutionary divergence.
There remained a residue of generally
poorly known Early Paleozoic shells that did
not fit this new scheme. The concept of an
angulation acting as an anal emargination in
these forms was another generalization that
could be used to interpret the early history
of the class. Given sufficient study, this con-
cept of the function of the angulation held
the promise of yielding as much interpreta-
tion as the slit. It is a human trait to want to
do better than a predecessor. A reinterpre-
tation of the Macluritacea and, indirectly, the
Euomphalacea provided yet another oppor-
tunity to improve upon the classification by
Wenz, who considered them both as family-
level taxa. Unfortunately, Knight's knowl-
edge of the V-shaped notch was based on the
work of others, and the premise of hyper-
strophic coiling was based on a single genus.
Because of the paucity of data his conclusions
are particularly subject to reinterpretation, as
I have done below and in past papers. He
would have been the first to insist that no
ones idea's in any field are sacrosanct, par-
ticularly in the light of new findings.
NEW DATA ON LESUEURILLA
AND LECANOSPIRA
Through the kindness of Dr. R. Courtes-
sole and Prof. C. Babin, I have been studying
the Lower and Middle Ordovician gastro-
pods of the Montagne Noire region of south-
ern France. The fauna is predominantly bel-
lerophontacean but contains a few other
forms. Following earlier workers, I identified
the euomphalacean Lesueurilla prima (Per-
ner) in the fauna. I also found a few specimens
of the pleurotomariacean Pararaphistoma
(Climacoraphistoma) vaginati (Koken and
Perner), a species that has a flush slitband on
the upper surface (Figure 1F). Upon com-
paring the two, I found strikingly similar
growth lines.
The specimens are external molds pre-
served in nodules, and the impressions of
growth lines are remarkably good. This sim-
ilarity in growth lines of genera supposedly

FIGURE
I-A, LesueurillainfundibuliformKoken, a small topotype in oblique dorsal view for general
orientation, x 1.5. USNM 316119. Ordovician, Lower Gray Orthoceras Limestone, at northeast end
of Neptunes Anker, 2.5 miles north of Byxelkrok, Oland, Sweden. B, Lesueurilla prima (Perner), an
oblique basal view of a latex impressionof a topotype shows curvatureof growthlines high on the
outer surface, x 3. USNM 316120. Ordovician,SarkaBeds, Osek near Rokycany,Czechoslovakia.
C, Detail of upper whorl of specimen illustrated in A, to show growth lines curving back abruptly
just before the narrow flattened peripheral keel; thickness of the shell is indicated to the upper left,
x 3. D, Detail of outerwhorlface of specimenillustratedin A, to showgrowthlines, sweepingstrongly
forwardnear upperpart of face, x 3. E, Obliqueview of externalmold illustratedin B to show the
growth lines, x 3. F, Pararaphistona (Climacoraphistoma) vagenati (Koken and Perner), a finely
preserved external mold to show the curvature of growth lines on upper and outer whorl surfaces
toward a narrow flat peripheral slit band, x 3. Universit& de Bretagne Occidentale LPB-9026. Or-
dovician, Arenigian,La Maurerie"Les Pierrils,"Herault,France. G, Lesueurillaprima (Perner),a
finely preserved external mold showing the curvature of growth lines on upper and outer whorl
surfaces, x 3. Universit6 de Bretagne Occidentale LPB-9029. Ordovician, Arenigian, Roquebrun-
L'Escougoussou, Herault, France. H, Lecanospira compacta (Salter) a finely preserved external mold
showing the curvature of growth lines on upper and outer whorl surfaces, x 2. USNM 316921. Lower
Ordovician, Nittany Dolomite, 2 miles north of Spruce Creek, Tyrone quadrangle, central Penn-
sylvania.

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 PALEOZOIC GASTROPOD REVISION 265

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266 ELLIS L. YOCHELSON
far separated systematically was so surpsising
that it needed additional confirmation. An
available topotype specimen of L. prima from
Czechoslovakia is preserved in even sharper
detail than are the French specimens (Figures
1B, E); it shows growth lines sweeping to-
wards an exceedingly narrow area that forms
a sharp keel on the periphery, seen on other
specimens which preserve the upper surface.
The growth lines on either side of this keel
have different inclinations, and I am unable
to trace the growth lines across the feature.
By definition, this keel is the site of a slitband.
I have examined a large collection of L.
infundibulum Koken, the type species of Le-
sueurilla. Although these specimens are com-
monly preserved as steinkerns, enough shell
remains on one individual to show that the
growth lines bend strongly and, just below
the upper surface, abruptly backward near the
upper keel (Figures 1A, C, D). This feature
is not a notch in the sense of Ulrich, but a
more deeply emarginated structure. I am un-
able to trace the growth lines across the keel,
so that, by definition, this area is a selenizone.
More accurately, the area is a slitband, for no
lunulae are to be seen. Interestingly enough,
Knight's 1941 illustration and description of
the type species provide exactly the same in-
formation. It is the interpretation that is dif-
ferent.
Having established in my mind this re-
markable similarity in fundamental mor-
phology of the aperture between a Middle
Ordovician low trochiform pleurotomari-
acean (Pararaphistoma) and a Middle Or-
dovician shell having a flatly coiled upper
surface (Lesueurilla), I then searched older
material. Hundreds of specimens of the mid-
dle Early Ordovician Lecanospira were as-
sembled by Ulrich, for this genus has great
stratigraphic utility. In his collections, I have
not yet found one specimen perfectly enough
preserved to see the precise nature of the
growth lines on the keel. In the best pre-
served, the trend of the growth lines on the
two whorls surface, inward- and outward-in-
clined because of the shape of this genus, is
different (Figure 1F). Had this form of growth
line been observed in a more conventionally
coiled trochiform shell, I would have inter-
preted the course of the growth lines as evi-
dence of an obscure selenizone. The overall
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similarity in curvature of growth lines is suf-
ficient to convince me that if a narrow slit
occurred in Lesueurilla, it also occurred in
Lecanospira.
The shape of the aperture, as indicated by
growth lines, is important. As a consequence
of the angle at which a shell is carried, the
various parts of the aperture may have dif-
ferent inclinations (Linsley, 1977). In a tri-
angular aperture, geometrical considerations
would not necessarily require that the inner
and outer lips have the same amount of in-
clination. However, no theoretical position
of the shell on the foot would account for the
curvature of aperture near the angulation seen
in both Lesueurilla and Lecanospira.
Another aspect of apertural shape in Le-
sueurilla and Lecanospira may be noted. It
is evident that a nonflexible calcareous plate
could not effectively seal this form of aper-
ture. However, a corneous operculum that
would be retracted some distance into the
whorl could be effective. As discussed, a few
Macluritacea and some Euomphalacea are
known to have had calcified opercula. No
calcified opercula are known among living
pleurotomariids, and no fossil opercula have
ever been demonstrated to be associated with
pleurotomariaceans. This evidence is nega-
tive and weak, but the shape of the aperture
when considered from the standpoint of
opercula does not rule out the assignment of
Lesueurilla and Lecanospira to the pleuro-
tomariaceans, as suggested by the growth
lines.
IMPLICATIONS OF INTERPRETATIONS
FOR CLASSIFICATION
The data presented produce several diverse
lines of speculation: First: If Lesueurilla and
Lecanospira have a slit or a slitlike feature,
they had two gills and in a functional sense
are allied with the pleurotomariaceans. At the
same time, the overall shell shape is not like
that of Pleurotomaria or any of its close allies.
I believe that Pleurotomariacea as used in the
"Treatise" should be broken into several su-
perfamilies to allow for major conchological
differences, the presence of a slit or slitlike
feature being a characteristic at the subor-
dinal level. Murchisoniacea would be includ-
ed within this suborder. Under this scheme,
the Bellerophontacea also could be accom-
 PALEOZOIC GASTROPOD REVISION
modated in the Suborder Pleurotomariina.
The current Suborder Bellerophontina should
be abandoned. It contains the Helcionellacea,
which I do not judge to be related to the
Bellerophontacea and not even to the class
Gastropoda (Yochelson, 1978, p. 182; 1979b,
p. 343), and the Bellerophontacea, which by
accident also include a few coiled putative
monoplacophorans that should be removed.
The concept of Bellerophontacea as Pleuro-
tomariacea sensu lato seems conchologically
reasonable, for the shape of Lecanospira is
no more different from that of Pleurotomaria
than is the shape of Bellerophon.
Second: There is a problem about the gen-
era to be included within the Suborder Ma-
cluritina. Lesueurilla may be regarded as a
"swing" genus, for in 1952, Knight placed it
within the Macluritacea, and in 1960, Knight,
Batten, and Yochelson moved it to the
Euomphalacea. Lecanospira, however, was
considered a "good" macluritacean in both
classifications. If neither genus is in the cor-
rect suborder, it would seem wise to look
critically at other genera and see whether a
new interpretation of the shape of the aper-
ture would move other taxa to the Pleuro-
tomariacea sensu lato.
Third: The Suborder Macluritina that links
together the Macluritacea and Euomphalacea
should be abandoned. The principle upon
which the two superfamilies are joined is the
interpretation that some of the Euomphala-
cea were hyperstrophically coiled; in retro-
spect, it makes no sense whatsoever to have
ever suggested both hyperstrophic and or-
thostrophic soft parts within the Euompha-
lacea. Even moving at a snail's pace, knowl-
edge of the Euomphalacea has increased
somewhat during the last two decades. It is
a reasonable presumption that many of these
animals were sedentary and that some may
have been filter feeders rather than browsers,
for the shell shape is not suited to crawling.
Animals that live with the shell lying on the
substrate require stability and a wide low shell
with a nearly flat base is well-suited to this
life mode. I have argued that if the bottom
is soft, the coiling might even grow slightly
upward to keep the aperture out of the mud
(Yochelson, 1971). With these concepts in
mind, the few euomphalaceans, such as Om-
phalocirrus, that seemingly coil in a hyper-
strophic manner are readily reinterpreted as
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267
orthostrophic. The opercula that are known
within the Omphalocirridae, the most hy-
perstrophic-appearing of the Euomphalacea,
are those of an orthostrophic coiled shell. The
vast majority of the genera within the Euom-
phalacea do not remotely resemble Maclu-
rites.
Fourth: The highly speculative issue of the
number of gills in Macluritacea and Euom-
phalacea ought to be discussed again. The
operculum of Maclurites has two attach-
ments for retractor muscles. From this paired
feature and from the assumption that in hy-
perstrophic shells the angulation is " ... the
trace of the dorsal angulation" (Knight 1952,
p. 38), paired gills were assumed for the Ma-
cluritacea. Such an assumption is plausible
but, like other such aspects of the soft-part
anatomy of fossils, essentially unprovable. It
has also been tacitly assumed that the sup-
posed descendent Euomphalacea also had
paired gills. The subgenus Euomphalus and
a few other euomphalaceans have a strong
nearly right-angle shoulder to the whorl,
which theoretically could have acted to chan-
nel water from the gills. Most euomphala-
ceans do not have such a prominent shoulder,
and some have a more rounded whorl sec-
tion. Indeed, for water circulation within the
aperture, the interior shape of the whorl, not
the exterior, is important. The shell of euom-
phalaceans that possess an angulated shoul-
der is thicker in that area so that the interior
of the aperture is rounded. There is no pro-
nounced exit channel for exhalant water.
Thus, no compelling conchological evidence
exists of the possible presence of two gills.
The Euomphalacea ought to be judged an
independent superfamily possibly derived
from the pleurotomariacean stock, in the same
way that the Trochacea are thought to have
been derived from that stock. Lesueurilla-
like forms could have lost one gill and given
rise to the Euomphalacea.
Fifth: The content of the Macluritacea de-
serves reconsideration. Although there are
many specimens of Maclurites, the Maclu-
ritidae is not a taxonomically diverse family;
when Omphalocirrus and now Lecanospira
are removed, it is much diminished. All that
remains are Palliseria, Maclurites, Teiichi-
spira and the inadequately known Macluri-
tella; calcified opercula are known from the
first three genera, but not from the fourth.
268 ELLIS L. YOCHELSON
No evidence of a calcified operculum exists
for any of the Onychochilidae, the other fam-
ily of Macluritacea currently recognized. This
family has been reconsidered by McLean
(1981) who would not interpret any of the
included genera as hyperstrophic. That may
be an extreme view, but his work deserves
careful evaluation. Certainly some of the
"high-spired" macluritaceans are better
placed elsewhere among the gastropods; per-
haps as McLean suggests, the Trochacea be-
gan earlier than we realize, and that there are
early sinistral trochaceans. To attempt to dis-
tinguish a hyperstrophic shell from a sinistral
shell on conchological features alone is far
more difficult a task than Knight thought.
Hyperstrophy may have been an important
event in the morphologic evolution of the
gastropods, but it may have been limited to
a few genera and not have had any long-range
implications.
To summarize, in part, one well-preserved
specimen of Lesueurilla has cast doubt on a
substantial part of our current classification
of Paleozoic gastropods. It raises the possi-
bility of greater conchological diversity with-
in the Pleurotomariacea and it destroys a link
of Euomphalacea with Macluritacea. If my
interpretation is correct, several genera should
be reexamined, and a new basis for classifi-
cation is needed.
Recently, Morris and Cleevely (1981) pro-
posed a major revision of the Euomphalacea.
On some of the points suggested by them I
agree, such as "It is by no means certain that
they share any character with the Macluri-
tacea apart from the coiled shell." On some
points I disagree, such as in placing the Ophi-
letidae and the Trochonematidae within the
Euomphalacea; if nothing else, the difference
in shape between the predominately flat dis-
coidal euomphalids and the lenticular ophi-
letids on one hand and steplike trochone-
matids on the other would seem to transcend
the limits of a superfamily. On some points
I am dubious, for these authors tentatively
place the Euomphalacea in the suborder
Pleurotomariina "... as they at least share
the assumed primitive character of paired
ctenidia." This assumption is based on the
presence of a sinuate angulation in many gen-
era. Whether this feature is homologous with
a distinct narrow sinus or a selenizone is a
real question. It is an act of faith--of that I
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have been as guilty as others-to base clas-
sification on presumptions concerning the soft
parts. It is far better to keep our interpreta-
tions distinct from our facts, such as size,
shape and geologic occurrence. In my view,
the Euomphalacea are not close to the Pleu-
rotomariina and they are better put in a dis-
tinct suborder or at least within the Suborder
Trochina, of which all living members have
a single gill; this would have the advantage
of using Pleurotomariina for those shells in
which the presumption of paired gills has a
bit stronger morphologic evidence to support
the presumption.
On the other hand, if all facts were known
or if we all agreed on the same interpretation
of the store of facts, classification would be
static. Not only is healthy disagreement stim-
ulating, it leads to clarification of thought and
perhaps to new concepts which bring us a step
closer to a better understanding of the diver-
sity of former living animals.
REFERENCES
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 PALEOZOIC GASTROPOD REVISION
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MANUSCRIPT
RECEIVED 21, 1981
SEPTEMBER
REVISED MANUSCRIPT RECEIVED APRIL 12, 1982