ArticlePDF Available

About the supra-generic classification of the Marginelliform Gastropods: a morphological study

Authors:
  • Independant Searcher
Biodiversity Journal, 2019, 10 (3): 221–236
About the supra-generic classification of the Marginelliform
Gastropods: a morphological study
Franck Boyer
4, impasse Pasquier, 30190 Garrigues Sainte Eulalie, France: e-mail: franck.boyer7@orange.fr
ABSTRACT
Received 26.08.2019; accepted 14.09.2019; published online 18.09.2019
The supra-generic classification of the marginelliform gastropods is reevaluated on the ground
of a critical review of the main general revision works dealing with the combined analysis of
the conchological characters, of the external anatomy and of the radular morphology, and less
systematically of the organization of the alimentary tract. The disparate nature of several
groups is demonstrated, and it provides grounds for the description of three new supra-generic
taxa: Marginellonidae fam. nov., Plesiocystiscinidae fam. nov. and Canalispirinae subfam.
nov.
INTRODUCTION
The integrative taxonomy of the marginelliform
gastropods has found probably its first hearty
demonstration in the precursory work of Coan
(1965) who proposed a relatively innovative clas-
sification confronting the conchological data with
some available elements concerning the external
anatomy of the animals and the types of the radulae.
Coan (1965) proposed to recognize three sub-fam-
ilies within a single family Marginellidae J. Flem-
ing, 1828: Marginellinae J. Flerming, 1828 and
Cystiscinae Stimpson, 1865 implicitely derived re-
spectively from Marginellidae and from Cystisci-
dae Stimpson, 1865, as well as Marginelloninae
subfam. nov., explicitely proposed for sheltering the
two monospecific genera Marginellona Martens,
1904 and Afrivoluta Tomlin 1947.
In this formative period of the integrative tax-
onomy of the marginelliforms, the definition pro-
duced for each of these three sub-families remained
eminently summary: the Marginellinae were de-
fined as being composed of the largest species”,
with a radular plate said to be “flat or curved with
many cusps, whereas the Cystiscinae were defined
as being composed of very small species with
characteristic white shellsand provided with “a
small and arched radular plate with fewer cusps”,
when the Marginelloninae were recorded as “rela-
tively large species with radular plates bearing nu-
merous cusps”.
The family Marginellidae did save his unifying
status, whereas the family Cystiscidae went down-
graded at the rank of sub-family within the Margin-
ellidae. In his conclusion, Coan (1965) detailed the
composition of these three sub-families in genera
for the fauna of the Panamic Province (“West Amer-
ican species”), and he annexed a list of all the avail-
able genera, as well as their synonymy.
The anatomical study of Ponder (1970) about
four marginelliform species from Indo-Pacific
Province and its exploitation in the frame of a
broader reflexion about the taxonomy of the
neogastropods (Ponder, 1974), followed by a work
KEY WORDS
Marginellidae; Cystiscidae; Granulinidae; taxonomic classification; morphology; cladistics.
https://doi.org/10.31396/Biodiv.Jour.2019.10.3.221.236
of operculum, polished surface of the shell, vitreous
suture), Harasewych & Kantor (1991) report the
convergent anatomic features which seem to sup-
port “the original assignation of this taxon (A’s N:
M. gigas) to the family Marginellidae (presence of
a buccal caecum, uniserial radula with broad
rachidian comblikeplate, wide aperture of the
digestive gland, wide rectal gland and palleal
oviduct with separated ingestive gland and seminal
receptacle)”, before to underline some significant
divergency traits, especially the non-specialized al-
imentary system of M. gigas, with a strong Gland
of Leiblein without terminal bulb as well as an im-
portant Valve of Leiblein, which presents features
of primitive neogastropods so far not-recorded in
the Marginellidae”. Furthermore, M. gigaswas said
to be “deprived of both a glandular duct bypassing
the Valve of Leiblein and as well as a salivary
gland, which are present in the documented Mar-
ginellidae, including in the single species of Cys-
tiscinae studied (A’s N : for this feature) hitherto”.
According to Harasewych & Kantor (1991), these
particularities suggest that the group Marginelloni-
nae may constitute “the most primitive of the Mar-
ginellidae subfamilies”, that means more primitive
than the Cystiscinae, considered itself by Coovert
(1989) as a primitive group within the Marginelli-
dae from the fact of the inclusion of a triserial radu-
late species.
Coovert & Coovert (1995) published their
supraspecific revision of the marginelliforms, split-
ting the group in two families Marginellidae and
Cystiscidae, the latter being restored for the occa-
sion. The Marginellidae received two subfamilies:
Marginelloninae Coan, 1965 and Marginellinae
Flerming, 1828, the Marginelloninae remaining
composed of their two monospecific genera Mar-
ginellona and Afrivoluta, and the Marginellinae
being distributed in three Tribes (Austroginellini, tr.
nov., Prunini tr. nov. and Marginellini J. Fleming,
1828) constitued of many genera. Four sub-families
were recognized in the Cystiscidae: Plesiocystisci-
nae subfam. nov., Cystiscinae, Persiculinae subfam.
nov. and Granulininae subfam. nov. This organiza-
tion was based on the comparison of relatively
broaden compiled data concerning the shell mor-
phology, the external anatomy of the animals (prin-
cipally the structural shape of the head) and the
morphology of the radulae, data completed for
some species and some genera by the description of
of Fretter (1976) about the Panamic species Volva-
rina taeniolata Mörch, 1860 all together ushered a
true comparative anatomy applied to the marginel-
liforms, lifted the documentary investigations to
higher level of detail and thoroughness, and pro-
vided the framework of a radical reconstruction of
the organization of the group. Ponder (1974) did
conclude about an intermediary position of the Mar-
ginellidae (sensu lato) between the Volutidae and
the Volutomitridae, while pointing out at the same
time a greater proximity with the latter, based on
the similarity of the columellar plaits and of several
shared anatomic characters (one single ingestive
gland, right diverticula of the renal organ approach-
ing or penetrating the renal-genital duct, lateral
radular teeth lacking in the Marginellidae versus
faint or lacking in the Microvolutidae, as section of
the Volutomitridae).
Directly inspired by these seminal works,
Coovert (1986), continued by Coovert & Coovert
(1990), did enter into a methodic work of documen-
tation about the marginelliforms from various as-
pects, which will play as pieces preparing their
“Revision of the Supraspecific Classification of the
Marginelliform Gastropods” published on 1995.
Among the most prominent of these preparatory
monographies: an essay on the genus Cystiscus
Stimpson, 1865 (Coovert, 1986), another on the
genus Afrivoluta (Coovert, 1987a), a compilation
about the external anatomy of the marginelliforms
(Coovert, 1987b), another compilation devoting to
the conchological characters (Coovert, 1988), a
third compilation about the general radulae
(Coovert, 1989), followed by a focus on the radulae
in the Volvarina/Prunum group (Coovert &
Coovert, 1990), displaying a good number of un-
published data.
Harasewych & Kantor (1991) did extend the
Ponder’s method, applying it to the elusive species
Marginellona gigas (Martens, 1904) from south-
eastern Asia. The authors confirm the belonging of
Marginellona (and by extension: of the Marginel-
loninae) to the family Marginellidae, however with
some consistant reserves. After to have remembered
the initial attribution of M. gigas and of Afrivoluta
pringlei Tomlin, 1947 to the family Volutidae (in
particular for the head shape and for the morphol-
ogy of the columellar plaits), and the original fea-
tures which led to their subsequent placement in the
family Marginellidae (radular morphology , lacking
FRANCK BOYER
222
About the supra-generic classification of the Marginelliform Gastropods: a morphological study
the radular odontophore and/or of the alimentary
canal.
This organization in families and sub-families
will be adopted as soon by the malaco-taxonomists
and for the most it will remain undisputed up to
2008, with the revision of the genus Serrata
Jousseaume, 1875 for New-Caledonia by Boyer
(2008), who considers the Tribes organization pro-
posed within the Marginellidae as a mainly eclectic
construction and suggests the transfer of the genera
Serrata and Hydroginella Laseron, 1957 from the
Tribe Austroginellini G.A. Coovert et H.K.
Coovert, 1995 to the Tribe Prunini G.A. Coovert et
H.K. Coovert, 1995. Some new genera have been
proposed also after the general revision of Coovert
& Coovert (1995), but without changing of the ar-
chitecture of the marginelliforms organization they
proposed, in particular: three new genera in the Ple-
siocystiscinae G.A. Coovert & H.K. Coovert, 1995
for the Caribbean Sea (Intelcystiscus Ortea et Es-
pinosa, 2001, Inbiocystiscus Ortea et Espinosa,
2001 and Ticocystiscus Espinosa et Ortea, 2002), a
genus Paolaura Smriglio et Mariottini, 2001 from
the Indian Ocean in the Granulininae, a genus De-
missa Boyer, 2015 from the Indo-Pacific Province
in the Marginellidae (implicitely associated to the
Prunini), the two genera Granulinopsis Boyer, 2017
from northeastern Atlantic and Granulinella Boyer,
2017 from New Caledonia in the Granulininae (the
latter genus with reserve).
A thesis worked by Souza (2007) about “a
cladistic analysis of the family Marginellidae based
on comparative morphology” and documenting
good number of new anatomical dissections, did
confirm the distinction between Marginellidae and
Cystiscidae”, the latter being prudently considered
as “placed in more basal position within the Muri-
coidea”. The same data have been subject to a new
study by Souza & Simone (in press), in view to give
way to stronger conclusions about the inner organ-
ization of the Marginellidae and the relations with
the most closely related groups (L. Simone, pers.
comm.).
A noticeable questioning of the Coovert &
Coovert’s suprageneric taxonomy will come on
2017 with the publication of a revision of the gran-
uliniforms by Boyer (2017), who proposes to re-
move the sub-family Granulininae from the family
Cystiscidae and to place it in a new family Gran-
ulinidae, said to hold several original features and
considered to be phyletically closer to the Margin-
ellidae and Volutidae than to the Cystiscidae. The
principal traits adopted by Boyer (2017a) as diag-
nosis of the new family are: first the external
anatomy (animal head of the Type 2 in Coovert &
Coovert) and the pattern of the alimentary canal
studied by Perez-Dionis et al. (2001) (“organ of
Leiblein made of a small bag provided with a short
duct rejoining the digestive tract after to have
passed through the oesophageal nervous ring”),
these two characters being considered as similar to
the pattern found in the Marginellidae, versus the
slit head pattern occurring in the sub-families Cys-
tiscinae and Persiculinae (the Type 2 animal being
however found also in the Plesiocystiscinae, but
constituting a plesiomorphic character, possibly
analogous and widespread among non-closely re-
lated species); secondly the uniserial rachiglossan
radula with central plate wearing “dorsal tubers or
cusps” and the sculpture patterns of the shell (gran-
ulous microsculpture and overlying macrosculp-
ture), these two features showing as perfectly
original among the marginelliform gastropods and
allowing the taxonomic separation of the granulin-
ids at the family rank towards the Marginellidae as
well as the Cystiscidae. In his revision of the Gra-
nunidae of Masirah (Oman), Boyer (2018) attrib-
utes the family in erroneous terms as “Granulinidae
Boyer, 2017”, a whereas the correct attribution is
naturally “Granulinidae Coovert et Coovert, 1995”,
at the benefit of the Article 36 of the Code of Zoo-
logical Nomenclature.
On the ground of available documents about
general morphology, we propose to complete the re-
organization of the marginelliform gastropods in
creating two new families for the respective place-
ment of the sub-families Marginelloninae and
Plesiocystiscinae, and in creating also a new sub-
family within the Cystiscidae for the placement of
the genus Canalispira Jousseaume, 1875 and pos-
sible close relatives.
RESULTS
Discussion of the proposed taxonomies
Harasewych & Kantor, 1991
Harasewych & Kantor (1991: 37–38) are focus-
ing their demonstration in a comparative table of 15
223
anatomical features documented for Marginellona
gigas, 9 other Marginellidae species and one Cys-
tiscidae species (Granula sp., as Duliculum sp. in
Ponder, 1970, both junior synonyms of Gibberula
Swainson, 1840), but without any Volutidae neither
any Volutomitridae in the outgroup, while the com-
ments from their Discussion mostly centres on the
affinities of Marginellona with these two families.
The 15 tested features are dealing with external
anatomy for three of them, with the alimentary canal
for 9 of them, and with the reproductive system for
the last three. While the attribution of Marginellona
to the Marginellidae by Martens (1904) was de-
fended on the ground of the radular morphology
(modified rachiglossan, with exclusion of the later-
als, comblike central plate with short, thin, subequal
and very numerous cusps), of the lacking of an op-
erculum, of the polished shell surface and of the vit-
reous suture, these features have not been considered
in the comparative table of Harasewych & Kantor
(1991), no more than the occurrence of a “deep
groove on the dorsal side of the Marginellona head
(Fig. 1), despite the fact that all these characters are
discussed in different places of their article.
Among the 15 features tested in their table, sev-
eral of them seem to work as poorly discriminant
in the prevailing context: for instance Harasewych
& Kantor (1991) admit that the fusion of the siphon
with the head just seems to play as a frequent envi-
ronmental adaptation to the burying move, found in
fact in each of the three Marginellidae sub-families,
and they moreover imply that such an adaptation
does not hold any phyletical value. The extent and
the position of the siphon fusioningare by the
way not detailed, what is introducing a real ambi-
guity, from the fact that all the intergrading situa-
tions are in reality occurring, especially among
some of the tested species (pers. obs.). The separa-
tive dorsal canal or gutter observed on the dorsal
side is interpreted as a “tentacles fusion”, what
looks as a questionable wording and introduces a
further ambiguity: the table reports the same feature
in Granula (as Cystiscinae in Coan, 1965), without
supporting this suggested homology in their com-
ments, while the authors report the occurrence of a
similar feature in at least three groups of Volutidae.
At last, the authors underline that the third tested
feature (the lateral lobes of the head) are totally
original to Marginellona by reference to the rest of
the Marginellidae group, but widely spread in the
Volutidae. By themselves, these three features lead
to regret that any Volutidae neither Volutomitridae
were not used in the outgroup for this integrative
test.
Other features tested by Harasewych & Kantor
(1991) seem unfitting in the context, like for in-
stance the presence or absence of a radula, while
the authors admit by themselves that the loss of
the radula is supposed to be of polyphyletic order”.
The presence or absence of aciniform salivary
glands as well as the shape of the penis do not hold
no more an evident discriminant value at the scale
of the family or of the sub-family. By the way, the
comparative table shows that the tested Marginel-
lidae are disparate for these features, and most of
the features and states tested in the table prove to
have not discriminant value in the prevailing con-
text.
Despite the fact that the authors do not notify
the matter, we note that the sole Cystiscinae
species tested here is sharing 9 of the 15 anatom-
ical characters possessed by Marginellona gigas,
while only two of the other tested Marginellidae
present a better proximity (10/15 for Haloginella
mustelina and 11/15 for Mesoginella pygmaea),
one species of Marginellidae scoring about the
FRANCK BOYER
224
Figure 1. Cephalic morphology of Marginellona gigas (fol-
lowing Harasewych & Kantor, 1995 fig. 15: s = siphon, t =
tentacle, c = = channel along mid-line of head, lp = lateral
lappets.
same order than the Cystiscinae species, 7/13 for
Prunum martini (Petit de la Saussaye, 1853, and
the 6 further Marginellidae scoring lower or very
lower, from 0/5 for Hyalina pallida (Linnaeus,
1758) to 5/12 for Prunum marginata (Born, 1778).
By themselves, these results lead to question about
the pertinence and the efficiency of the compara-
tive method applied, like about the choice of the
characters and of their modes, if not about the aim
really pursued.
We observe moreover the introduction of notice-
able biases, for instance concerning the apparent
absence” of such or such character, which might
be in fact displaced, nanified or even badly
searched: that is concretely the case for the sup-
posed absence of a radula in Hyalina pallida, which
has been finally proved to be a radulate species, like
apparently all its genus (Boyer, 2015a), as well as
for the genus Dentimargo Cossmann, 1899 reported
as being non-radulate in the whole, whereas at least
two Dentimargo species have been proved to be
radulate (Boyer, 2005, 2017b). The cladistic prac-
tice is revealing herein some of its limits: the relia-
bility of the observations worked, the discriminant
value attributed to the characters, the ambiguity of
the modes adopted for the definition of the charac-
ters states, and the real comparability of individual
results.
Coovert & Coovert, 1995
Coovert & Coovert (1995) provide their analysis
from a more important number of characters, deal-
ing principally with the shell morphology, the ex-
ternal mophology of the animal (mostly the pattern
of the head), the radular type, the type of odon-
tophore and the anatomy of the alimentary canal.
The study of the characters is more deepened than
in Harasewych & Kantor (1991) and it is also ap-
plied to a larger number of species. However, only
the anatomical features are computed in a compar-
ative table (Coovert & Coovert, 1995: 54, Table 3),
excluding the shell morphology features and often
using excessively simplifying discriminatory
modes, if not teleological ones. The integrative
table proposed by Coovert & Coovert (1995) only
crosses in fact 14 characters, to be related to the 15
characters crossed by Harasewych & Kantor (1991)
in their own compative table.
The analysis of Coovert & Coovert (1995) about
the shell morphology does value some characters
felt as “representative” of special evolutionary lin-
eages, while shelving the variability factors which
might trouble the demonstration, and avoiding,
minimizing or relativising the interest either the im-
portance of further conchological features. For in-
stance the authors give a major place to the “partial
About the supra-generic classification of the Marginelliform Gastropods: a morphological study 225
Figure 2. Internal architecture of the shells, following Coovert & Coovert (1995, figs. 1–4).
resorption of the internal whorls of the shell” (Fig.
2), what constitutes from their point of view a
shared and unifying characters among the Cystisci-
dae. Boyer (2017a) demonstrated that this feature
is noticeably variable within the granuliniforms,
sometimes poorly marked or dubious in this group,
and may result from a simple convergence linked
to similar architectural constraints. In support to
their demonstration, Coovert & Coovert (1995: 49,
figs. 1–4) present by the way a figure of Granulina
hadria where the supposed partial resorption of
the internal whorls” is not at all evidenced (Fig. 2:
fig. 4). Coovert & Coovert (1995) make also an
eclectic use of further conchological characters:
they insist for instance on “the absence of a external
labial varixin the Cystiscidae, with the noticeable
exceptions of the Granulininae where it is well-pre-
sent and much marked, and of the genus Persicula
where its presence should be irregular, whereas the
external varix can be also factually represented in
some Gibberula species, what is suggesting in the
whole that this feature holds an eclectic status in the
Cystyiscidae better than a dominant and character-
istic status. The occurrence of labial teeth said to
characterize some marginelliform genera is in fact
most often facultative at the scale of some genera
or at the level of several species groups, if not spo-
radic within such or such species. The shell macro-
sculptures are spotted as constituting a frequent
or constant feature at the scale of several genera
of Marginellidae (Austroginella Laseron, 1957,
Glabella Swainson, 1840, etc.) but any diagnostic
value is given to the micro and macro-sculptures
found in the Granulininae, whereas the other groups
attributed to the Cystiscidae are totally deprived of
such features.
The study of the morphological types of the an-
imal head in Coovert & Coovert (1995) are subject
to similar biases. For instance, the “slit or bilobate
head (Fig. 3) are recognized as constituting a
shared character in the Cystiscidae groups consid-
ered as the most evolvedones (Cystiscinae and
Persiculinae), but the similarity presented by the
cephalic gutter in the Marginelloninae is not in-
quired in phyletic terms, despite the fact that
Coovert & Coovert (1995) did admit henceforth
that this gutter is also occurring in Afrivoluta
pringlei Tomlin, 1957 as it is in Marginellona
gigas, what Coovert (1987a) did not remark previ-
ously.
The comparative analysis of the radular patterns
(Fig. 4) is clearly flawed by a poorly argumented
grouping trend, if not tarnished by implausibility.
Coovert & Coovert (1995) consider for instance
that the “comblike radula with many cusps” of the
Marginelloninae constitutes a plesiomorphic stage
of the “comblike radula with fewer cusps” found in
several Marginellinae groups, and from this state-
ment they infer that the Marginellinae would be de-
rivated from the Marginelloninae. This supposed
filiation is just resting in fact on the intuition and
on an apparent “common senseagainst whom the
taxonomical research is precisely supposed to de-
fend itself: besides the fact that this character might
be only analogous, it may more probably prove to
come from the parallel evolution of very anciently
separated modified rachiglossan lineages, or even
it may play in the contrary (and more plausible) di-
rection, with the numerous-cusps radula derivating
from fewer-cusps radula. As result, the occurrence
of this comblike radula with many cusps” does not
allow neither justify by itself to unify the Margin-
ellinae and the Marginelloninae within a common
family, and no more to speculate about the close-
ness of their kinship. In the same vein, the similarity
alleged between the arched radula of the Granulin-
inae and that of the Cystiscinae and Persiculinae
Coovert & Coovert (1995) seems to be quite unsub-
stantial: besides the fact that the analogy explana-
tion remains once more possible, it must be
observed that the radula of the Granulininae pres-
ents a deep originality in the details, especially con-
cerning the dorsal tubers or cuspsof the plates
(Boyer, 2017a), any equivalent being known among
the other groups of the marginelliform gastropods.
At last, the Plesiocystiscinae are considered as
a “primitivegroup of cystiscids which would have
conserved a triserial radula, while the Cystiscinae
brother-groups would loose the laterals, but would
save the central tooth in heritage. In fact the shape
of the central tooth in Plesiocystiscus Coovert et
Coovert, 1995 differs noticeably from the uniseral
plates found in Cystiscus, Gibberula and Persicula,
from a sculpture with uneven reliefs not necessarily
homologous with the Cystiscinae pattern, but fi-
nally more comparable to the pattern found in the
Granulininae, where the sculpture of “dorsal tubers
or cuspsis however more marked and well-differ-
ent from an architectural point of view (Boyer,
2017a).
FRANCK BOYER
226
About the supra-generic classification of the Marginelliform Gastropods: a morphological study 227
Figure 3. Types of the external anatomy (head), following Coovert & Coovert (1995, figs. 5–12).
The analyse of the odontophore allows Coovert
& Coovert (1995) to shed a new light from original
observations, this character being neglected until
then in the study of the marginelliforms. The au-
thors recognize two principal states in the odon-
tophore of the marginelliforms: the occurrence of
two separated plates versus the occurrence of two
plates fusioned at the tip or fusioned in two points,
the first state seeming to be general in the Cystisci-
nae and the second state seeming to be general in
the Marginellinae, the absence of an odontophore
being naturally acted in the non-radulate species or
supposed to be. The data brought to light as well as
the demonstration are convincing by themselves,
but the number of species studied for this character
remain extremely narrow: in fact Coovert &
Coovert (1995) did study directly the odontophore
of only two species in the Cystiscinae (both attrib-
utable to Persiculinae) and of five species in Mar-
ginellinae (all attributable to the Volvarina/Prunum
complex). For the rest, Coovert & Coovert (1995)
are dealing about faintly supported undirect sources
for a very limited number of species (Ponder, 1970;
Fretter, 1976; Ponder & Taylor, 1992). In the whole,
FRANCK BOYER
228
Figure 4. Types of the external anatomy (head), following Coovert & Coovert (1995, figs. 5–12).
Coovert & Coovert (1995) can refer to pieces of in-
formation (from direct observation and full-drawing
down to short allusion) just for few species belong-
ing to three genera or super-genera: Persicula/Gib-
berula, Volvarina/Prunum and Austroginella, out
of various other Marginellidae genera holding
comblike radulae (Marginellona/Afrivoluta,
Cryptospira Hinds, 1844, Serrata, etc.) or strongly
modified radulae (Hydroginella, Demissa, etc.), and
also out of the majority of the genera placed by
them in the Cystiscidae (Cystiscus/Crithe, Gran-
ulina, Plesiocystiscus). From this fact, the general-
ization supported by Coovert & Coovert (1995) at
the scale of their two proposed families Marginell-
idae and Cystiscidae looks as poorly argumented
and quite hazardous.
The synthetic analysis of the anatomical charac-
ters looks as unbalanced from several points of
view, particularly about the panel of tested species,
constituted of three Cystiscidae (in fact only Persi-
culinae and not any Cystiscinae, Plesiocystiscinae
or Granulininae) and of twenty-one Marginellidae
(including Marginellona gigas, but out of numerous
atypical” genera such as Closia Gray, 1857, Cryp-
tospira, Demissa, Hydroginella or Serrata). This
lesser documentation of the Cystiscidae (sensu lato)
results probably from anthropic biases: the lesser
sampling of small-sized groups and their more dif-
ficult observation in vivo like on dissection duty
both lead to a lesser recognizing of the taxonomic
diversity and of morphologic disparity at work.
Even if understandable by itself, the diversity bias
introduced in the comparative Table is however im-
portant and it seems to be evident that the relative
homogeneity suggested in the Cystiscidae would be
certainly much lower (like prevailing in the 21
tested Marginellidae) if the groups Cystiscinae, Ple-
siocystiscinae and Granulininae would be inte-
grated in the comparison.
The choices made about the testable characters
and their modes calibration introduce also major bi-
ases, inducing for instance more important synapo-
morphies between Marginellona and the other
Marginellidae than with the panel of tested Cystis-
cidae (in fact only Persiculinae), this from a deeply
artificial manner.
As representative examples:
Character 1 (Buccal pouch) is splitted in the two
modes “absentversus present”, but the absence
if buccal pouch does not present necessarily an ho-
mologous feature: we observe incidentally that the
buccal pouch is reported as absentin the three
radulate Persiculinae as well as in the seven non-
radulate (or supposed to be) Marginellidae, what is
giving account of two deeply different states and
amount s to proceed to an eclectic classification.
Character 2 (Neogastropods radular sac) is also
splitted in the absent” versus “present” modes, but
it introduces a double ambiguity: from one hand we
do not understand if absent means absence of
any radular sac” or if it means “radular sac differ-
ent from the Neogastropods sac pattern”, what
should refer to very constrasted solutions depending
of the kind of radular sac implied; on the other
hand, the Cystiscidae as well as the Marginellidae
being Neogastropods, so how to define any
Neogastropods radular sacwhich would be oc-
curring in some marginelliform groups (or Neogas-
tropods ones) but not in other ones ?
Character 3 (General radular type) is splitted in
the three modes cystiscid”, “marginellid” or ab-
sent”: first such discrimination is perfectly tauto-
logic (Cystiscidae having cystiscid radulaeand
Marginellids having marginellids radulae”); sec-
ondly the “absence of the radula” does not consti-
tute a feature from the same biologic and taxonomic
nature, as representing a secondary or regressive
stage, that means a subcategory playing equally
within both families and not a valid category by its
own; at last, the absence of a radula can come from
various processes and to unit this state as a distinct
category or mode is deprived of any signification.
Same kind of remarks could be tackled for Charac-
ter 4 (Specific radular type).
Character 6 (Siphon) is splitted in the two
modes “attached left of head” versus “fused to
head”. Only Marginellona gigas and Prunum mar-
ginatum (Born, 1778) would share the second
mode (siphon fused to head) while all the other
Marginellidae as well as the three tested Persiculi-
nae would share the first mode (siphon attached left
of head). This definition is ambiguous because
fused” is by itself a kind of attachment”, and
overall because very numerous species are observed
at the live state as well as under dissection (pers.
obs.) as displaying various attachment intergrades.
Harasewych & Kantor (1991) did observe abroad
that the (partial) fusion of the siphon is essentially
About the supra-generic classification of the Marginelliform Gastropods: a morphological study 229
correlated to burrowing habits, and that such a
plastic” feature is poorly or not significant in the
phyletic field.
Character 9 (Gland of Leiblein) is splitted in
three modes: 1) “small, narrow gland, no terminal
bulb”; 2) “large, sacculate gland, no terminal
bulb”; 3) “with terminal bulb and long convoluted
duct”. The two first modes are questioning, by the
fact that their differences seems to be very subtile
and possibly not really discriminating. In fact the
mode 1 concerns exclusively the three Persiculinae
and the mode 2 concerns only Marginellona gigas,
while mode 3 is uniting all the other tested Margin-
ellidae. Such a subtile distingo operated between
the modes 1 and 2 (better than to consider that they
are variant stages in the same kind of gland) equal
to create an artificial caesura for the sake of the
cause: the Marginelloninae must have nothing to
deal with the Cystiscidae and much to do with the
Marginellidae.
We observe moreover that the absence of the
Character “external anatomy/shape of the head” in
the Comparative Table of the anatomic characters
is not justified by Coovert & Coovert (1995), and
that it introduces another important bias, together
because it masks a deep heterogeneity within the
Cystiscidae (Type 2 animal sensu Coovert, 1987b
in the Plesiocystiscinae and the Granulininae, Type
3 in the Cystiscinae, and Type 4 in the Persiculinae),
but also because it is affecting the comparative
analysis for Marginellona gigas, whose cephalic
gutter suggests possible affinities with the slit head
occurring in the Cystiscinae and the Persiculinae,
and does not have homologues in the other Margin-
ellidae which all have a Type 2 head.
If we put aside the first 4 Characters of the syn-
thetic Table for the reasons presented hereabove and
if we take in account the remark expressed about
the Character 9, the Cystiscinae (= Persiculinae)
just hold one original character among the 10 other
integrated characters : the possession of an odon-
tophore constituted by two separated plates (char-
acter non-tested in Marginellona). For the 9 other
considered characters (from 6 to 14), at least one of
the modes retained by Coovert & Coovert (1995) is
shared between the Cystiscidae (= Persiculinae) and
the Marginellidae. More generally, we can consider
that the Comparative Table of Coovert & Coovert
(1995) for the anatomical characters just demon-
strate the separation between the Cystiscidae and
the Marginellidae in a partial and not robust man-
ner. By contrast, this Comparative Table should tell
us about the position of Marginellona gigas, here
incorporated in the Marginellidae but sharing an ex-
clusive feature with two of the three Persiculinae
(presence of the Valve of Leiblein) and apparently
another exclusive feature with the three Persiculi-
nae (Gland of Leiblein without a terminal bulb). If
we consider the whole documented characters, the
three Persiculinae do share no less features with
Marginellona gigas than a good number of the
tested Marginellidae species (in fact than the ma-
jority of them). If we agree with the equivalence of
the modes 0 and 1 in the Character 9 (= Gland of
Leiblein without terminal bulb) and if we restrict to
the Coovert & Coovert’s accounting, the Persiculi-
nae only share 4 to 5 of the 13 features attributed to
Marginellona gigas, but 11 of the 20 other Margin-
ellidae are in the same case.
Souza, 2007
Souza (2007) is focusing is thesis work on the
Marginellidae in testing 22 species within an inte-
grative Table of 93 characters, in the intention to re-
fine the previous cladistic works, especially the
supra-specific revision of Coovert & Coovert
(1995). Beside 18 species of Marginellidae (includ-
ing Marginellona gigas newly dissected on two
specimens), Souza (2007) propose an interesting
outgroup of 4 species, including a Volutidae,
Adelomelon brasilianus (Lamarck, 1811), and a
Persiculinae, “Persiculasagittata (Hinds, 1844).
Among the 93 tested characters, 12 conchological
characters are associated to 81 anatomical charac-
ters. Beside other valuable results, we note the study
of the odontophore performed in Marginellona
FRANCK BOYER
230
Figure 5. Synthetic patterns of odontophores (in cut),
following Souza (2007, figs. 60–62).
gigas: this odontophore proves to be composed of
two separated plates, like it was observed in the Per-
siculinae, but in a special condition of attachment
of the horizontal muscle, named as “model m6 mod-
ified” (Fig. 5: fig. 61) and differing from the usual
model m6 found generally in the Caenogas-
tropoda”. This special odontophore model might be
considered as transitionaltowards the fusioned
model found in the other Marginellidae. Souza
(2007) reports that the odontophore found in Mar-
ginellona gigas is very similarto the one found
in the Volutidae Adelomelon brasilianus.
We note also the observation made by Souza
(2007) about the similarity presented by the “comb-
like radula with numerous cuspsfound in Margin-
ellona (83-85 cusps) and the “comblike radula with
lesser cuspsfound in other Marginellidae groups
(Volvarina/Prunum, etc). Souza (2007) gives the
average number of 45 cusps for the tested Margin-
ellidae species, but in reality the average number is
much lower in the documented Prunini species
(Coovert & Coovert, 1990; pers. obs.). Despite this
point, the topographic test applied by Souza (2007)
leads to the conclusion that the comblike radula
features” came out independently in the Marginel-
loninae and in the Prunini, and does not seem to
play as a derived character.
In conclusion, Souza (2007) confirms however
the Marginellidae (including Marginellona gigas)
as constituting a monophyletic group supported by
17 synapomorphies, the Cystiscidae being consid-
ered as a more primitive group (more “basal
cladistic position) in the group Muricoidea.
Despite the fact that this kind of result is much
conditioned by the methodologic and sampling
choices initially operated, we can consider that 17
synapomorphies among a total of 93 tested charac-
ters constitute a rather modest and poorly conclu-
sive score. Souza (2007) does not conclude nothing
specifically bout the placement of Marginellona
gigas, despite the heavy average divergence of its
characters from the characters tested in the other
Marginellidae. The synthetic Table of Souza (2007)
let us see Marginellona gigas as equidistant be-
tween Adelomelon brasilianus, Persiculasagit-
tata and the group constituted by the 17 other
Marginellidae species.
Boyer, 2017a
In his revision of the granuliniforms, Boyer
(2017a) does not bring new documentation, he is
testing and commenting just a limited number of
characters, he does not present any synthetic Com-
parative Table and he is just playing in fact a de-
ductive approach on the ground of some data
previously displayed in the literature. His propo-
sition about the autonomization of the granulini-
forms at the family level is principally resting on
three general antinomies with the other Cystisci-
dae groups (internal shell whorls poorly or not re-
sorbed; presence of micro and macro shell
sculptures; organ of Leiblein with duct bypassing
the oesophagic ring), one special antinomy with
the Plesiocystiscinae (uniserial radula), two spe-
cial antinomies with the Cystiscinae and the Per-
siculinae (non-slitted Type 2 head; embossed
dorsal tubers or cusps on the radular plate) and two
antinomies with the Marginellidae considered in
their whole (shell micro-sculptures; narrow and
arched radula with embossed tubers on the radular
plate). For lack of available documentation about
the odontophore at that time, this feature was not
tested.
Proposition of the creation of two new fami-
lies and of one new subfamily
We propose to reevaluate the position of the
Marginelloninae and of the Plesiocystiscinae in the
way initiated by Boyer (2017a), identifying the
original features recognizable for these two groups
and their taxonomic compatibility with other groups
of the marginelliform gastropods. The eclectic com-
position observed in the Cystiscidae leads to reeval-
uate in particular the position of the genus
Canalispira among them.
MARGINELLONIDAE fam. nov.
DESCRIPTION. Shell morphology: the type of the
columellar pleats and the consistency of the surface
lacquer in the Marginelloninae look as closer to
those found in some Volutidae and Volutomitridae
(Harasewych & Kantor, 1991) better than to the
models found in the Marginellidae or in the Cystis-
cidae, in which the porcellanous shell seems to have
a different composition (pers. obs.).
External anatomy of the animal (shape of the
head): the dorsal cephalic gutter has no equivalent
in the Marginellidae, and it is not necessarily ho-
About the supra-generic classification of the Marginelliform Gastropods: a morphological study 231
mologous to the full-separation of the head in two
distincts lobes as found in the Cystiscinae and Per-
siculinae (Harasewych & Kantor, 1991; Souza,
2007). The lateral dewlaps are comparable to simi-
lar structures found in several groups of Volutidae;
this character is clearly not homologous with the
rather spread position of the posterior head ob-
served in some Persiculinae or in the Olividae (pers.
obs.).
Radula: the uniserial radula with wide central
cusps bearing very numerous cusps, looks as simi-
lar to the basic model represented in Prunini and
related groups (the genus Serrata, some Hy-
droginella, etc), but it does not seem to be a shared
character or even to play as an ancestral model
for them. The primitive” state alleged for this
radula bearing very numerous cusps does only rest
itself on the prejudice attributing a great seniority
to the Marginelloninae from the fact of their very
limited diversity in the Recent together with their
very patchy distribution are both connoting a relict-
ual status, and on some anatomical features some-
times considered as archaic, however not factually
in a different way that various archaic features re-
tained in all the organisms, beside recently derived
ones.
Odontophore: constituted of two separated
plates, like in the pattern documented for the
Persiculinae and antithetical with the pattern
of “fusioned plates” documented for the Volva-
rina/Prunum group (Souza, 2007). The inter-plates
muscular attachment shows to be of a particular
type, possibly “transitional” between the “free
state” and the fusioned state”, and the homology
with the pattern found in the Persiculinae is not
proved. For this character, Marginellona presents
apparently a special affinity with some Volutide
(Souza, 2007).
Alimentary canal: the presence of a non-
through Valve of Leiblein conciliates Marginellona
with the two Persicula proved to hold the same
pattern, and it differentiates it from all the other
Marginellidae, as well as from the sole Gibberula
documented in this field (Harasewych & Kantor,
1991; Coovert & Coovert, 1995). If this character
contributes to distinguish Marginellona from the
other Marginellidae, it does not allow to confer a
particular proximity with the Cystiscidae or with
part of them, the ontogeny, the anatomy and the
conservation of this organ presenting very hetero-
geneous situations within the neogastropods (Kan-
tor & Fedosov, 2009). The absence of accessory
salivary gland in Marginellona constitutes a less
ambivalent datum, in so far as, if this gland is pres-
ent in the three documented Persiculinae, it seems
to be present in only the half-part of the Marginel-
lidae species documented in this field by Coovert
& Coovert (remarkably in all the tested Aus-
troginelini).
REMARKS. Taxonomic placement: in the whole,
the Marginelloninae show to be in higher proximity
with the Volutidae and the Volutomitridae, specially
concerning the shell morphology, the shape of the
head and the odontophore pattern, even if for these
two latter characters more or less important affini-
ties are suggested with the Cystiscinae and the Per-
siculinae. The shape of their radula suggests that the
Marginelloninae might have a more or less close re-
lationship with the Prunini and related groups, but
it does not state necessarily a direct filiation : the
type of “comblike radula” might be inherited from
a more or less undirect common ancestor, or even
to play as a alternative evolutionary occurrence in
parallel lineages of uniseriate rachiglossan gas-
tropods. The presence of a Valve of Leiblein differ-
entiates clearly the Marginelloninae from the other
Marginellidae, but if it is shared apparently with all
the Persiculinae, these two characters look as too
much heterogeneous for a valuable statement about
a possible close relationship. In the whole, the Mar-
ginelloninae do not suggest more affinities with the
Marginellidae than with the Cystiscidae, and they
seem to range in fact rather far from them. In any-
way, the belonging of the Marginelloninae to the
family Marginellidae looks as poorly convincing,
and to consider them as possible subfamily of the
Volutidae or of the Volutomitridae would play itself
as a poorly supported taxonomical solution in the
present state. In the precise case, the ambivalence
of the established affinities leads better to state in
favour of an original combination of characters de-
serving the attribution of an autonomous status to
Marginelloninae group, better than in favour of an
artificial attachment which plays overall as a nega-
tion of the morphological and anatomical disparities
at work. In the state of the documentation, we con-
sider the group of the Marginelloninae as presenting
the conditions for a tentative attribution to a new
family Marginellonidae within the Superfamily
Muricoidea Rafinesque, 1815.
FRANCK BOYER
232
PLESIOCYSTISCINIDAE fam. nov.
DESCRIPTION. Shell morphology: the resorption
of the internal whorls, the absence of external labial
varix and the absence of the siphonal notch ob-
served in the Plesiocystiscinae are shared with the
Cystiscinae, but these two latter features are not
shared with the Persiculinae, which holds a more or
less marked siphonal notch, and a more or less
thickened external varix in several groups, and in
the whole flatter and less lamelliform columellar
plaits. On the other side, the total absence of exter-
nal labial varix in some groups of Persiculinae
(Gibberula philippii group, etc.) does not seem to
be homologous with the absence of external varix
in the Plesiocystiscinae and in the Cystiscinae (pers.
obs.).
External anatomy (shape of the head): the Type
2 head (Fig. 6: non-bifurcated headwith extended
tentacles and siphon) differs deeply from the Type
3 (bilobed headwithout tentacles neither visible
siphon) and from the Type 4 (bilobed head with
more or less extended tentacles ans siphon) respec-
tively found in the Cystiscinae and in the Persiculi-
nae, and it is not necessarily homologous with the
Type 2 found in the Granulinidae and the Margin-
ellidae, this pattern of V-shaped head with pointed
tentacles and siphon being widely spread in the
neogastropods, including in several families placed
in the group Muricoidea.
Radula: the triserial radula of the Plesiocys-
tiscinae has been interpreted as being anances-
tral state within the Cystiscidae” (Coovert &
Coovert, 1995), but nothing is materially testifying
such a statement, even if the loss of the laterals
does not constitute by itself a more complex or ex-
traordinary process than the full loss of the radula
in some Marginellidae lineages. The shape of the
central plate in the Plesiocystiscinae seems to be
morphologically less close to the uniserial plate of
the Cystiscinae and Persiculinae than to the plate
of the Granulinidae (nowadays considered as an
autonomous family presenting more affinities with
the Marginellidae and the Volutidae than with the
Cystiscidae), any evident conclusion not seeming
to play, due to the important plasticity of this organ
in most of the considered groups (the disparity of
the radular types at work in the Marginellidae
plays by itself as a very demonstrative experience
field).
Odontophore: unknown.
Alimentary canal: unknown.
REMARKS. Taxonomic placement: despite the
lacking of data about their internal anatomy, the Ple-
siocystiscinae present by themselves an important
originality compared to the other Cystiscidae groups
among which they were placed until now: their shell
morphology presents similarities with that of the
Cystiscinae, but the radula of the Plesiocystiscinae
cannot be stated as constituting the evolutionary
stage immediately preceding the uniserial radula
found in the Cystiscinae (or the other Cystiscidae as
well), and overall the external anatomy of the Ple-
siocystiscinae is sufficiently divergent from that of
the Cystiscinae for to invalidate their grouping
within the same family. We consider that the Persi-
culinae are by far more divergent from the Plesio-
cystiscinae. Without totally reject of a possible
ancestral status” of the Plesiocystiscinae towards
the Cystiscinae, or even towards the Persiculinae in
a more remote perspective, the important evolution-
ary distance suggested towards these two groups
through their external anatomy justifies by itself the
integration of the Plesiocystiscinae within an new
autonomous family Plesiocystiscidae, itself placed
in the Superfamily Muricoidea.
The distribution of the Plesiocystiscinae is
known to range along the Panamic Province and on
both sides of the tropical Atlantic Ocean (being ap-
parently much rarer on the eastern side), but the
group seems to be much more elusive in the Indo-
Pacific Province. Two species from New Caledonia
have been however attributed to Plesiocystiscus by
Boyer (2003), on the ground of the characters of the
shell associated to a Type 2 animal (Fig. 7), but
without a research of the radula. As most of the nu-
merous Cystiscus species ranging in the Indo-Pa-
cific waters were only studied on the ground of their
shell, we can expect that a good number of these
so-said “Cystiscus species” will prove to belong to
Plesiocystiscus or to brother-genera.
CANALISPIRINAE subfam. nov.
In the course of this study, the Persiculinae were
reported as constituting a specially heterogeneous
group, by reference in particular to the various tran-
sitional forms occurring between Persicula and
Gibberula, but also within the Gibberula them-
selves (Gibberula philippii group, etc.). Among the
About the supra-generic classification of the Marginelliform Gastropods: a morphological study 233
Persiculinae, the genus Canalispira presents a set
of specially composite characters, combinating a
sharp and slender bilobed head, without appendices
and no visible siphon (like occurring in the Cystisci-
nae), with a radula similar to that found in the Per-
siculinae (Type 3 sensu Coovert, 1987b) and a shell
rather similar to that of this group, even for the col-
umellar plaits, but however without siphonal notch
(Coovert & Coovert, 1995; McCleery & Wakefield,
2007; pers. obs.).
In most of the Indo-West Pacific species, the
Canalispira species present a more or less deeply
canaliculated spire and undecored white shells,
whereas in Caribbean similar species with the same
general shell shape, same kind of external anatomy
and same kind of radula are shown to be not
canaliculated and to wear zic-zac shell decoration
(McCleery & Wakefield, 2007). The occurrence of
several brother-genera may occur, but at least an in-
tergrading situation was observed in the Maldives,
with a full-white shelled species presenting a non-
canaliculated spire (Boyer, 2018b).
The original combination of features in the ex-
ternal anatomy and in the shell morpology of
Canalispira (Fig. 8: lectotype of the type-species
Canalispira olivellaeformis Jousseaume, 1875)
leads to propose to integrate this genus in a new au-
tonomous subfamily Canalispirinae within the Cys-
tiscidae, beside the subfamilies Cystiscinae and Per-
siculinae.
The species described from São Tomé as “Gib-
berulacucullata by Gofas & Fernandes (1998)
presents together a sharp and slender bilobed head
deprived of tentacles, perfectly similar to the Canal-
ispira pattern, but also a short but visible siphon and
a shell belonging to the classic Gibberula-pattern,
with an evident siphonal notch and prominent labial
teeth. The possible replacement of this transitional-
looking species will deserve to be studied by itself.
CONCLUSIONS
The new supra-generic organization proposed
for the marginelliform gastropods consists to rec-
ognize five families in a group which must be con-
sidered in the present state as polyphyletic within
Muricoidea, and to proceed to the creation of a new
subfamily in the Cystiscidae, conserving for now
the rest of the organization proposed previously by
Coovert & Coovert concerning the composition in
subfamilies and tribes (with the replacement of Ser-
rata and Hydroginella in the Prunini, as reported
before).
FRANCK BOYER
234
Figure 6. Plesiocystiscus jewetti (Carpenter, 1857), Cave Landing, Santa Barbara, Calif., by J. Goddard
Figure 7. “Plesiocystiscusbavayi Boyer, 2003, Touho, New Caledonia, in Boyer (2003, fig. 78).
Familia MARGINELLIDAE Fleming, 1828
Subfamilia MARGINELLINAE Fleming, 1828
Tribe AUSTROGINELLINICoovert et Coovert, 1995
Tribe PRUNINI Coovert et Coovert, 1995
Tribe MARGINELLINI Fleming, 1828
Familia GRANULINIDAE Coovert et Coovert,
1995
Subfamilia GRANULININAECoovert et Coovert,
1995
Familia MARGINELLONIDAE fam. nov.
Subfamilia MARGINELLONINAE Coan, 1965
Family CYSTISCIDAE Stimpson, 1865
Subfamilia CYSTISCINAE Stimpson, 1865
Subfamilia PERSICULINAE Coovert et Coovert, 1995
Subfamilia CANALISPIRINAE subfam. nov.
Familia PLESIOCYSTISCIDAE fam. nov.
Subfamilia PLESIOCYSTISCINAE Coovert et
Coovert, 1995
The revision worked presently by Souza & Si-
mone (in press) will surely bring new useful mor-
phological and taxonomical elements about the
organization of the Marginellidae. A supra-specific
revision of the marginelliforms based on molecu-
lar data is also at work by a research team
managed by A. Fedosov in the Muséum national
d’Histoire naturelle (Paris, France): their demon-
strations and conclusions will be also highly useful
to address.
ACKNOWLEDGEMENTS
I wish to acknowledge here Luiz Ricardo Simone
(Museu de Zoologia, Universidade de São Paulo)
and Alexander Fedosov (Severtzov Institute, Russ-
ian Academy of Sciences, Moscow) which both con-
tributed to the clarification of my aims and to the
rationalization of my method in this work. I must
thank also Jeffrey Goddard (Marine Science Insti-
tute, University of California, Santa Barbara) for the
photos of live specimens of Plesiocystiscus jewetti.
As often, I am indebted to Alain Robin (Le Mesnil-
Saint-Denis, France) for his help in the mounting of
the iconography, and to Walter Renda (Amantea,
Italy) for his indefectible support. The photogra-
phies of the types of Canalispira olivellaeformis
were performed by Manuel Caballer in the frame of
the E-Recolnat program (ANR-11-INBS-0004).
REFERENCES
Boyer F., 2003. The Cystiscidae (Caenogastropoda) from
upper reef formations of New Caledonia. Iberus, 21:
241–272.
Boyer F., 2005. The discovery of a radula in a Denti-
margo species and its taxonomic implications. Iberus,
23: 47–52.
Boyer F., 2008. The genus Serrata Jousseaume, 1875
(Caenogastropoda: Marginellidae) in New Caledonia.
In: Héros V., Cowrie R. H. & Bouchet P. (Eds.), Trop-
ical Deep-Sea Benthos, 25. moires du Muséum
national d’Histoire naturelle, 196: 386–436.
Boyer F., 2015a. Révision des marginelles de Linné
(Mollusques prosobranches: Marginellidae et Cystis-
cidae). Xenophora Taxonomy, 8: 33–55.
Boyer F., 2015b. Etude d’un nouveau genre de Margin-
ellidae (Mollusca: Neogastropoda) de l’Indo-Paci-
fique. Xenophora Taxonomy, 9: 31–48.
Boyer F., 2017a. Révision de l’organisation supra-spéci-
fique des gastéropodes granuliniformes. Xenophora
Taxonomy, 16: 25–38.
Boyer F., 2017b. Révision des Marginellidae du récifal
supérieur de l’île de Masirah (Oman). Xenophora
Taxonomy, 17: 3–31.
Boyer F., 2018a. Révision des Granulinidae du récifal
supérieur de l’île de Masirah. Xenophora Taxonomy,
20: 3–13.
Boyer F., 2018b. Révision des gastéropodes marginelli-
formes dans le récifal supérieur des Maldives.
Xenophora Taxonomy, 21: 22–47.
Coan E., 1965. A Proposed Reclassification of the Family
Marginellidae. The Veliger, 7: 184–194.
About the supra-generic classification of the Marginelliform Gastropods: a morphological study 235
Figure 8. Canalispira olivellaeformis Jousseaume, 1875,
lectotype MNHN, 4.1 mm.
Coovert G.A., 1986. Notes on the Genus Cystiscus and
its Type Species. Marginella Marginalia, 1: 20–24.
Coovert G.A., 1987a. The genus Afrivoluta and a review
of the type species Afrivoluta pringlei Tomlin, 1947.
Marginella Marginalia, 2: 1–8.
Coovert G.A., 1987b. A literature review and summary
of marginellid external anatomy. Marginella Margin-
alia, 3: 8–25.
Coovert G.A., 1988. Taxonomic characters in the family
Marginellidae: conchological characters. Marginella
Marginalia, 4: 43–47.
Coovert G.A., 1989. A literature review and summary of
published marginellid radulae. Marginella Margina-
lia, 7: 1–37.
Coovert G.A. & Coovert H.K., 1990. A study of margin-
ellid radulae. Part I : type 6 radula, “Prunum / Volva-
rina” type. Marginella Marginalia, 8–9: 1–68.
Coovert G.A. & Coovert H.K., 1995. Revision of the
Supraspecific Classification of Marginelliform Gas-
tropods. The Nautilus, 109: 43–110.
Espinosa J. & Ortea J., 2002. Nuevas especies de mar-
ginelliformes de Cuba, Bahamas y el Mar Caribe de
Costa Rica. Avicennia, 15: 101–128.
Fretter V., 1976. The anatomy and feeding of the volu-
tacean prosobranch Volvarina taeniolata Mörch.
Journal of Molluscan Studies, 42: 327–336.
Gofas S. & Fernandes F., 1988. The Marginellidae of São
Tomé, West Africa. Journal of Conchology, 33: 1–30.
Harasewych M.G. & Kantor Y., 1991. Rediscovery of
Marginellona gigas (Martens, 1904), with notes
on the anatomy and systematic position of the sub-
family Marginelloninae (Gastropoda: Marginellidae).
Nemouria, 37: 1–19.
Kantor Y. & A. Fedosov, 2009. Morphology and devel-
opment of the valve of Leiblein: Possible evidence
for paraphyly of the Neogastropoda. The Nautilus,
123: 73–82.
Marten E. von, 1904. Die beschalten Gastropoden der
deutschen Tiefsee-Expedition 1898–1899. A system-
atisch-geofraphischer Teil. Wissenschaftliche Ergeb-
nisse der deutschen Tiefsee-Expedition auf dem
Dampfer “Valvidia”, 7: 1–146 ; pls 1–4.
McCleery T. & Wakefield A., 2007. A review of the enig-
matic genus Canalispira Jousseaume, 1875 (Gas-
tropoda, Cystiscidae) with the description of three
new species from the western Atlantic. Novapex, 8:
1–10.
Ortea J. & Espinosa J., 2001. Intelcystiscus y Inbiocys-
tiscus (Mollusca: Neogastropoda: Cystiscidae) dos
nuevos géneros del Atlantico occidental tropical. Avi-
cennia, 14: 107–114.
Ortea J. & Espinosa J., 2016. La subfamilia Plesiocys-
tiscinae G.A. Coovert & H.K. Coovert, 1995 (Mol-
lusca: Gastropoda) en La Guadeloupe, Antillas
Menores. Revista de la Academia Canaria de Cien-
cias, 28: 65–78.
Perez-Dionis G., Ortea J., Moro L. & J.J. Bacallado J.J.,
2001. Anatomía básica de Granulina guancha (d’Or-
bigny, 1829) (Mollusca: Neogastropoda: Cystisci-
dae). Vieraea, 29: 125–130.
Ponder W.F., 1970. Some Aspects of the Morphology of
Four Species of the Neogastropod Family Marginel-
lidae with a Discussion on the Evolution of the Tox-
oglossan Poison Gland. Journal of the Malacological
Society of Australia, 2: 55–81.
Ponder W.F., 1974. The Origin and Evolution of the
Neogastropoda. Malacologia, 12: 295–338.
Ponder W.F. & Taylor J.D., 1992. Predatory shell
drilling by two species of Austroginella (Gas-
tropoda: Marginellidae). Journal of Zoology, 228:
317–328.
Smriglio C. & Mariottini P., 2001. Description of Pao-
laura gen. nov. and of three new species of Cystisci-
dae from the Indian Ocean. La Conchiglia, 33 (299):
11–17.
Souza P.J.S. Jr., 2007. Análise cladística da família Mar-
ginellidae (Mollusca, Gastropoda) baseada em mor-
fologia comparada. Tese doutorale, Instituto de
Biociências, Universidade de São Paulo. I–XI, 1–
128, figs. 1–115.
Souza P.J.S. Jr & Simone L.R., in press. Cladistic analy-
sis of the family Marginellidae (Mollusca, Gas-
tropoda) based on phenotypic features. Zootaxa.
FRANCK BOYER
236
ResearchGate has not been able to resolve any citations for this publication.
Article
Full-text available
The discovery of a comb-like radula in specimens of Dentimargo cf. aureocinctum (Stearns, 1872) is recorded. The occurrence of such a radula in a presumed non-radulate genus suggests a very close relationship between the genus Dentimargo and the genus Volvarina Hinds, 1844. This matter invalidates the taxonomic organisation currently accepted within the Marginellinae, and especially that between the Tribes Marginellini and Prunini. The conchological differenciation of Dentimargo from the Volvarina-Dentimargo common stem is shown to have arisen before the loss of the radula, and this loss is considered to have little taxonomic value within the mar-ginellid gastropods. Due to its high conchological similarity with the type species D. dentifera (Lamarck, 1803), D. cf. aureocinctum is conserved in the genus Dentimargo, which is provisionally considered as being composed both of radulate and of non-radulate species. RESÚMEN Se reseña el descubrimiento de una rádula en forma de peine en ejemplares de Dentimargo cf. aureocinctum (Stearns, 1872). La presencia de tal rádula en un género que se considera carecer de la misma indica una relación estrecha entre el género Dentimargo y el género Volvarina Hinds, 1844. Ello invalida la ordenación taxonómica actualmente aceptada entre los Marginellinae y particularmente entre las Tribus Marginellini y Prunini. Se muestra que la diferenciación conquiológica de Dentimargo a partir del estirpe común Volvarina-Denti-margo se ha producido antes de la pérdida de la rádula y que esta pérdida es de escaso valor taxonómico en gasterópodos marginéllidos. Considerando su gran semejanza con-quiológica con la especie tipo D. dentifera (Lamarck, 1803), D. cf. aureocinctum se man-tiene en el género Dentimargo, considerando en este tanto especies provistas de rádula cómo careciendo de la misma.
Article
Full-text available
Les six espèces de mollusques prosobranches marginellifor-mes décrites par Linné en 1758 sont révisées, sur la base de leur description originale, des figures de Gualtieri (1735), du matériel linnéen déposé à la Linnean Society of London, et de la documentation réunie sur le chromatisme des animaux, la morphologie des radulas et la variabilité naturelle des formes considérées. Quatre de ces espèces, Voluta persicula L., V. monilis L., V. faba L. et V. glabella L., font pour la première fois l'objet d'une révision formelle. Des lectotypes sont désignés pour V. monilis L. et pour V. faba L. Des néotypes sont désignés pour V. persicula et pour V. glabella. Le statut élusif attaché à Bulla pallida L. comme à V. monilis L. est clarifié par l'apport d'éléments inédits. Le taxon Voluta pallida décrit par Linné en 1767 est révisé, et il est proposé d'en abandonner l'emploi au bénéfice de la stabilité de la nomenclature. Le placement gé-nérique des espèces considérées est discuté et leur distribution géographique est précisée.
Article
Full-text available
As part of a project intended to review the taxa of Marginellidae worldwide, the phylogenetic relationships of its main representatives are provided based on comparative phenotypy. Characters from most structures and organs are investigated and used for a phylogenetic analysis, resulting in the following cladogram: ((((Marginellona gigas ((Eratoidea watsoni ((Prunum sp—Leptegouana guttata) (Volvarina brasiliana (Prunum prunum—P. rubens)))) (Austroginella muscaria (Marginella ealesae ((Marginella rosea (M. glabella—M. sebastiani)) (Dentimargo bruneolus (Eratoidea scalaris (Dentimargo sp—D. aureocincta)))))))) Pachycymbiola brasiliana*) Persicula sagitatta*) Buccinanops gradatus*). Those marked with * are outgroups functionally analyzed as part of the ingroup (respectively a volutid, a cystiscid and a nassariid); the root is based on Trophon geversianus (Muricidae). The genera Prunum, Marginella and Dentimargo are revealed as non-monophyletic. The monophyly of the family Marginellidae is supported by 17 synapomorphies. The volutids appear to be its sister taxon, and the possibility of Marginellidae being only a branch of Volutidae is discussed.
Article
Full-text available
ABSTRACT Three new species of three genera of the Plesiocystiscidae subfamily are described from specimens collected in the island of Guadeloupe, Lesser Antilles. Of the three new taxa illustrations of live animals and radular architecture two of them are provided.