International Journal of Paleobiology & Paleontology

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Convergent and Parallel Evolutionary Traits in Early Cretaceous

Rudist Bivalves (Hippuritidina)

Masse JP* and Fenerci Masse M

Review Article
Aix-Marseille University, France
Volume 4 Issue 1
Received Date: December 07, 2020
*Corresponding author: Jean Pierre Masse, Aix-Marseille University, Place Victor Hugo.
Published Date: January 12, 2021
13331 Marseille Cedex 03, France, Email: jean-pierre.masse@hotmail.com

Abstract

Early Cretaceous Hippuritida clades, requieniide (family Requieniidae) and hippuritide (families Radiolitidae, Polyconitidae,
Caprinidae, “Caprinulidae” and Caprinuloideidae), show distinctive myophoral arrangements and shell structures.
Nevertheless they share some characters, such as the transverse shell thickening of the myophores of the attached valve
which are convergent traits in Lovetchenia (Requieniidae) and Homopleura (Monopleuridae). The bent posterior myophore
of the right valve of Pseudotoucasia (Requieniidae) closely resemble the posterior myophore of the left valve of Horiopleura
and Polyconites (Polyconitidae). The shell cellular structure is one of the key attributes of the family Radiolitidae (e.g.
Eoradiolites) but this structure is also present in some advanced Requieniidae (“Toucasia-Apricardia “group). Canaliculate
shell structures are convergent evolutionary traits which are common in the Caprinidae and Caprinuloideidae and also exist in
the Polyconitidae and “Caprinulidae”. In most of the foregoing canaliculated groups, two trends are well expressed, reflecting
parallel evolution: expansion of canals into the entire shell and increasing complexity of canal architecture. Convergent taxa
took some advantages by using former innovations. An Albian peak of convergence coincided with the emergence of new
clades, which suggests a reset following the mid-Aptian extinction event.

Keywords: Monopleuridae; Eoradiolites; Caprinulidae; Hippuritidine Clade; Myophoral; Polyconitidae; Caprinidae

Introduction canaliculated shell structures have also been regarded as

convergent traits [6,7].
Evolutionary convergence is implicated when two
or more lineages with distinct ancestors independently The objectives of the present paper are to compile
evolve a similar morphological trait, a concept close to examples of convergent versus parallel evolution in
evolutionary parallelism which postulates that similarity Lower Cretaceous rudist bivalves (order Hippuritida)
may appear in lineages having a common origin and/or belonging to distinctive suborders or families from the
changed together in roughly similar fashion [1,2]. In bivalves, Mediterranean, Caribbean and Asiatic paleobiogeographic
evolutionary convergence was established for post-Paleozoic entities. We first address two clades, the requieniide clade
Crassatellidae and Astartidae [3]. It was also documented and the hippuritidine clade [8,9] (Figure 1) in which the
for the acquisition of an opisthogyrate rostrate shell habit arrangement of myophores and the shell structure show
observed in infaunal taxa belonging to unrelated lineages, as some similar patterns. We will also reappraise the similarity
an adaptation to shallow burrowing and allowing the animal in distribution of canaliculated shell structures in distinctive
to reach the water-sediment interface while the main body families of the hippuritidine clade, and subclades, including
remained below the sediment surface [4,5]. In rudist bivalves their evolution through time. We discuss the relevance of the

Convergent and Parallel Evolutionary Traits in Early Cretaceous Rudist Bivalves (Hippuritidina) Int J Paleobiol Paleontol
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concept of convergence and/or parallelism applied to the myophoral organisations recorded in distinctive clades may
foregoing pattern of characters and analysed their relative suggest phylogenetic affinities or may afford the opportunity
timing. In the following we use the family names and authors to evaluate other patterns.
proposed by Skelton [8,9].
The Requieniide Clade
In the requieniide clade the shell is attached by the
left valve (LV). Myophoral evolutionary patterns are well
illustrated in the family Requieniidae Kutassy, which includes
five genera having contrasting myophoral arrangements
[10,11]. In Matheronia right valve (RV) myophores are located
on low, salient bulges, whereas the LV myophores are on shell
wall. In Requienia the RV is flattened and bears a posterior
myophoral plate, the anterior is a low bulge; LV myophores
being inconspicuous. Toucasia is characterized by posterior
myophoral plates on both valves and the posterior side of
the RV is raised above the commissure. The myophores of
Lovetchenia are represented by shell transverse thickenings
on both valves, and the RV is salient (Figure 2A) As Toucasia,
Pseudotoucasia has posterior myophoral plates on both
valves but the myophore of the RV is bent with an elbow
shape (Figure 3A).

The Hippuritidine Clade

In the hippuritidine clade the shell is attached by the
RV. Myophoral evolutionary patterns are well illustrated
in two families, Monopleuridae Munier -Chalmas and
Polyconitidae MacGillavry. As stated by Masse and Fenerci-
Masse [12], the family Monopleuridae includes two groups
of genera. To the first group belong Monopleura Matheron
emend. and Homopleura Masse and Fenerci-Masse [12]
with RV myophores located on flat transverse shell
thickenings present either only on the posterior side, as in
Monopleura or on both the anterior and posterior sides, as
in Homopleura (Figure 2B). In the family Polyconitidae the
Figure 1: Suprageneric classification of Hippuritida [8,9].
LV posterior myophore is a bent lamina which consists of
The requieniide clade is in yellow, the hippuritidine clade
a plate subparallel to the commissure, attached basically to
is represented by two colours: green for the radiolitoidine
the shell by a pedicle, (Figure 3B), the anterior myophore is
subclade and orange for the caprinoidine subclade.
a vertical plate, in Horiopleura the RV posterior myophore is
on a transverse shell thickening , the anterior on shell wall
[13]. In Polyconites the myophoral organisation of the LV
Abbreviations and codes used for the paleontological
is nearly the same, the RV myophores being on shell wall.
descriptions: BC, body cavity; A, anterior side; P, posterior
Pediculate myophores also exist in some Caprinuloideidae
side; m, anterior and posterior muscles; am, anterior
Damestoy, this is the case for the LV posterior myophore of
myophore; pmp, posterior myophoral plate; pmc, posterior
Amphistricoelus warringi Harris and Hodson , represented
myophoral cavity pt, posterior tooth; at, anterior tooth; as,
by a vertical, slightly convex, plate supported by a pedicle
anterior socket; ps, posterior socket. Muscles are figured in
attached to the posterior tooth [14] (Figure 3C).
pale blue.
Convergent Patterns in Myophoral Arrangement
Myophoral Arrangement in Rudist Bivalves
The foregoing shows convergent patterns between
Myophores are fundamental in rudist systematics
the two clades, requieniide (family Requieniidae) and
and their diversity and phylogenetic conservatism are
hippuritidine (families Monopleuridae, Polyconitidae
remarkable and possess a high taxonomic value [6,7]. Similar
and Caprinuloideidea), whereas the convergent traits are

Masse JP, et al. Convergent and Parallel Evolutionary Traits in Early Cretaceous Rudist Bivalves Copyright© Masse JP, et al.
(Hippuritidina). Int J Paleobiol Paleontol 2021, 4(1): 000120.
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recorded on the RV of the first group, they are located in the the Requieniidae and primitive forms of the Polyconitidae,
LV of the second group. it has a generic significance in Pseudotoucasia whereas in
the Polyconitidae it is a marker of the family. The posterior
In Monopleura (Monopleuridae), Matheronia and myophore of the LV of Amphistricoelus (Caprinuloideidae) is
Requienia (Requieniidae), the myophoral arrangement of also pediculate (Figure 3C), the pedicle supports a vertical
the attached valve is similar. This similarity is recorded plate facing the posterior shell side [14].
in primitive taxa of the two families. The transverse shell
thickening of the myophores of the attached valve are
convergent traits in Lovetchenia, i.e. L. lovetchensis (Zklatarski)
(Requieniidae) (Figure 2A) and Homopleura, i.e. H.balkanica
Masse and Fenerci-Masse (Monopleuridae) (Figure 2B); this
morphology characterises relatively advanced taxa. In that
case morphological identity corresponds with functional
similarity. Inflated transverse myophores appear first in the
requieniide clade in the Hauterivian then in the hippuritidine
clade in the Barremian, it may reflect the resurgence of a
primitive trend (atavism) recorded in diceratids [15].

Figure 3: Convergent myophoral arrangement: bent (elbow

shape) or pediculate myophores. A, B, longitudinal sections
of bivalve specimens. A, Pseudotoucasia santanderensis
(Douvillé), from Santander (Spain) [17]. B, Horiopleura
gemmellaroi (di Stefano), from Northern Tunisia [13].
C, transverse section of a RV of Amphitriscoelus warringi
Harris and Hodson, fromTrinidad [14]. Arrow indicates
the pediculate portion of the myophore. For Figure A and
B, the shell layers are represented as in Figure 2.

Shell Structures
Figure 2: Convergent myophoral arrangement: transverse
shell thickening, figures based on longitudinal sections of The basic shell structure of rudist bivalves is two layers,
bivalve specimens. A, Lovetchenia lovetchensis (Zlatarski), i.e. an inner shell layer, originally made of aragonite, from
from Lovetch, Bulgaria (Requieniidae) [15]. B, Homopleura which myocardinal features are built, and an outer shell
balkanica Masse and Fenerci-Masse, from Emen, Bulgaria layer made of prismatic calcite [18]. In most taxa, but one,
(Monopleuridae) [12]. Inner aragonitic shell layer is in belonging to the requieniide clade, the double layer has
grey, the calcitic outer shell layer in black. a compact microstructure. In the Lower Cretaceous the
hippuritine clade is also dominated by taxa having a compact
The bent posterior myophore of the RV of Pseudotoucasia double layering, but two additional arrangements have their
(Requieniidae) closely resembles the posterior myophore FO during this time interval: cellular structures present in
of the LV of Horiopleura and Polyconites (Polyconitidae). In the calcitic outer shell layer, and canaliculated structures
Pseudotoucasia santanderensis (Douvillé) the myophore restricted to the inner formerly aragonitic shell layer. During
is elbow shaped (Figure 3A) and its tip points posteriorily, the early Cretaceous the two basic structures didn’t coexist
which mimics that of Horiopleura (Figure 3B). A similar whereas they were coeval in some late Cretaceous forms, e.g.
myophoral organisation was described by Ayoub-Hannaa the Campanian Pseudosabinia klinghardti (Boehm) [19,20].
and Fürsich [16] in the Cenomanian requieniid “Apricardia”
noncarinata from Egypt. The morphological similarity Cellular Versus Compact Structures
does not imply a functional identity in Pseudotoucasia for
The so-called cellular structure has been attributed
instance the muscle is inserted at the inflated apex of the
to the association of funnel plates and radial plates [21],
myophore whereas in the Polyconitidae the muscle insertion
forming a network which is one of the key attributes of
is on the plate itself which parallels the commissural plan.
the family Radiolitidae d‘Orbigny [22]. Early Cretaceous
This myophoral geometry typifies an advanced taxon of

Masse JP, et al. Convergent and Parallel Evolutionary Traits in Early Cretaceous Rudist Bivalves Copyright© Masse JP, et al.
(Hippuritidina). Int J Paleobiol Paleontol 2021, 4(1): 000120.
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representatives of the family are represented by Eoradiolites in Sellaea on the RV anterior shell margin, but not in the LV
with a quadrangular outer shell layer structure and perimyophoral cavity, usually canaliculated in the Caprinidae.
Archaeoradiolites with radially arranged branching walls. The existence of anterior and posterior canals is recorded in
Neocaprina raghawiensis, (Figure 4C) a putative descendant
It is worth noting that the primitive, Barremian member of Sellaea [35]. Canals are not uncommon in Polyconitidae.
of the family, i.e. Agriopleura, has a compact outer shell The RV of Polyconites operculatus (Roulland) ( pro Radiolites
[23,24]. A similar acellular structure, is recorded in the polyconilites d’Orbigny, 1842) has dorsal,vertical canals
late Aptian-Albian radiolitid genus Auroradiolites, assumed (see d’Orbigny, 1842, pl. 574, Figure 3 ).The LV posterior
to be a marker of the Asian-Pacific province [25,26]. perimyophoral cavity of Magallanesia canaliculata Sano, et al.
The acellular Auroradiolites was considered as a direct [36] (Figure 4D) and M. rutogensis Rao, et al. [25], bear a row
descendant of Agriopleura [26], in that case the compact of radial canals. In the LV posterior perimyophoral cavity of
shell structure might be just inherited. Given the distinctive Praecaprotina yegashii (Yehara) canals are present in some
paleogeography (Asiatic versus Mediterranean) and the specimens [37]. The foregoing data are a perfect illustration
time gap (Barremian-early Aptian versus late Aptian-Albian) that canals evolved independently in several different clades
between the two genera, an alternative interpretation is [18,7]. So they are typical convergent evolutionary traits.
proposed. Auroradiolites might derive through convergence
from a former cellular form, e.g. Eoradiolites. Actually the
development of cellular structures in advanced radiolitids
shows that this pattern follows a juvenile compact stage
through ontogeny [27], heterochrony is thus a more likely
interpretation of the compact structure of Auroradiolites.
Another case of convergence is illustrated by the genus
Gorjanovicia which documents the re-inception of acellular
Radiolitidae in the late Cretaceous [28].

Cellular structures tend to characterize the hippuritidine

clade, nevertheless they have been also recorded in
“Toucasia like” requieniids (unpublished data from the
authors). Quadrangular cellular structures are present on
the ventral side of the LV of specimens of Albian age from
Laguna Colorada-El Madrono (Queretaro, Mexico) [29] and
southern Italy. Similar structures were described by Palmer Figure 4: Pallial canals in Caprinidae, “Caprinulidae”
[30] in the requienid “Apricardia” asymmetrica Palmer from and Polyconitidae, based on transverse sections. A, B
Paso del Rio, Colima (Mexico) in beds of Albian age (formerly Caprinidae. A, LV of Praecaprina varians Paquier, from
assigned to the Cenomanian). SE France [14]. B, LV of Offneria simplex Chartrousse
and Masse, from SE France [38]. C “Caprinulidae”. LV of
Canaliculate Structures Neocaprina raghawiensis Steuber and Bachmann, from
Sinai Peninsula [35]. D, Polyconitidae. LV of Magallanesia
Canals tend to develop between vertical blades, when
canaliculata Sano et al., from Cebu island, Philippines [36].
aligned along the outer shell margin one refers to pallial
Arrow indicates the canal-free ventral side.
canals. They originate from the partitioning of cavities or
gutters and have a significant record in several families
with Lower Cretaceous representatives, e.g. Caprinidae
Evolutionary Parallelism
d’Orbigny and Caprinuloideidae. Canals have their FO in In the families Caprinidae, Caprinuloideidae and
Pachytraga tubiconcha Astre (Caprinidae) of Hauterivian “Caprinulidae” the geometry, mode of development and
age [31,32]. They play a major role in Caprinidae, e.g. expansion of canals in the shell are nearly identical and
Praecaprina varians Paquier (Figure 4A), Offneria (Figure evolved similarly. As shown by Coogan [39] and Coogan in
4B) and the Caprinuloideidae, e.g. Amphistroelus warringi Dechaseaux, et al. and Mitchell [40,41], canals of primitive
(see Figure 3C). Pallial canals are present in the unrelated forms are characterized by few simple radial walls, as in
clade of “Caprinulidae” Yanin, 1990 (a family not recognized Praecaprina (Caprinidae) (Figure 4A) , Amphitriscoelus
by Rineau, 2017, but independent from the Caprinidae), (Caprinuloideidae)(Figure 3C) and Neocaprina
including Sellaea and Neocaprina [33,34]. Assuming a (Caprinulidae) (Figure 4C ). In these families the ventral
phylogenetic relationship between Himeraelites and Sellaea side of primitive forms lack pallial canals, this is the case
it is worth noting that canals, absent in Himeraelites, settled for Praecaprina (Figure 3A), Amphistriscoelus (Figure 3C),

Masse JP, et al. Convergent and Parallel Evolutionary Traits in Early Cretaceous Rudist Bivalves Copyright© Masse JP, et al.
(Hippuritidina). Int J Paleobiol Paleontol 2021, 4(1): 000120.
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Neocaprina raghawiensis Steuber and Bachman [35] (Figure invasion of the ventral margin, 2- the increasing complexity
4C) and the primitive Offneria simplex Chartrousse and Masse of canal architecture through time, two phenomena
(Figure 3B). By contrast, in more advanced forms, the number reflecting parallel evolution of distinct lineages rather than
of canals increases and in some cases invade the ventral side, convergence.
as in advanced species of Offneria (Figures 5B, 5C) [14,42],
and Coalcomana (Figure 5D) [14]. This trend is not only
recorded in caprinids but also in the Polyconitidae. This is
the case of species of Magallanesia: the derived M.rutogensis
Rao, et al. [25] has more canals than its putative ancestor
M. canaliculata Sano et al. In the “Caprinulidae” too, the
advanced Cenomanian species Neocaprina gigantea Plenicar
[43-45] is more canaliculated than its Albian ancestor N.
raghawiensis (Figure 4C and 5A). Quantitative changes in
canal number are associated with modifications of their
geometry: they bifurcate and tend to be subdivided, this is
the case in the Caprinidae for species of Offneria (Figures
5B, 5C) [38], and in the Caprinuloideidae with Coalcomana
ramosa (Boehm) (Figure 5D) and Caprinuloidea [14]. The
geometrical changes are associated with an increasing
structural complexity and the existence of several canal
rows with distinctive geometries, a trend well illustrated
in species of Offneria with O. simplex, a Barremian species, Figure 5: Canaliculate architectures in advanced forms of
followed by O. rhodanica of late Barremian-early Aptian age “Caprinulidae”, Caprinidae and Caprinuloideidae, based on
and O. prebetica of the latest early Aptian (Figure 4B, 5B, 5C) transverse sections. A, RV of Neocaprina gigantea Plenicar,
[24,38]. fromTurkey [46]. B, LV of Offneria prebetica Masse et al.,
from Southern Spain [24]. C, LV of Offneria rhodanica
Convergence appears somewhat unlikely to account Paquier, from SE France [14]. D, LV of Coalcomana ramosa
for: 1- the expansion of canals into the entire shell, i.e. the (Boehm), from Mexico [14].

Timing of Evolutionary Convergent Traits (Figure 6)

Figure 6: Timing of evolutionary convergent and parallel traits in early Cretaceous rudists. Red points mark the convergent
taxa, green points their ancestors.

Masse JP, et al. Convergent and Parallel Evolutionary Traits in Early Cretaceous Rudist Bivalves Copyright© Masse JP, et al.
(Hippuritidina). Int J Paleobiol Paleontol 2021, 4(1): 000120.
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Pediculate posterior myophores are recorded in the requieniide (family Requieniidae) and hippuritide (families
Mediterranean hippuritide clade, in the Barremian -early Radiolitidae, Polyconitidae, Caprinidae, “Caprinulidae”
Aptian genus Horiopleura, their Caribbean counterparts are and Caprinuloideidae), possess distinctive myophoral
nearly contemporaneous, e.g. the early Aptian Amphitriscoelus arrangements and shell structures. But they share some
warringi. The convergent requieniide Pseudotoucasia, is similar characters located in the RV of the first group and in
younger, late Aptian- Albian. Cellular structures have their the LV of the second group. The transverse shell thickening
FO in the late Aptian radiolitid genera Archaeoradiolites of the myophores of the attached valve are convergent
and Eoradiolites, but the convergent requieniide traits in Lovetchenia (Requieniidae) and Homopleura
representative “Toucasia-Apricardia group”, is younger, (Monopleuridae). The bent posterior myophore of the RV
Albian. Acellular species of Radiolitidae, i.e. Auroradiolites, of Pseudotoucasia (Requieniidae) closely resembles the
are late Aptian-Albian. Canal system developed in late posterior myophore of the LV of Horiopleura and Polyconites
Hauterivian- Barremian and early Aptian, in Caprinidae and (Polyconitidae). The shell cellular structure is one of the key
Caprinuloideidae, and evolved parallel patterns. By contrast attributes of the family Radiolitidae (e.g. Eoradiolites) but
canaliculate Polyconitidae and “Caprinulidae” started to this structure is also present in some advanced Requieniidae
expand in the Albian and Cenomanian. (“Toucasia-Apricardia” group). The restatement of acellular
Radiolitidae is a late Aptian -Albian event connected with the
The foregoing shows that a peak in convergent traits Asian -Pacific faunal province. Canaliculate shell structures
is in the late Aptian- Albian, mostly in the context of the which are common in the Caprinidae and Caprinuloideidae
“Albian radiation” of Rineau [7]. This radiation is marked by are also present in Polyconitidae and “Caprinulidae”, and
emergence of new clades and, in the Caribbean, by a diversity have long been considered as convergent evolutionary traits.
peak in the caprinuloids [47] with a strong turnover of the In the foregoing canaliculated groups, with the exception
Youngicaprininae [41]. The late Aptian-Albian events suggest of the Polyconitidae two trends are well expressed, 1- the
a reset following the mid-Aptian extinction event [33,48]. expansion of canals into the entire shell, i.e. the invasion
into the ventral margin, 2- the increasing complexity of canal
architecture through time, these two phenomena reflect
Discussion parallel evolution. Convergent taxa took some advantages
by using former innovations. The late Aptian-Albian peak of
The strategy of pediculation implies that by contrast
convergence coincides with the emergence of new clades,
with a myophore on shell wall, muscles were not supported
mostly in the Caribbean and South Tethyan Mediterranean
by a double (calcite+aragonite) shell layer, but a single nacre-
margins, it suggests a reset following the mid-Aptian
made layer. Actually nacre consists of aragonitic nannobricks
extinction event.
and a complex organic phase, having a high tensile strength,
high elasticity and high resistance to breaking [49,50].
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(Hippuritidina). Int J Paleobiol Paleontol 2021, 4(1): 000120.
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