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Phylogenetic relationships of the family Carditidae (Bivalvia:

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DOI: 10.1080/14772019.2018.1532463

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Phylogenetic relationships of the family Carditidae

(Bivalvia: Archiheterodonta)

Damián Eduardo Pérez

To cite this article: Damián Eduardo Pérez (2019): Phylogenetic relationships of the
family Carditidae (Bivalvia: Archiheterodonta), Journal of Systematic Palaeontology, DOI:
10.1080/14772019.2018.1532463

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 Journal of Systematic Palaeontology, 2019
Vol. 0, No. 0, 1–37, http://doi.org/10.1080/14772019.2018.1532463

Phylogenetic relationships of the family Carditidae (Bivalvia: Archiheterodonta)

Dami
an Eduardo P
erez



Museo Argentino de Ciencias Naturales Bernardino Rivadavia, Divisi
on Paleoinvertebrados, Av. Angel Gallardo 470 (C1405DJR),
Buenos Aires, Argentina
(Received 15 September 2017; accepted 6 September 2018)

Carditidae is a highly speciose group of bivalves but is one of the more neglected and poorly studied families. This work
presents the first phylogenetic study of the entire family, using 77 species-level taxa (including fossil and recent terminals) and
144 new morphological characters. Multiple searches were conducted, including comparisons of discrete and continuous
characters and equal and implied weighting. Morphological disparity analysis, phylomorphospace construction and
phylogenetic signal analysis were performed using the character matrix. A new systematic scheme is suggested on the basis of
these results, including the already proposed subfamily Palaeocarditinae, the re-defined Carditinae, Thecaliinae (included
within Carditinae in some searches), Carditamerinae (with reduced diversity) and Venericardiinae (with extended diversity).
The new subfamily Scalaricarditinae and the new clades Eucarditidae (all carditids excluding Palaeocarditinae),
Carditida (Carditinae þ Thecaliinae þ Carditamerinae) and Cyclocardida (Cyclocardia þ Pleuromeris þ Scalaricarditinae
þ Miodomeridinae) are proposed. The monophyly of the planicostate and alticostate lineages are confirmed within the
Venericardiinae (now formally named Venericorini and Venericardiini, respectively). The relationships of Palaeocarditinae with
other carditids and the origin of the family are discussed, as well as the phylogenetic placement of other carditid genera not
included in the analyses. Homoplasy and the phylogenetic signals of different morphological characters are also explored
and discussed.

Keywords: Carditidae; Bivalvia; Archiheterodonta; phylogeny; systematics

Introduction 2004). According to Coan (1977, p. 375), scarcity of

Carditidae F
erussac, 1822 is a large group of molluscs
that is distributed worldwide, but is one of the more
neglected and poorly studied bivalve families (Huber
2010). Recent revisions recognized 22 living genera and
nearly 140 species within this family (Huber 2010),
with approximately 34 known fossil genera. According
to Chavan (1969), the fossil record of Carditidae begins
in the Devonian with the presence of the questionable
species Carditomantea spinata Quenstedt, 1926
(Svalbard, Arctic Ocean), followed by a sparse
Mesozoic record. The maximum diversity of the group
is found during the Palaeogene, with the emergence of
most of the modern taxa. Many fossil carditids were
assigned to the genera Cardita or Venericardia but most
of these assignments were never reviewed. Despite poor
systematic research on the group, carditid bivalves were
the subject of several studies, including reproductive
biology (Yonge 1969), evolutionary and ontogenetic
analyses (Stanley 1970; Gould 1977) and, mainly, bio-
stratigraphy (Conrad 1830; Burckhardt 1902; Gardner &
Bowles 1939; Camacho & Fern
andez 1956; del R
ıo

Corresponding author. Email: trophon@gmail.com
knowledge on this family is a consequence of their
‘bewildering degree of morphological variability’. The
reason for this variability is the existence of sexual
dimorphism in several species, high intraspecific vari-
ation and the presence of ontogenetic changes (Heaslip
1968; del R
ıo 1995; P
erez & del R
ıo 2012; P
erez et al.
2017b). Due to these issues, and the long evolutionary
history of the group, the phylogenetic relationships of
Carditidae remain obscure.
The aim of this contribution is to generate a new and
comprehensive systematic scheme, and morphological
delimitation of carditid lineages, using a set of different
quantitative phylogenetic tools and approaches, and to
shed light on the evolutionary history of this particular
group of archiheterodont bivalves.
Early work on Carditidae relationships
Lamarck (1806) grouped the genera Cardita Bruguiere,
1792 and Venericardia Lamarck, 1801 together and
generated the first taxonomic concept of the family

# The Trustees of the Natural History Museum, London 2019. All rights reserved.

Published online 15 Jan 2019

2 D. E. P
erez
Table 1. Composition of subfamilies proposed by Chavan (1969). Some of these genera are now considered either as not
Carditidae(1) or junior synonyms of other taxa(2), see text for further details. The genera Birkelundita, Darwinicardia, Kalelia,
Kolmeris, Neovenericor, Powellina, and probably Fasciculicardia and Rotundicardia, were unknown to Chavan.
Subfamily
Palaeocarditinae
Carditinae
Carditamerinae
Miodomeridinae
Venericardiinae
Carditesinae
Thecaliinae
Carditidae. F
erussac (1822) erected a formal definition
and related this family to Mytilidae, Unionoidea and
Crassatellidae. Dall (1902) and Thiele (1935) placed the
families Carditidae and Condylocardiidae within the
group Carditacea. Chavan (1969) created the superfam-
ily Carditoidea, including Carditidae, Condylocardiidae
and the fossil family Permophoridae. Healy (1995)
related carditids with crassatellids based on spermato-
zoid ultrastructure. Recently, Carditidae was placed into
the newly described clade Archiheterodonta based on
molecular tools (Giribet 2008; Bieler et al. 2014).
Archiheterodonta includes the families Carditidae,
Astartidae, Crassatellidae and Condylocardiidae, but its
internal phylogenetic relationships remain poorly under-
stood (Giribet 2008; Gonzalez & Giribet 2015).
There are very few studies about the phylogenetic
relationships within carditids. The earliest study was
published by Conrad (1830), who proposed the exist-
ence of a group of North American carditids related to
the European species Venericardia planicosta Lamarck,
1801. These taxa are characterized by the presence of
large shells, sculptured with smooth radial ribs.
Subsequently, Harris (1919) named this lineage as the
‘planicosta group’ and also described the ‘rotunda-alti-
costata group’, consisting of North American carditid
species with small, subcircular shells and high radial
ribs covered by nodes. According to Harris (1919), the
hallmark of the ‘rotunda-alticostata group’ was the par-
ticular structure of the radial ribs, named as ‘terraced’
by him, with a central cord flanked by paracostal ribs
on each side. These two lineages include species previ-
ously assigned to Venericardia, but no Cardita species.
Stewart (1930) recognized these groups and proposed
the genus Venericor for the planicostate carditids.
Gardner & Bowles (1939), Verastegui (1953), Stenzel &
Genera included
Palaeocardita, Carditomantea, Schizocardita, Pseudopis1
Cardita, Jesonia2, Beguina
Carditamera, Lazariella, Glans, Centrocardita, Goossensia,
Carditella1, Carditellona1, Cyclocardia, Scalaricardita,
Plionema1, Vimentum, Fenestricardita, Vetericardiella2,
Miodontiscus, Tutcheria1, Choniocardia, Carditellopsis1,
Arcturellina, Izumicardia, Pleuromeris, Cossmannella,
Cretocardia, Cardiocardita, Bathycardita
Miodomeris, Chavanella2, Pteromeris, Coripia
Venericardia, Venericor, Leuroactis, Pacificor, Megacardita,
Trapezicardita
Glyptoactis, Claibornicardia, Baluchicardia, Ludbrookia2,
Xenocardita, Paraglans, Cardites
Thecalia, Milneria
Krause (1957) and Heaslip (1968) used this approach
for North American and European carditids, and
Maxwell (1969) considered it for New Zealand repre-
sentatives. After these authors, the clades began to be
named ‘planicostate’ and ‘alticostate’ carditids respect-
ively. Recently, McClure & Lockwood (2015) carried
out the first quantitative phylogenetic analysis of North
American carditids and discussed the monophyly of
these lineages. The placement of some taxa
(Megacardita Sacco, 1899; ‘Venericardia’ pectuncularis
Lamarck, 1806) within these lineages was little dis-
cussed (Stewart 1930; Verastegui 1953; McClure 2009).
The planicostate-alticostate proposal is, currently, the
only hypothesis for the phylogeny of Carditidae.
Dall (1902) was the first to propose a subfamiliar
classification within Carditidae, with the subfamilies
Carditinae and Thecaliinae, the latter including only two
genera: Thecalia Adams & Adams, 1858 and Milneria
Dall, 1881. The purpose of this author was to separate
the carditids with conspicuous sexual dimorphism into a
subfamily, because Thecalia and Milneria are character-
ized by the presence of an incubatory chamber in female
shells. Subsequently, the only systematic scheme for the
family was suggested by Chavan (1969) (Table 1). This
author re-defined the subfamily Carditinae (including
only mytiliform or modioliform genera), included the
subfamily Thecaliinae of Dall (1902), and defined five
new subfamilies, based mainly on shell outline and
hinge configuration: Carditamerinae, Miodomeridinae,
Palaeocarditinae, Venericardiinae and Carditesinae. The
morphological boundaries between some of these subfa-
milies are not clear, and the limits among different gen-
era and subgenera have been subject to argument and
used with dissimilar criteria by subsequent authors. The
systematic subfamiliar scheme of Chavan (1969) ignores
 Phylogeny of Carditidae
the previous proposals of the planicostate-alticostate lin-
eages, whose taxa were classified in the subfamilies
Carditamerinae, Carditesinae or Venericardiinae. Despite
this incongruence, there have been no revisions of the
subfamiliar classification scheme of Chavan (1969),
except for that of Scarlato & Starobogatov (1979). The
latter authors did not discuss the taxonomic composition
of Chavan’s subfamilies but proposed the rearrangement
of Carditidae, considering the family as a superfamily,
including the Mesozoic families Aenigmoconchidae and
Archaeocardiidae, and also the families Carditidae
(from Carditinae), Thecaliidae (from Thecalinae),
Miodomerididae (from Miodomeridinae) and
Carditameridae (including Carditamerinae,
Venericardiinae, Palaeocarditinae and Carditesinae).
Adegoke (1977) pointed out that this arrangement failed
to clarify the taxonomic problems of the family. Due to
the absence of revisions, the first phylogenetic analysis
of the whole family Carditidae is performed here, based
on morphological shell-characters. This analysis includes
representatives of all known subfamilies and proposed
lineages with fossil and recent taxa, and represents the
first attempt to revisit the previous subfamiliar system-
atic scheme.
Material and methods
Taxonomic sample
The systematics of the carditids is a complex subject
due to the lack of consensus in the delimitation of gen-
era and subgenera (Adegoke 1977). On one hand, the
family includes only a few genera (e.g. Venericardia
Lamarck, 1801; Cardita Bruguiere, 1792), which
include a large number of subgenera (e.g.
Claibornicardia Stenzel & Krause, 1957; Venericor
Stewart, 1930; Megacardita Sacco, 1899). On the other
hand, some of these taxa have been considered as gen-
era (for a comparison of different approaches see Sacco
1899; Dall 1903; Chavan 1969; Moore 1992; McClure
& Lockwood 2015). The use of subgenera implies close
phylogenetic relationships among species or species-
groups within a genus (Dubois 1982) and, as a result, in
order to avoid a priori phylogenetic inferences, all cardi-
tid genera or subgenera are considered to have generic
status in this work.
The phylogenetic analysis was performed using 77
species-level terminals. Four non-carditid species were
included as outgroups: the fossil crassatellids
Crassatella ponderosa (Gmelin, 1791) and Spissatella
trailli (Hutton, 1873) and the astartids Astarte sulcata
(da Costa, 1778) and Laevastarte (¼Astarte) basteroti
(de la Jonkaire, 1823). These outgroups represent two
3
widely accepted, closely related-groups to carditids, the
archiheterodont families Crassatellidae and Astartidae
(Giribet 2008; Gonzalez & Giribet 2015). The fourth
archiheterodont family, Condylocardiidae, is problematic
because of the tiny size of its shells and a lack of infor-
mation (Middelfart 2002; Gonzalez & Giribet 2015).
The fossil record of this family is poor (Olsson 1942;
Ludbrook 1953; Kensley & Pether 1986; Ward &
Blackwelder 1987; Maxwell 1992; Middelfart 2002) and
molecular phylogenies place it within Carditidae
(Gonzalez & Giribet 2015), but they have distinctive
morphologies not recognized in true carditids, like the
presence of an internal resilifer. Further studies on this
group are needed.
The ingroup comprises 73 carditid species that belong
to 38 genera. This represents 66% of the currently valid
carditid genera described in the bibliography. All
Carditidae subfamilies proposed by Dall (1903) and
Chavan (1969) are represented in this analysis, including
their type genera and other representatives, as well as
representatives of the planicostate and alticos-
tate lineages.
The Palaeocarditinae is represented by two species of
Palaeocardita Conrad, 1867, namely P. crenata
(M€unster, 1841) (Late Triassic, Europe) and P.
stoecklini Hautmann, 2001 (Late Triassic, Iran), and
Schizocardita cristata K€orner, 1937 (Late Triassic,
Per
u). Chavan (1969) proposed Carditomantea
Quenstedt, 1929 (Early Devonian, Svalbard) as a palaeo-
carditine, but this assignment is doubtful because of the
sparse information on hinge morphology in this
Palaeozoic taxon. Morris & Eagar (1978) proposed that
there is no reason to place Carditomantea in Carditidae.
Another putative palaeocarditine is Pseudopis Cox, 1946
(Early Jurassic, UK), which probably belongs to the
Astartidae because of its hinge configuration and exter-
nal sculpture (Cox 1946; Zakharov 1970). Tutcheria
Cox, 1946 (Late Triassic–Early Jurassic, Eastern Asia,
Europe, South America and North America) was placed
within Carditamerinae by Chavan (1969) but its hinge
morphology resembles that of Pseudopis and it may rep-
resent another Astartidae, as mentioned by Ichikawa &
Maeda (1963). Further revisions are needed on these
genera in order to discuss their relationships.
Septocardia Hall & Whitfield, 1877 (Late Triassic,
South America, North America and Russia) was placed
as a basal member of Carditidae (Schneider 1995;
Schneider & Carter 2001; Ros-Franch et al. 2014) or
Cardiidae (Keen 1969; Damborenea & Mance~nido
2012) because of its hinge configuration, but its rela-
tionships also require new studies.
Two genera of Carditinae were included, represented
by three species: Cardita Bruguiere, 1792, with C.
4 D. E. P
erez
variegata Bruguiere, 1792 (type species of the genus:
Pleistocene–Recent, Indian Ocean) and C. crassa
Lamarck, 1819 (Miocene, France), and Beguina R€oding,
1798 with B. semiorbiculata (Linnaeus, 1758) (type spe-
cies of the genus: Recent, Indian Ocean and Australia).
The third carditine genus, Jesonia (Gray, 1847) (Recent,
Western Africa), was synonymized with Cardita by
Lamy (1922) and Huber (2010). There is some confu-
sion about the type species of Cardita because
Bruguiere (1792) originally did not designate any spe-
cies name and more recent authors named C. sulcata
(Children, 1823) or C. calyculata (Gray, 1847) as its
type species, whereas others considered C. variegata as
a synonym of C. calyculata (Lamarck 1801; Reeve
1843). Fleming (1818) was the first to establish C. vari-
egata as the type species of the genus (Winckworth
1929; Maxwell 1992).
The highly diverse subfamily Carditamerinae includes
24 genera according to Chavan (1969), but some of
them are not considered carditids nowadays, namely
Carditella Smith, 1881, Carditellopsis Iredale, 1936 and
Carditellona Iredale, 1936, (included in
Condylocardiidae: Huber 2010) and Plionema Conrad,
1872 (included in Crassatellidae: Gerhardt 1897).
Others, such as Miodon Carpenter, 1863 and
Vetericardiella Chavan, 1969, are not valid due to syn-
onymy (with Miodontiscus and Pseudocardia, respect-
ively: Vokes 1967). This analysis considered the genus
Carditamera Conrad, 1838, with the species C. arata
(Conrad, 1838) (type species of the genus:
Miocene–Recent, North America), C. floridana (Conrad,
1838) (Holocene–Recent, North America), C. affinis (G.
B. Sowerby I, 1823) (Pliocene–Recent, Central and
North America) and C. gracilis (Shuttleworth, 1856)
(Pliocene–Recent, Central America), Arcturellina
Chavan, 1951 as represented by A. asperula (Deshayes,
1825) (type species of the genus, late Eocene, France),
Cardiocardita Anton, 1838 by Ca. ajar (Bruguiere,
1792) (type species of the genus: Recent, western
Africa), Bathycardita Iredale, 1925 by B. raouli (Angas,
1872) (type species of the genus: Recent, Australia),
Cyclocardia Conrad, 1867 with the species Cy. borealis
(Conrad, 1831) (type species of the genus:
Pleistocene–Recent, Eastern North America), Cy. awa-
moensis (Harris, 1897) (early Miocene, New Zealand),
Cy. ventricosa (Gould, 1850) (Recent, western North
America), Cy. mesembria (Stilwell & Zinsmeister, 1992)
(Eocene, Antarctica), Cy. compressa (Reeve, 1843) (late
Pleistocene–Recent, southern South America), Cy. can-
nada (Ihering, 1907) (early Miocene, Argentina), Cy.
dalek P
erez & del R
ıo, 2017b (early Miocene,
Argentina) and Cy. nortensis del R
ıo, 1986 (late
Miocene, Argentina), Scalaricardita Sacco, 1899 with S.
scalaris (J. de C. Sowerby, 1825) (type species of the
genus: late Miocene–late Pliocene, Europe) S. laciarina
(Feruglio, 1954) (early Pliocene, Argentina) and S.
camaronesia (Ihering, 1907) (early Miocene, Argentina),
Vimentum Iredale, 1925 by V. dilectum (Smith, 1885)
(Recent, Australia), Glans Megerle, 1811 by G. trapezia
(Linnaeus, 1758) (type species of the genus: Recent,
Mediterranean Sea and Western Africa), Centrocardita
Sacco, 1899 by Ce. aculeata (Poli, 1795) (type species
of the genus: Recent, Mediterranean Sea), and
Pleuromeris Conrad, 1867 with P. decemcostata
Conrad, 1867 (type species of the genus:
Pliocene–Pleistocene, southern North America), P. tri-
dentata (Say, 1826) (Pliocene–Recent, southern North
America), P. fueguina (Steimann & Wilckens, 1908)
(early–middle Miocene, Argentina), P. sulcolunularis
(Ihering, 1907) (early Miocene, Argentina) and P.
zelandica (Deshayes, 1854) (Pleistocene–Recent, New
Zealand). The former carditamerine genera
Choniocardia Cossmann, 1905, Cossmanella Mayer-
Eymar, 1896, Cretocardia Conrad, 1877,
Fenestricardita Casey, 1961, Goossensia Cossmann,
1885, Izumicardia Ichikawa & Maeda, 1963 and
Miodontiscus Dall, 1903 were not considered because
the incompleteness or unavailability of their material.
All genera belonging to the subfamily
Miodomeridinae were included in the analysis:
Miodomeris Chavan, 1936 by M. cossmanni (Chavan,
1936) (type species of genus: late Eocene, France),
Pteromeris Conrad, 1862 by P. perplana (Conrad, 1841)
(type species of genus: Pliocene–Recent, North
America) and Coripia de Gregorio, 1885 by C. uniden-
tata (Basterot, 1825) (type species of genus:
Pliocene–Recent, Europe). The correct type species of
Coripia is C. unidentata as properly explained by Pras
(2013), although many works mention C. corbis
(Philippi, 1836) in this respect. The genus Chavanella
Jaworski, 1938 is a synonym of Miodomeris
(Vokes 1967).
Chavan (1969) established within the subfamily
Venericardiinae the Cenozoic genera Venericardia
Lamarck, 1801, Leuroactis Stewart, 1930, Pacificor
Verastegui, 1953, Venericor Stewart, 1930, Megacardita
Sacco, 1899 and the Mesozoic Trapezicardita Casey,
1961. With the exception of the latter (because material
of this species was not available), all of these genera
were included in the analysis. Leuroactis is represented
by L. pilsbry Stewart, 1930 (type species of the genus:
late Paleocene, North America), Pacificor by P. mulleri
Verastegui, 1953 (type species of the genus: late
Eocene, North America), Venericor by V. planicosta
(Lamarck, 1801) (type species of the genus: late
Eocene, France), V. aposmithii Gardner & Bowles, 1939
 Phylogeny of Carditidae
(late Paleocene, North America) and V. bashiplata
Gardner & Bowles, 1939 (early Eocene, North America)
and Megacardita by M. jouanetti (Basterot, 1825) (type
species of the genus, Miocene, Europe). Many species
were assigned to Venericardia in the bibliography but,
according to Huber (2010), this is a fossil taxon,
restricted to the Eocene of France. In this analysis this
genus is represented by the type species, V. imbricata
Lamarck, 1801 (late Eocene, France). The recently re-
validated genus Neovenericor Rossi de Garc
ıa, Levy &
Franchi, 1980 (P
erez et al. 2017b) was not formally
included within Venericardiinae but its proposed rela-
tionships with Venericor or Megacardita (Gardner &
Bowles 1939; Feruglio 1954; Camacho & Fern
andez
1956) allow it to be included in this subfamily.
Neovenericor is represented in the analysis by N. aus-
troplata (Gardner & Bowles, 1939) (type species of the
genus: early Miocene, Argentina), N. paranensis
(Borchert, 1901) (late Miocene, Argentina), N. carreren-
sis (Griffin, 1991) (Eocene, Argentina) and N. ponder-
osa (Suter, 1913) (late Oligocene, New Zealand).
Among the former Carditesinae, the genus Cardites
Link, 1807 is represented by three species: C. antiquatus
(Linnaeus, 1758) (type species of the genus:
Miocene–Recent, Europe), C. partschii (Goldfuss, 1840)
(Miocene, Europe) and C. feruglioi (Petersen, 1946)
(early Paleocene, Argentina), the genus Glyptoactis
Stewart, 1930 is represented by G. hadra (Dall, 1903)
(type species of the genus) (early–middle Miocene,
North America), the genus Baluchicardia Rutsch &
Schenk, 1943 is represented by B. beaumonti (d’Archiac
& Haime, 1854) (type species of the genus: Late
Cretaceous–Paleocene, India), the genus Claibornicardia
Stenzel & Krause, 1957 is represented by Cl. alticostata
(Conrad, 1833) (type species of the genus) (Eocene,
North America), Cl. acuticostata (Lamarck, 1805)
(Eocene, France) and Cl. paleopatagonica (Ihering,
1903) (early Paleocene, Argentina) and the genus
Paraglans Chavan, 1941 is represented by P. calcitra-
poides (Lamarck, 1806) (type species of the genus:
Eocene, France). Ludbrookia Chavan, 1951 is consid-
ered a synonym of Pseudocardia. The genus
Xenocardita Vokes, 1946 was not included due to the
lack of information about its hinge morphology.
The subfamily Thecaliinae is composed of two genera
according to Dall (1902, 1903) and Chavan (1969):
Thecalia Adams & Adams, 1858 and Milneria Dall,
1881. Recently, a new thecaliine genus Powellina
Huber, 2010 was described. The species T. concamerata
(Bruguiere, 1792) (type species of the genus: Recent,
South Africa) and P. brookesi (type species of the
genus: Finlay, 1926) (Recent, New Zealand) were con-
sidered in the phylogenetic analysis. The third taxa,
5
Milneria, shows a distinct morphology (its brood cham-
ber may not be homologous and its outline and external
sculpture are different to all known carditids) and Huber
(2010) proposed that its correct placement in this sub-
family was doubtful.
Other carditid genera mentioned in the literature were
never assigned to any of the subfamilies proposed by
Chavan, but they were included in this analysis.
Birkelundita Heinberg, 1993 was represented in the ana-
lysis by B. turoniense Heinberg, 1993 (type species of
the genus: Late Cretaceous, Europe). Also, the recently
described genus Kolmeris P
erez & del R
ıo, 2017b is
represented in the analysis by K. tehuelchana (Ihering,
1907) (type species of the genus: early Pliocene,
Argentina). Many former alticostates were considered,
including Fasciculicardia Maxwell, 1969 represented by
F. subintermedia (Suter, 1917) (type species of the
genus: early Miocene, New Zealand) and F. acanthodes
(Suter, 1917) (late Eocene, New Zealand) and F. sp.
(late Miocene, Argentina) (P
erez & del R
ıo 2017b),
Rotundicardia Heaslip, 1968 represented by R. rotunda
(Lea, 1833) (type species of the genus, late Eocene,
North America), R. diversidentata (Meyer, 1885) (late
Eocene, North America) and R. mariobrosorum P
erez &
del R
ıo, 2017a (early Paleocene, Argentina),
Purpurocardia Maxwell, 1969 by P. purpurata
(Deshayes, 1854) (type species of the genus:
Pliocene–Recent, New Zealand and Australia), P. ele-
gantoides (Ortmann, 1899) (Eocene, Argentina and
Chile) and P. leonensis del R
ıo, 1986 (late Miocene,
Argentina) and the recently described genera Kalelia
P
erez & del R
ıo, 2017a represented by K. multicostata
(Lamarck, 1806) (type species of the genus: late
Paleocene, France), K. pectuncularis (Lamarck, 1806)
(late Paleocene, France) and K. burmeisteri (B€ohm,
1903) (early Paleocene, Argentina), and Darwinicardia
P
erez & del R
ıo, 2017b represented by D. patagonica
(G. B. Sowerby I, 1846) (type species of the genus:
early Miocene, Argentina) and D. angusticostata
(Deshayes, 1824) (¼V. granulata Defrance, 1828; late
Eocene, France).
According to Gardner & Bowles (1939), Verastegui
(1953), Stewart (1930), McClure & Lockwood (2015)
and Camacho & Fern
andez (1956), the genera
Venericor, Leuroactis, Pacificor and Neovenericor rep-
resent planicostate carditids. Also, Stewart (1930) dis-
cussed the placement of Megacardita within
planicostates. Harris (1919), Stewart (1930), Stenzel &
Krause (1957), Heaslip (1968) and Maxwell (1969) con-
sidered within alticostate carditids the genera
Venericardia, Claibornicardia, Baluchicardia,
Glyptoactis, Rotundicardia, Purpurocardia and
6 D. E. P
erez
Fasciculicardia. P
erez et al. (2017a, b) also included
Kalelia and Darwinicardia in this group.
Studied material and repositories
A summary of specimens used in the phylogenetic anal-
yses is listed in the Supplementary material, Appendix
1. Studied material includes specimens from the follow-
ing repositories: MACN-Pi and CIRGEO-PI, Divisi
on
Paleoinvertebrados, Museo Argentino de Ciencias
Naturales ‘Bernardino Rivadavia’, Buenos Aires,
Argentina; MACN-In, Divisi
on Invertebrados, Museo
Argentino de Ciencias Naturales ‘Bernardino
Rivadavia’, Buenos Aires, Argentina; CPBA, C
atedra
de Paleontolog
ıa of the Universidad de Buenos Aires,
Buenos Aires, Argentina; MLP, Museo de La Plata,
Argentina; MHNSR.Pi, Museo de Historia Natural of
San Rafael, San Rafael, Argentina; SGN, Servicio
Geol
ogico Minero, Buenos Aires, Argentina; MAS-Pi,
Museo de Ciencias Naturales y Antropol
ogicas ‘Prof.
Antonio Serrano’, Paran
a, Argentina; DMT-Pi,
Colecci
on Paleoinvertebrados, Centro de Investigaciones
Cient
ıficas y de Transferencia Tecnol
ogica a la
Producci
on, Diamante, Argentina; SGO.PI, Colecci
on
Paleoinvertebrados, Museo de Historia Natural,
Santiago, Chile; PZ-NRM.Mo, Samling Paleobiologi,
Naturhistoriska Riksmuseet, Stockholm, Sweden;
NHMUK, The Natural History Museum, London, UK;
MNHN, Mus
eum national d’Histoire naturelle, Paris,
France; GMNH, Mus
eum d’Histoire Naturelle de
Geneve, Switzerland; NMR, Natural History Museum
of Rotterdam, the Netherlands; USNM.Mo, Mollusk
Collecion, National Museum of Natural History,
Smithsonian Institution, Washington DC, USA; PRI;
Palaeontological Research Institution, Ithaca, NY, USA;
ANSP-IP, Paleoinvertebrates Collection, Academy of
Natural Sciences of Drexel University, Philadelphia, PA,
USA; UF, University of Florida, Gainesville, FL, USA;
YPM, Yale Peabody Museum, New Haven, CT, USA;
TM, Museum of New Zealand Te Papa Tongarewa,
Wellington, New Zealand; AMS, Australian Museum,
Sydney, Australia.
Character sampling and scoring
The character list is composed of 144 morphological
characters, including both external and internal shell
morphology, which are listed in Supplementary material,
Appendix 2. The scorings are included in
Supplementary material, Appendices 3A and 3B.
Twenty-one characters sample the general shell outline,
seven the umbon, four the escutcheon, nine the lunule,
three the general morphology of the hinge, 27 the right
hinge features, 16 the left hinge features, three the
general morphology of the external sculpture, five the
comarginal external sculpture, 23 the radial external
sculpture, 10 the comarginal nodes, eight the internal
morphology (including muscle scars and the pallial
line), and seven the ventral margin crenulations. Two
characters refer to sexually dimorphic features and 14 to
ontogenetic variations. Hinge features, traditionally con-
sidered the main systematic feature for many bivalve
groups, including carditids (Lamy 1922; Chavan 1969),
correspond to 31.9% of all the characters, while external
sculpture, an important feature in carditid systematics
for many authors (Harris 1919; Verastegui 1953;
Maxwell 1969), comprises 28.47%. Characters 1 (num-
ber of radial ribs) and 2 (mean size of adult shells) were
considered as continuous in some search strategies and
as discrete characters in others. They were discretized
using raw data and performing a frequency analysis in
PAST 3.14 (Hammer et al. 2001) to obtain discretized
states. The following 13 characters were treated as addi-
tive (¼ordered): 1, 2, 5, 6, 23, 24, 29, 57, 62, 70, 75,
105 and 135. Characters 1 and 2 were considered as
additive in the discrete matrix approach and the rest of
them represent progressive and nested changes of
a feature.
The characters were scored in Mesquite v3.2
(Maddison & Maddison 2017). Character-states were
scored as polymorphic if different states were present in
multiple individuals of a single taxon, using the follow-
ing notation: [state A state B]. Unknown characters
were scored as missing data using a quotation mark (?).
The latter occurred with the unknown right hinge of
Fasciculicardia sp. and the partially unknown left hinge
of C. mesembria. If tripartite radial ribs were not devel-
oped or the lunule was absent, subsidiary characters
were considered as inapplicable and scored using a
hyphen (–). Only 0.722% of the character-states were
scored as missing data or inapplicable. All characters
were scored based on adult well-preserved shells,
including ontogenetically variable characters, because
the bivalve shell retains a complete record of external
traits through all post-larval growth stages (Crampton &
Maxwell 2000).
Character definitions
General features of the carditid shell are summarized in
Figure 1. External morphologies of different carditids
are shown in Figure 2 and internal and hinge morpholo-
gies in Figure 3. Details of different features are illus-
trated in Figure 4. Morphological terminology follows
Verastegui (1953) and Heaslip (1968), which are con-
sistent with the bivalve morphological terms of Cox
(1969) and Bernard (1895) for carditids. The different
features were assembled in the following character-
 Phylogeny of Carditidae
Figure 1. General features of the carditid shell. Claibornicardia paleopatagonica (Ihering, 1903) (MACN-Pi 5197). A, external view
of right valve; B, left hinge; C, internal view of right valve. Scale bar equals 10 mm.
groups: general shell outline and shell shape (characters
2–22), umbones (characters 23–29), escutcheon (charac-
ters 30–33), lunule (characters 34–42), hinge (characters
43–88), sculpture generalities (characters 89–91), comar-
ginal sculpture (characters 1, 97–119), comarginal nodes
(characters 119–128), internal morphology (characters
130–137) and ventral margin crenulations (characters
138–144). Characters that describe the number of radial
ribs remains virtually fixed throughout the lifespan of
an individual, and this number shows little intraspecific
variation. To test the non-independence of the number
of ribs with respect of the sampled species, an analysis
using a generalized linear model (GLM) was carried out
in R (R Development Core Team 2017). This analysis is
described in Supplementary material, Appendix 4.
Tree searches and branch support
The two data matrices, matrix 1 with all discrete charac-
ters (‘all disc’ approach) and matrix 2 with characters 1
and 2 as continuous characters (‘cont þ disc’ approach),
were analysed using TNT v1.5 (Goloboff & Catalano
2016). Due to the number of taxa included in this phylo-
genetic analysis, heuristic searches were conducted.
They involved the generation of an initial tree via ran-
dom addition sequence that was subjected subsequently
to TBR branch-swapping, retaining a maximum of 10
trees per replicate. This procedure was replicated 100
times. Search strategies include two weighting arrange-
ments: equal weights and implied weighting (Goloboff
1993) using k values from 1 and 100 by increments of
1. A strict consensus tree was generated only for the
results of the equal weighting searches because a single
tree was found for each search with implied weights.
Clades recovered in different search strategies were
illustrated through sensitive grids (Wheeler 1995).
7
As a measure of branch support, jackknife indices
(jk) (Felsenstein 1985) were calculated. For jackknife,
difference of frequencies (GC) and p ¼ 0.14 were used,
performing 1000 pseudoreplicates (Goloboff et al.
2003). Consistency (CI) and retention (RI) indices were
also calculated (Farris 1989). Synapomorphies common
to all the recovered trees for each strategy are listed in
the Supplementary material, Appendix 5.
Posterior analysis of tree results
Posterior to the searches, the most representative top-
ology obtained from the all-discrete matrix (implied
weighting, k ¼ 31: see Results, below) was used in stat-
istical analyses using R (R Core Development Team
2017). For reading the matrix and tree topology the
packages ‘ape’ (Paradis et al. 2004; Popescu et al.
2012), ‘phylobase’ (Bolker et al. 2016), ‘paleotree’
(Bapst 2012), ‘strap’ (Bell & Lloyd 2014) and ‘Claddis’
(Lloyd 2016) were used. The phylogenetic tree was
time-calibrated (functions: ‘timePaleoPhy’ and
‘geoscalePhylo’) using the minimum branch length
(mbl) method to maintain the temporal structure of
events (Laurin 2004). The stratigraphic ranges of the
taxa used in this analysis are included in Supplementary
material, Appendix 6. A morphological disparity ana-
lysis was performed (Lloyd 2016), in which a distance
matrix was calculated using generalized Euclidean dis-
tances (GED), which allows for missing data (function:
‘MorphDistMatrix’), and these data were used to per-
form a principal coordinate analysis (PCoA). With these
results, a phylomorphospace was generated (function:
‘MorphspacePlot’).
Phylogenetic signals for all the characters were calcu-
lated using the package ‘picante’ (function:
‘phylosignal’) (Kembel et al. 2010), considering all
8 D. E. P
erez
 Phylogeny of Carditidae 9

characters as unordered and using the most representa- Kolmeris (Clade 5) is present under equal weighting and
tive topology (implied weighting, k ¼ 31: see Results, k ¼ 8–100 and another formed by Coripia, Vimentum
below). Phylogenetic signal is a measure of the degree and Scalaricardita (Clade 4) in all of the searches. The
to which phylogeny predicts the ecological similarity of genera Cyclocardia (Clade 10) and Cardites (Clade 7)
species. By means of the K statistic this measure com- are monophyletic in all of the searches, but
pares the observed signal in a trait with the signal under Scalaricardita is non-monophyletic under equal weight-
a Brownian motion model of trait evolution on a phyl- ing and k ¼ 23–74. The genus Pleuromeris (Clade 6) is
ogeny (Blomberg et al. 2003). K values greater than 1 polyphyletic under k ¼ 23–74 and monophyletic in the
indicate the presence of phylogenetic signal. Results remaining searches. A clade formed by Neovenericor,
were grouped and analysed in the character-groups Megacardita, Bathycardita, Venericor, Leuroactis and
defined previously. Pacificor (Clade 8), i.e. all planicostate taxa, is recov-
ered in all of the searches and is placed within the
remaining carditids under equal weighting and
Results k ¼ 23–100 (Clade 13). In other searches, these taxa are
placed with Cardites, the Centrocardita clade þ Cardita
Matrix with all discrete characters clade or separated from the rest of the carditids
Tree searches with this approach are summarized in (k ¼ 1–22). Among planicostates, two lineages are
Figure 5A. Under equal weights, 50 most parsimonious recovered. The first lineage is composed of Leuroactis,
trees (MPTs) were obtained, with tree lengths of 834 Pacificor and Venericor species (Clade 14) in all of the
steps, CI of 0.333 and RI of 0.796. Clades recovered in searches, and the second one is composed of
equal weighting and implied weighting are included in Megacardita, Bathycardita and Neovenericor (Clade 15)
the sensitive grids shown in the tree with k ¼ 23 under equal weighting and k ¼ 75–100. A clade formed
(fit ¼ 18.76211, CI ¼ 0.333, RI ¼ 0.795). Jackknife sup- by the remaining carditids, i.e. Venericardia,
ports are also shown for this implied weighting tree. Arcturellina, Paraglans, Purpurocardia, Darwinicardia,
All of these trees (Fig. 5A) show a clear distinction Kalelia, Rotundicardia, Claibornicardia, Baluchicardia,
between Palaeocardita spp. þ Schizocardita (Clade 1) Cardiocardita, Glyptoactis and Fasciculicardia (Clade
and the remaining carditids. Former Carditinae (Clade 9), is recovered in k ¼ 1–75. The genus Cardites is
3) and Birkelundita are placed as the sister clade of nested within Clade 9 or it appears as the sister clade of
Thecaliinae (Clade 11) in most searches, and in others this group under equal weighting and k ¼ 1–7, 23–100.
the Thecaliinae members are nested within Carditinae. An internal clade formed by Kalelia, Paraglans,
This group is the sister clade of Arcturellina, Cardiocardita and Venericardia (Clade 16)
Carditamera þ Glans þ Centrocardita (Clade 2). The and another including Claibornicardia, Baluchicardia,
species of the genera Cyclocardia, Pleuromeris, Rotundicardia, Glyptoactis and Fasciculicardia (Clade
Scalaricardita, Vimentum, Coripia, Pteromeris, 17) are obtained in some searches (both under
Miodomeris and Kolmeris (Clade 12) are grouped under k ¼ 5–74). These groups agree with the traditional sub-
equal weighting and k ¼ 14–100 searches. Within Clade families Palaeocarditinae and Thecaliinae, but
12, a clade formed of Pteromeris, Miodomeris and Miodomeridiinae, Carditinae, Carditesinae,
3
Figure 2. External morphologies among carditid species. A, Crassatella ponderosa (Gmelin, 1791) (GMNH 45970), left valve; B,
Laevastarte basteroti (de la Jonkaire, 1823) (MACN-Pi 3011), left valve; C, Palaeocardita crenata (M€unster, 1841) (MACN-Pi
3007), right valve; D, Beguina semiorbiculata (Linnaeus, 1758) (MACN-In 17486), left valve; E, Cardita crassa Lamarck, 1819
(MACN-Pi 2353), right valve; F, Carditamera floridana (Conrad, 1838) (MACN-In 41173), left valve; G, Thecalia concamerata
(Bruguiere, 1792) (MACN-In 13778), right valve; H, Carditamera gracilis (Shuttleworth, 1856) (MACN-In 2623), left valve; I,
Carditamera arata (Conrad, 1838) (YPM IP 560023), right valve; J, Cyclocardia dalek P
erez & del R
ıo, 2017b (MACN-Pi 5770),
left valve; K, Cyclocardia compressa (Reeve, 1843) (MACN-Pi 363), left valve; L, Cyclocardia borealis (Conrad, 1831) (YPM IZ
033525), left valve; M, Miodomeris cossmanni (Chavan, 1936) (MNHN FJ09318), right valve; N, Pleuromeris tridentata (Say, 1826)
(UF 189442), left valve; O, Pteromeris perplana (Conrad, 1841) (UF 49642), left valve; P, Darwinicardia patagonica
(Sowerby,1846) (CPBA 15948), right valve; Q, Purpurocardia purpurata (Deshayes, 1854) (MACN-In 28255), right valve; R,
Megacardita jouannetti (Basterot, 1825) (MACN-Pi 2355), right valve; S, Scalaricardita camaronesia (Ihering, 1907) (MACN-Pi
349), right valve; T, Coripia unidentata (Basterot, 1825) (FJ14611), right valve; U, Venericor planicosta (Lamarck, 1801) (MACN-Pi
961), right valve; V, Kolmeris tehuelchana (Ihering, 1907) (MACN-Pi 371), left valve; W, Neovenericor austroplata (Gardner &
Bowles, 1939) (USNM.Mo 327924), right valve; X, Venericardia imbricata Lamarck, 1801 (GMNH 46023), right valve; Y, Kalelia
multicostata (Lamarck, 1806) (MNHN A07711), right valve; Z, Cardites antiquatus (Linnaeus, 1758) (MACN-In 1163), right valve;
A’, Rotundicardia rotunda (Lea, 1833) (ANSP-IP 5267), left valve; B’, Rotundicardia mariobrosorum P
erez & del R
ıo, 2017a
(CPBA 17276), left valve. Scale bar equals 10 mm.
10 D. E. P
erez

Figure 3. Internal and hinge morphologies among Carditidae species. A, Crassatella ponderosa (Gmelin, 1791) (GMNH 45970), left
hinge; B, Crassatella ponderosa (Gmelin, 1791) (GMNH 45970), right valve; C, Laevastarte basteroti (de la Jonkaire, 1823)
(MACN-Pi 3011), left hinge; D, Laevastarte basteroti (de la Jonkaire, 1823) (MACN-Pi 3011), right valve; E, Beguina
semiorbiculata (Linnaeus, 1758) (MACN-In 17486), right hinge; F, Beguina semiorbiculata (Linnaeus, 1758) (MACN-In 17486), left
valve; G, Cardita crassa Lamarck, 1819 (MACN-Pi 2353), left hinge; H, Cardita crassa Lamarck, 1819 (MACN-Pi 2353), right
valve; I, Carditamera floridana (Conrad, 1838) (MACN-In 41173), left hinge; J, Carditamera floridana (Conrad, 1838) (MACN-In
41173), right valve; K, Carditamera gracilis (Shuttleworth, 1856) (MACN-In 2623), right hinge; L, Carditamera gracilis
(Shuttleworth, 1856) (MACN-In 2623), left valve; M, Carditamera arata (Conrad, 1838) (YPM IP 560023), right valve; N, Kalelia
multicostata (Lamarck, 1806) (MNHN A07711), right valve; O, Rotundicardia rotunda (Lea, 1833) (ANSP-IP 5267), left valve; P,
Venericardia imbricata Lamarck, 1801 (GMNH 46023), left hinge. Q, Venericardia imbricata Lamarck, 1801 (GMNH 46023), right
valve; R, Venericor planicosta (Lamarck, 1801) (MACN-Pi 961), right hinge; S, Venericor planicosta (Lamarck, 1801) (MACN-Pi
961), left hinge; T, Megacardita jouannetti (Basterot, 1825) (MACN-Pi 2350), left hinge; U, Neovenericor paranensis (Borchert,
1901) (PZ-NRM Mo. 118089), right hinge; V, Cardites antiquatus (Linnaeus, 1758) (MACN-In 1163), right valve; W, Scalaricardita
camaronesia (Ihering, 1907) (MACN-Pi 349), right valve; X, Cyclocardia borealis (Conrad, 1831) (YPM IZ 033525), right valve; Y,
Miodomeris cossmanni (Chavan, 1936) (MNHN FJ09318), left valve; Z, Pteromeris perplana (Conrad, 1841) (UF 49642), left valve.
The same scale bar applies to all parts, but represents different lengths: 10 mm for A–H, K–N, P–V, X; 5 mm for I, J, O; 3 mm for
W; 2 mm for Y, Z.
 Phylogeny of Carditidae 11

Figure 4. Details of different features among Carditidae species. A, Crassatella ponderosa (Gmelin, 1791) (GMNH 45970), dorsal
view of left valve showing escutcheon; B, Palaeocardita crenata (M€unster, 1841) (MACN-Pi 3007), dorsal view of valve showing
escutcheon; C, Purpurocardia purpurata (Deshayes, 1854) (MACN-In 28255), dorsal view of valve showing absence of escutcheon;
D, Thecalia concamerata (Bruguiere, 1792) (MACN-In 13778), internal view of right valve showing incubatory chamber; E,
Neovenericor paranensis (Borchert, 1901) (PZ-NRM Mo. 118089), dorsal view of right hinge showing finely striations on surface of
middle tooth; F, Kolmeris tehuelchana (Ihering, 1907) (MACN-Pi 371), stacked radial ribs on posterior flank of left valve; G,
Glyptoactis hadra (Stewart, 1930) (UF 116940), left hinge showing little knob below ventral extreme of lunular margin; H,
Laevastarte basteroti (de la Jonkaire, 1823) (MACN-Pi 3011), comarginal undulations; I, Venericor planicosta (Lamarck, 1801)
(MACN-Pi 961), obsolete radial ribs towards ventral margin; J, Carditamera gracilis (Shuttleworth, 1856) (MACN-In 2623), radial
ribs with posteriorly inclined cross section; K, Venericor planicosta (Lamarck, 1801) (MACN-Pi 961), entire radial ribs; L,
Neovenericor austroplata (Gardner & Bowles, 1939) (USNM.Mo 327924), entire radial ribs; M, Darwinicardia patagonica
(Sowerby,1846) (PRI.66405), tripartite radial ribs with circular nodes; N, Claibornicardia paleopatagonica (Ihering, 1903) (MACN-Pi
5197), tripartite radial ribs; O, Rotundicardia mariobrosorum P
erez & del R
ıo, 2017a (MACN-Pi 5762), funginate nodes; P,
Fasciculicardia sp. (MACN-Pi 361b), saw-like nodes; Q, Scalaricardita camaronesia (Ihering, 1907) (MACN-Pi 349), rectangular
nodes; R, Darwinicardia angusticostata (Deshayes, 1824) (MNHN A25071), extreme tips of radial ribs poking out among
crenulations. The same scale bar applies to all parts, but represents different lengths: 10 mm for A, L; 6 mm for B, C, E, I, K; 5 mm
for G, J, M, P; 4 mm for N; 3 mm for D, F, O, Q, R; 2.5 mm for H.
12 D. E. P
erez

Figure 5. Results of the phylogenetic analyses. A, tree resulting from all discrete matrix approach (k ¼ 23); B, tree resulting from
continuous þ discrete matrix approach (k ¼ 14). Sensitive grids include presence or absence of clades in EW (equal weighting) and
IW (implied weighting with k between 1 and 100) searches. Clades explained in Results section of text. Jackknife support values
indicated below branches or grids. Only values above 50 are reported.
 Phylogeny of Carditidae
Venericardiinae and Carditamerinae are not recovered,
contra Chavan (1969).
The monophyly of Carditidae is strongly supported
(jk ¼ 100). Palaeocarditinae has a high support value
(jk ¼ 88) and the separation between this and the
remaining carditids is strong (jk ¼ 100). Clade 2
(jk ¼ 99), Clade 4 (jk ¼ 96), Clade 5 (jk ¼ 82), Clade 7
(jk ¼ 100), Clade 8 (jk ¼ 99), Clade 14 (jk ¼ 100) and
Thecaliinae (jk ¼ 88) also show high support values.
The genera Kalelia, Rotundicardia, Fasciculicardia,
Purpurocardia, Carditamera and Palaeocardita have
support values above jk ¼ 80.
Matrix with continuous and discrete characters
Tree searches found using this approach are summarized
in Figure 5B. Three MPTs with tree lengths of
1324.348 steps, CI of 0.283 and RI of 0.744 were found
under equal weights. The clades recovered under equal
weighting and implied weighting are included in the
sensitive grids shown in the tree recovered under k ¼ 14
(fit ¼ 26.1667, CI ¼ 0.263, RI ¼ 0.717). The jackknife
supports are also indicated on this tree.
Clades recovered with this approach are similar to
those found using all discrete characters (Fig. 5B).
Palaeocarditinae (Clade 1) is separated from the remain-
ing carditids, and Thecaliinae (Clade 11) is recovered
nested within Carditinae (Clade 3) in k ¼ 30–100
searches. Clade 2 (Carditamera þ Glans þ
Centrocardita) and Clade 4 (Scalaricardita þ Coripia
þ Vimentum) are present in all searches. The genus
Cyclocardia (Clade 10) is monophyletic under implied
weighting searches, but under equal weighting is para-
phyletic. A group composed of Kolmeris, Miodomeris
and Pteromeris (Clade 5) is recovered under equal
weighting and k ¼ 14–100 searches, and the genus
Pleuromeris is monophyletic only under equal weighting
and k ¼ 1–30 searches. Clade 12, composed of Clades 4,
5, 6 and 10, is recovered under equal weighting and
k ¼ 30–100 searches. The genus Cardites is monophy-
letic in all of the searches and nested within Clade 9 in
k ¼ 14–100 searches. The planicostate clade (Clade 8)
and the remaining carditids (Clade 9) are recovered in
all of the searches. These two clades are sister taxa to
each other under equal weighting and k ¼ 14–100
searches. A lineage composed of Leuroactis, Pacificor
and Venericor (Clade 14) is recovered in all of the
searches, and other including Megacardita, Bathycardita
and Neovenericor only in the k ¼ 1–30 searches. In
Clade 9, two major internal lineages are recovered.
Clade 16 (Paraglans, Arcturellina, Cardiocardita,
Venericardia and Kalelia) is recovered in all of the
searches and Clade 17 (Claibornicardia, Rotundicardia,
Baluchicardia, Fasciculicardia and Glyptoactis) only
13
under implied weighting. As with the previous approach,
only the recovery of Palaeocarditinae and Thecaliinae
are congruent with the earlier hypothesis of Chavan
(1969) sensu stricto.
The monophyly of Carditidae is strongly supported in
the cont þ disc approach (jk ¼ 100). Palaeocarditinae
(jk ¼ 93) and Thecaliinae (jk ¼ 94) have high support
values. Clades 2 (jk ¼ 99), 4 (jk ¼ 96), 7 (jk ¼ 100), 8
(jk ¼ 98) and 14 (jk ¼ 100) show high support values.
Clade 5 (jk ¼ 62) and the genus Cyclocardia (Clade 10,
jk ¼ 57) have lower support values than the others. As
in the previous approach, the genera Kalelia,
Rotundicardia, Darwinicardia, Purpurocardia,
Carditamera and Palaeocardita have support values
above jk ¼ 80. Fasciculicardia shows a slightly lower
support value (jk ¼ 78) and Scalaricardita is more
poorly supported (jk ¼ 65).
Synapomorphies
A detailed summary of the synapomorphies of each
clade is provided in Supplementary material,
Appendix 5, accompanied by the list of synapomorphies
obtained from the two search approaches and described
for the implied weighting trees of k ¼ 23 and k ¼ 14,
respectively.
Time-calibrated tree
The time-calibrated tree (Fig. 6) shows a long temporal
separation between two clades: Clade 1
(Palaeocarditinae), which was restricted to the Late
Triassic, and the remaining carditids, which are distrib-
uted from the Late Cretaceous to the Recent. This separ-
ation indicates a long ghost lineage of about 100 million
years. Species of the genera Pleuromeris and
Cyclocardia, and members of Clades 4 and 5, are
restricted to the Eocene–Recent and Clade 2 is distrib-
uted from the Neogene–Recent. Clade 3 and Clade 11
have a late Neogene–Recent distribution, with the
exception of Birkelundita turoniense, a Late Cretaceous
species. Planicostate taxa (Clade 8) are restricted to the
Paleocene–Miocene and the remaining carditids occur in
Maastrichtian–upper Miocene units, with the exception
of the extant species Cardiocardita ajar and
Purpurocardia purpurata.
Morphological disparity analysis
The PCoA recovered a first coordinate that accounts for
19.22% of the total variance and the second coordinate
accounts for 13.7% of the variance. The phylomorpho-
space obtained from this analysis (Fig. 7) shows three
discrete groups. The first one is composed of
14
Figure 6. Time-calibrated tree. Ages in millions of years. Bars indicate the stratigraphical range of each taxon. Q ¼ Quaternary.
palaeocarditines (Clade 1) and outgroups and is sepa-
rated from the other two groups along the first coordin-
ate. The second group is composed of Clade 4 and
Clade 5 and the genera Pleuromeris and Cyclocardia
(the latter forming Clade 12) and the third group
includes the remaining carditids. The second and third
groups share a distribution along the first coordinate but
are separated along the second one. A fourth group is
slightly separated from the third group and includes spe-
cies of Centrocardita, Glans and Carditamera
(Clade 2).
Phylogenetic signal of the characters
The analysis of phylogenetic signal was performed on
the all disc matrix and its results are showed in
Figure 8. The general shell outline and shell shape char-
acter groups exhibit low values of phylogenetic signal
(only the development and elongation of the dorsal mar-
gin have a significant signal). The relative position of
the umbones is the umbonal character with the highest
value. All of the escutcheon characters have high values
D. E. P
erez
of phylogenetic signal. In contrast, the lunule characters
have low values. Regarding the hinge, general characters
do not have high values of phylogenetic signal, with the
exception of the presence of an internal resilifer. The
right hinge shows high values, including the presence of
laminar lateral teeth, the presence and development of
an anterior tooth, the development of middle and poster-
ior teeth and the extension of a ventral edge of the
hinge. The left hinge presents low values, with the
exception of some features of the anterior tooth and the
presence of laminar lateral teeth. The types of sculpture
on the valves have high values, as occurs with the
comarginal sculpture. Regarding the radial sculpture, the
most conspicuous ornamentation among carditids, a few
characters show high values (type of radial ribs, pres-
ence of non-reduced interspaces, presence of postero-
ventral prosocline radial ribs). Comarginal nodes present
very high values of phylogenetic signal (for example in
their presence, type, size and separation). The internal
morphology includes some characters with a high value
(prominence, size and placement of the adductor muscle
scars, shape of the pallial line, presence of inner
 Phylogeny of Carditidae 15

Figure 7. Phylomorphospace resulting from the morphological disparity analysis. Red ¼ First group (Clade 1 in the text).
Yellow ¼ Second group (Clades 4, 5 and 12 in the text). Light blue ¼ Third group. Dark blue ¼ Fourth group (Clade 2 in the text).
B ¼ Beguina, Ba ¼ Baluchicardia, Bi ¼ Birkelundita, C ¼ Cardita, Ca ¼ Cardites, Cl ¼ Claibornicardia, Cy ¼ Cyclocardia,
D ¼ Darwinicardia, F ¼ Fasciculicardia, K ¼ Kalelia, L ¼ Leuroactis, N ¼ Neovenericor, P ¼ Purpurocardia, Pa ¼ Pacificor,
Pl ¼ Pleuromeris, Po ¼ Powellina, Pt ¼ Pteromeris, R ¼ Rotundicardia, S ¼ Scalaricardita, T ¼ Thecalia, Ve ¼ Venericor.

crenulations). When values are considered for the
groups, the general characters of the shell outline, escut-
cheon, lunule, hinge generalities, right hinge, left hinge,
radial sculpture and comarginal nodes have significant
values, with the right hinge and commarginal having the
highest values.
Discussion
Previous subfamiliar scheme
The subfamiliar scheme proposed by Chavan (1969) and
followed by subsequent authors (e.g. Bernard 1983;
Bouchet et al. 2010; Huber 2010; Carter et al. 2011; La
Perna et al. 2017) is not fully recovered by the phylo-
genetic analysis (Fig. 9). Some of the genera or species
included in the analyses were unknown at the time of
the analysis of Chavan (1969) and they are listed in
Table 1 and Figure 9.
The subfamily Palaeocarditinae sensu Chavan (1969)
is recovered in all the analyses with high support values.
Chavan (1969) indicates an elongate shell-outline, ortho-
gyrate umbones and a particular morphology of the right
middle tooth as key features of this group. This subfam-
ily is consistent with Clade 1 of this phylogenetic ana-
lysis, but its synapomorphies (shell-outline and umbonal
characters) do not coincide with those proposed by
Chavan (1969). The taxa of this subfamily are all pre-
Cretaceous in age according to Chavan (1969) and in
this phylogeny is restricted to Triassic species.
The Carditinae (included in Clade 3) is recovered in
most searches, but is paraphyletic in a few of them by
including the subfamily Thecaliinae (Clade 11) nested
within it. These two subfamilies have very similar myti-
liform or trapezoidal shell-outlines according to Chavan
(1969). The close relationship between these subfamilies
is highly supported in all of the analyses and has been
previously reported by Gonzalez & Giribet (2015). The
main difference between them is the presence of an
incubatory chamber in the Thecaliinae females, a feature
mentioned by Dall (1903) and Chavan (1969).
Carditamerinae is the most diverse of Chavan’s
(1969) subfamilies, including more than 20 genera, but
this is not recovered in the phylogenetic analyses per-
formed herein. A monophyletic Carditamerinae must
include all carditids except Palaeocarditinae sensu
16
Figure 8. Results of the phylogenetic signal analysis. Vertical axis indicates values of signal. Values above one (1) are significant
(dashed line marked this limit). Numbers of characters are described in Supplementary material, Appendix 2. Colour of points
separates the different character-groups.
Chavan (1969). Huber (2010) suggests that this subfam-
ily is not monophyletic. The type genus of the subfam-
ily (Carditamera) is closely related to Glans and
Centrocardita and these three taxa were recognized as
members of Carditamerinae by Chavan (1969). This
group is recognized as Clade 2 herein. The other former
Carditamerinae are included in the other clades recov-
ered here. Chavan (1969) proposed the presence of
shells with trigonal or trapezoidal outlines, a triangular
right middle tooth triangular and a conspicuous lateral
tooth as characters of this subfamily. These characters
are present in most of the taxa in Clade 2.
The subfamily Miodomeridinae according to Chavan
(1969) includes three genera: Coripia, Pteromeris and
Miodomeris. The latter two are recovered together and
close to Kolmeris in the analysis (Clade 5), but Coripia
is grouped with the former Carditamerinae taxa
Scalaricardita and Vimentum in Clade 4. Despite this,
Huber (2010) proposed that Coripia and Miodomeris
were closely related taxa, excluding Pteromeris. Chavan
(1969) proposed the following characters as synapomor-
phies of Miodomeridinae: small-sized shells, astartiform
outline, large hinge without a right anterior tooth and
with an inclined right middle tooth. Clade 5 synapomor-
phies include some outline characters corresponding to
the ‘astartiform’ outline mentioned by Chavan (1969),
such as obtuse-triangular outline, slightly inclined pos-
terior margin and curved and descendent dorsal margin.
The Venericardiinae of Chavan (1969) comprises
Venericardia and the former planicostate taxa
D. E. P
erez
Leuroactis, Pacificor, Venericor and Megacardita. The
type genus of the subfamily is recovered as distantly
related to these other genera. The Venericardiinae sub-
family shares prosogyrous umbones, sunken lunule,
right anterior tooth laminar, curved hinge teeth and an
obsolete lateral tooth according to Chavan (1969), but
these characters are not recovered as synapomorphies of
any clade herein.
Chavan (1969) included in Carditesinae the genera
Cardites, Baluchicardia, Claibornicardia, Glyptoactis
and Paraglans, but the type genus of the subfamily is
recovered as distantly related to these other genera. The
genus Venericardia is placed among these latter taxa. If
a monophyletic definition of Carditesinae is proposed,
this subfamily partially matches with Venericardiinae.
Chavan (1969) recognized as key characters of this sub-
family the presence of rounded and bulky shells, inter-
spaces with a ‘V’-shaped section, a very convex lunule
and curved right anterior and middle teeth. Clade 7
(Cardites spp.) has a very convex umbonal region as
synapomorphy.
According to the phylogenetic results presented here,
only the subfamily Palaeocarditinae is recovered sensu
Chavan (1969). Thecaliinae is also recovered but
included in Carditinae in some searches. The subfami-
lies Carditamerinae, Miodomeridinae and Carditinae
need re-definition. Carditesinae and Venericardiinae
must be synonymous if their previous definitions are
maintained. The subfamiliar scheme of Chavan (1969)
does not fully correspond to the monophyletic groups
 Phylogeny of Carditidae 17

Figure 9. Comparison between the previous systematic scheme of Chavan (1969) and the new scheme proposed here. Asterisks
indicate carditid taxa unknown to Chavan (1969).
18 D. E. P
erez
recovered here and lacks utility for understanding the
phylogeny and evolutionary history of the Carditidae.
The proposal of Scarlato & Starobogatov (1979) for
Carditamerinae, Venericardiinae, Carditesinae and
Palaeocarditinae is not recovered either, with the excep-
tion of the relationship between Venericardiinae and
Carditesinae.
The Planicostate and Alticostate lineages
The proposal of two lineages among carditids predates
the Chavan (1969) subfamiliar arrangement (Conrad
1830; Harris 1919; Stewart 1930), but it is was not
included in Chavan’s (1969) scheme. Some subsequent
authors followed this approach (Gardner & Bowles
1939; Heaslip 1968; McClure & Lockwood 2015).
These authors considered the species of both lineages
to belong to Venericardia and Chavan (1969) included
Venericardia sensu stricto and the planicostate genera
in his subfamily Venericardiinae (Fig. 9). Heaslip
(1968) and McClure & Lockwood (2015) applied the
term ‘venericards’ to both lineages and considered them
as Venericardia relatives. It is probable that Chavan
(1969) did not know the work of Heaslip (1968)
regarding alticostate carditids (the genus Rotundicardia
proposed in that work is not included in the list of
Chavan 1969).
According to Gardner & Bowles (1939), the planicos-
tate lineage includes three genera: Venericor, Leuroactis
and Pacificor. Others authors propose a similar scheme
but add a fourth taxon, Megacardita (Stewart 1930;
Verastegui 1953). The South American taxon
Neovenericor was considered as closely related to plani-
costate taxa (Gardner & Bowles 1939; Feruglio 1954;
Camacho & Fern
andez 1956; P
erez et al. 2017a). All of
these taxa plus Bathycardita are grouped in a monophy-
letic group in the analyses presented here (Clade 8).
Gardner & Bowles (1939) and McClure & Lockwood
(2015) also considered the planicostate group as mono-
phyletic, but they are polyphyletic in Heaslip (1963). A
preliminary diagnosis of this lineage (Harris 1919)
included the presence of large shells sculptured with
smooth radial ribs, inflated umbones, thick shells, right
anterior tooth laminar and large muscle scars (Gardner
& Bowles 1939). All of these characters are present in
Clade 8 taxa. Other characters mentioned by Gardner &
Bowles (1939) are present in North American planicos-
tates (grouped in Clade 14), such as a trigonal outline,
anteriorly placed umbones, large lunule, hinge plate
high, scimitar-shaped right middle tooth, and very low
radial ribs. This lineage is distributed from the late
Paleocene–Recent but Bathycardita raouli is the only
extant species. All of the other planicostates became
extinct before the Pliocene.
The alticostate lineage was originally defined to
include small venericards from Europe and North
America (Harris 1919). First, Stewart (1930) included in
this group the genus Glyptoactis. Subsequently, Rutsch &
Schenck (1941) included Baluchicardia and Stenzel &
Krause (1957) added the genus Claibornicardia. Heaslip
(1968) considered Rotundicardia as a member of this lin-
eage. Later, Maxwell (1969) included within alticostates
the New Zealand genera Purpurocardia and
Fasciculicardia. In the phylogenetic analyses herein, all
of these taxa are part of Clade 9, with the addition of
Paraglans, Arcturellina, Cardiocardita, Kalelia,
Darwinicardia and Cardites (in some searches). Huber
(2010) proposed that Cardiocardita is related to
Venericardia and McClure (2009) recovered Arcturellina
and Paraglans among alticostates. Some authors (Rutsch
& Schenck 1941; Heaslip 1968) considered a monophy-
letic origin for this group. In contrast, McClure &
Lockwood (2015) considered alticostates as a paraphy-
letic group, but they did not include Baluchicardia within
the alticostate lineages and used it as outgroup in their
phylogenetic analysis. The paraphyly of the alticostates in
their analysis could be a consequence of this approach.
Harris (1919) mentioned that alticostates can be defined
by the presence of small shells covered with fine sculp-
tured and tripartite radial ribs. The main synapomorphy
of Clade 9 is the presence of paracostal ribs, forming tri-
partite radial ribs. However, some of these taxa do not
possess paracostal ribs (Cardites, Purpurocardia and
some Rotundicardia species). In some searches, Cardites
is found as basal to the remaining alticostates, and could
represent the plesiomorphic condition for the group.
Heaslip (1968) noted the absence of paracostal ribs in
derived members of Rotundicardia and considered this
feature as an apomorphy of some species of this genus.
Clade 9 extends from the Late Cretaceous (Baluchicardia
beaumonti) to the Recent (Purpurocardia purpurata and
Cardiocardita ajar). The maximum taxonomic diversity
of the group is achieved during the Eocene.
According to the phylogenetic analyses presented
here, there is a clear separation between planicostate
and alticostate carditids, as previously stated by Harris
(1919), and both lineages could be monophyletic, with
the inclusion of some additional genera: Bathycardita
among planicostates and Paraglans, Arcturellina,
Cardiocardita, Kalelia, Darwinicardia and possibly
Cardites among alticostates. Both lineages comprise
members of the subfamily Venericardiinae sensu
Chavan (1969), and alticostates also include genera of
the Carditesinae.
 Phylogeny of Carditidae
The phylogenetic position of Kalelia
The genus Kalelia was defined recently by P
erez & del
R
ıo (2017a) and includes three species from the
Paleocene of Argentina and France: K. multicostata, K.
pectuncularis and K. burmeisteri. This taxon has large
shells and lunule, and a high hinge with a wide ventral
development, which are characters common among pla-
nicostate carditids. Also, Kalelia has tripartite radial
ribs sometimes restricted to a portion of the valve, a
key character of alticostate carditids. This combination
of features led Stewart (1930) to discuss the placement
of K. pectuncularis (as Venericardia pectuncularis) and
‘Venericardia’ marylandica Clark & Martin, 1901, a
species possibly included within Kalelia according to
P
erez & del R
ıo (2017a), among planicostate or alticos-
tate carditids. He concluded that they are alticostate
carditids with large-sized shells. Rutsch (1936) consid-
ered ‘V.’ pectuncularis and ‘V.’ multicostata as a pos-
sible section of V. imbricata, stating their placement
among alticostates. McClure (2009), in her phylogen-
etic analysis, recovered the same species among alticos-
tate carditids and mentioned the possibility that ‘V.’
pectuncularis convergently developed some planicostate
characters. In this analysis, Kalelia is placed as the sis-
ter group of a clade composed of Paraglans,
Arcturellina, Cardiocardita and Venericardia, among
alticostate carditids (Clade 9). This position allows us
to consider Kalelia as an alticostate carditid. Among its
synapomorphies we can list the presence of thick
shells, large lunules, and radial ribs short and wide in
the adult region, all of which are convergent characters
with those of planicostate carditids.
The phylogenetic position of Venericardia
The genus Venericardia was defined by Lamarck
(1801) and several extant and fossil species were
included in this genus subsequently (Huber 2010). The
assignment of Recent species to this genus is uncer-
tain and many of these were never reviewed (Huber
2010; P
erez & del R
ıo 2017a). In this analysis,
Venericardia is considered to be a fossil taxon
restricted to the Eocene of France (Huber 2010).
Chavan (1969) placed this genus within
Venericardiinae, along with the former planicostate
carditids, but according to the phylogeny presented
here Venericardia is grouped within the clade that
includes Cardiocardita, Paraglans and Arcturellina,
with the latter two genera also from the Eocene of
France. Subsequently, this result forces redefinition of
the subfamily Venericardiinae.
19
The composition of Carditamerinae and its
phylogenetic relationships
The Carditamerinae is the most diverse of Chavan’s
(1969) subfamilies, with more than 20 genera. The for-
mer carditamerines Arcturellina and Cardiocardita are
grouped within the alticostate carditids and Bathycardita
within planicostates. Scalaricardita and Vimentum are
placed with Coripia in a separate clade (Clade 4).
Pleuromeris and Cyclocardia are placed in different
positions in the analyses. Glans and Centrocardita are
the only genera recovered as sister taxa of Carditamera,
the type genus of the subfamily. This latter clade (Clade
2) is the only possible monophyletic group of the
Carditamerinae. As stated above, synapomorphies of
this group partially match those proposed by Chavan
(1969). The monophyletic Carditamerinae is composed
of Carditamera spp., Glans and Centrocardita. The
fourth group observed in the phylomorphospace agrees
with this separation of Clade 2, with species of
Carditamera sharing a similar portion of morphospace
and Glans and Centrocardita very close to them.
Among the four species of Carditamera, two lineages
are found in all of the phylogenetic searches: C.
arata þ C. floridana and C. affinis þ C. gracilis. These
lineages correspond with the separation between
Carditamera and Byssomera Olsson, 1961 (type species
Cardita affinis), proposed by Olsson (1961). Some
authors synonymized both taxa (Coan & Valentich-Scott
2012; Gonzalez & Giribet 2013) and Vermeij (2013)
confirmed the distinction based on new characters. The
results of the phylogenetic analyses performed here sup-
port this taxonomic separation into two taxa because C.
arata is the type species of Carditamera, while C. affi-
nis is the type species of Byssomera. A relationship
between Glans and Centrocardita was previously pro-
posed by several authors that considered these genera to
be synonyms (Glibert & Van de Poel 1970) or with
Centrocardita as subgenus of Glans (Sacco 1899;
Chavan 1969; Moore 1992; Beu 2006). Clade 2 is a
more temporally restricted clade (early
Miocene–Recent) than the previously considered
Carditamerinae (Eocene–Recent).
A phylogenetic analysis of Archiheterodonta carried
out by Gonzalez & Giribet (2015) recovered
Centrocardita as the sister group of the species of
Cardita and Thecalia. Dall (1902, 1903) included Glans
and Carditamera as subgenera of Cardita, and related
these taxa to mytiloid forms, and Thiele (1935) included
both taxa within Beguina. Bernard (1983) considered
Byssomera as a subgenus of Cardita. The relationship
between Carditamerinae (Clade 2) and Carditinae (Clade
3) is recovered in all of the analyses with high sup-
port values.
20 D. E. P
erez
The phylogenetic position of Thecaliinae
Dall (1902) established a separation between the sub-
family Thecaliinae and the remaining carditids, but this
small clade has not been related to other groups. This
author mentioned the resemblance between males of
Thecalia and species of Glans, and Thiele (1935) indi-
cated the same condition between males of Thecalia and
Mytilicardita Anton, 1838 (an old synonym of Cardita
according to Dall 1902, 1903). Gray (1854) also referred
Thecalia to Mytilicardita. Thecaliinae and Carditinae
share the presence of mytiliform or modioliform out-
lined shells with a ventral notch (byssal notch) and have
very similar right hinge teeth. The relationship between
both subfamilies is strongly supported in all analyses,
and in most searches Thecaliinae is nested within
Carditinae. Gonzalez & Giribet (2015) recovered
Thecalia concamerata within species of Cardita in their
phylogenetic analysis. The main difference between the
members of both subfamilies is the presence of a brood-
ing structure in Thecaliinae (‘marsupium’). Dall (1903)
and Yonge (1969) interpreted the marsupium of
Thecalia as an enlarged byssal gape. This structure may
be an invagination of an interspace between radial ribs,
which probably corresponds in placement to the byssal
notch already present in male specimens. Taxa referred
to the subfamily Thecaliinae could be a particular group
of Carditinae that co-opted this structure for brooding
(Gould & Vrba 1982). There is only one record of incu-
bation in Carditinae, in Cardita aviculina Lamarck,
1819 (Schneider 1993). This is also true for some extant
species of Cyclocardia (Jones, 1963) and is presumed
for several species of alticostates (Heaslip 1968; P
erez
et al. 2017b). Dall (1902) indicated that incubation is a
supposed condition for all carditids. Yonge (1969)
inferred the presence of incubation in the phylogenetic-
ally close Glans carpenteri (Lamy, 1922), but Harvey
(1995) recognized the external retention of eggs and lar-
vae for Carditamera floridana, and interpreted ancestral
ovoviparity for the species.
In the case of Milneria, the relationships with the
Thecaliinae are unknown, and Huber (2010) considered
that they are distinct lineages only linked by the pres-
ence of a marsupium. The Milneria lineage could have
developed the marsupium independently.
A new carditid lineage?
A previously unrecognized clade has been recovered in
the phylogenetic analyses performed herein. This group,
Clade 4, is composed of the genera Coripia, Vimentum
and Scalaricardita. Chavan (1969) placed these taxa in
two different subfamilies: Vimentum and Scalaricardita
in his superdiverse Carditamerinae, and Coripia in
Miodomeridinae. Scalaricardita and Vimentum were
considered subgenera (Sacco 1899; Lamprell & Healy
1998) or synonyms (Thiele 1935) of Cyclocardia.
Coripia was considered a synonym (Dall 1902;
Cossmann & Peyrot 1912; Thiele 1935) or a subgenus
(Chavan 1969) of Pteromeris. Popov (1983) placed C.
unidentata (type species of Coripia) in the genus
Scalaricardita. This new lineage is strongly supported
and defined by several synapomorphies (see
Supplementary material, Appendix 5).
The new group is related to Cyclocardia in all of the
searches and the new lineage þ Cyclocardia is close to
Pleuromeris and Miodomeridinae in several results
(Clade 12). The monophyly of this group is poorly sup-
ported, but morphological similarities are recovered in
the disparity analysis, forming together the second group
and being positioned away from the remaining carditids
(see Results, above). This grouping could be the result
of convergent similarities, but they deserve further study
to discuss their evolutionary origin.
The monophyly of Pleuromeris and Cyclocardia
Pleuromeris is a very diverse, with around 30 species,
and is distributed from the late Oligocene to Recent. Its
geographical distribution includes South America, south-
eastern North America, Central America, the Caribbean
Sea, Australia and New Zealand. Five species of
Pleuromeris were included in the phylogenetic analyses,
representing three different regions (New Zealand,
south-eastern North America and South America) and
they are paraphyletic in some results. The South
American species, P. fueguina and P. sulcolunularis, are
recovered together in all searches, and the North
American (P. decemcostata and P. tridentata) and New
Zealand (P. zelandica) species form a monophyletic
clade. The composition of the genus has never been
reviewed since Conrad (1867) and Dall (1903) defined
its principal features. A more complete revision of the
phylogenetic relationships within Pleuromeris is needed
to resolve this question.
More than 100 species were assigned to Cyclocardia
in the literature, including living and fossil species cov-
ering a long time interval (Eocene–Recent) from South
America, eastern Asia, North America, Antarctica and
Europe (e.g. Maxwell 1969; Coan 1977; Stilwell &
Zinsmeister 1992; Kafanov et al. 2001; Janssen &
Moerdijk 2004; Huber 2010; G€uller & Zelaya 2013).
Cyclocardia (Clade 10) shows low support values and is
not monophyletic in some searches. From the morpho-
logical disparity analysis, the species of this genus
occupy a considerable portion of morphospace.
According to Maxwell (1969), New Zealand representa-
tives of Cyclocardia may belong to a different lineage,
called the ‘awamoensis’ group. del R
ıo (1986) placed
 Phylogeny of Carditidae
within this group the Miocene Cyclocardia nortensis,
and this species is recovered as the sister taxon of C.
awamoensis in these results. Maxwell (1969) and Beu &
Maxwell (1990) suggested that morphological similar-
ities between members of the ‘awamoensis’ group and
other Cyclocardia species are the result of convergence.
Coan (1977) mentioned at least two species-complexes
within the genus (one related to C. crebicostata from
the Bering Sea and other related to C. ventricosa from
the Gulf of Alaska to Baja California). Scarlato (1981)
related Alaskan and Japanese species with the genus
Crassicardia Savitzky, 1979, a taxon that separated C.
crassidens (Broderip & G. B. Sowerby I, 1829) from
Cyclocardia. Popov (1980) supported this separation
based on ontogenetic differences between C. crassidens
and C. borealis (type species of Cyclocardia). Later,
Popov (1983) erected the genera Lunulicardita and
Ainicardita for a group of several Russian species previ-
ously related to Cyclocardia or Venericardia but this
approach has not been followed by subsequent authors.
Bernard (1983) considered the existence of a different
subgenus for extant South American species assigned to
Cyclocardia. Other species also included in Cyclocardia
were placed in Megacardita by Popov (1983) but La
Perna et al. (2017) rejected these assignments.
According to the large amount of intraspecific variabil-
ity on this genus, Huber (2010) proposed the existence
of at least three subgenera within Cyclocardia. A com-
plete revision of the species of Cyclocardia is needed in
order to clarify the composition and phylogenetic rela-
tionships of this taxon.
The phylogenetic position of Palaeocarditinae
and the origin of Carditidae
Palaeocarditinae includes pre-Cretaceous carditids
according to Chavan (1969) and is the only subfamily
recovered sensu Chavan (1969) in the phylogenetic
results presented here. Clade 1 is consistent with this
subfamily and has a high degree of support, with several
synapomorphies. Many morphological features of these
species are shared with outgroups (astartids and crassa-
tellids), as can be seen in the phylomorphospace. These
features include: a straight dorsal margin; slightly proso-
gyrate umbones; presence of escutcheon; large lunule;
and right anterior tooth limited to a lamina adjacent to
the lunular margin. Other characters are shared with all
carditids, such as: the absence of an internal resilifer;
presence of radial sculpture; and correspondence of ven-
tral margin crenulations with interspaces.
Palaeocarditinae shows intermediate morphological fea-
tures between Carditidae and the rest of
Archiheterodonta.
21
The morphological distance of this group as depicted
in the phylomorphospace agrees with the temporal gap
between Palaeocarditinae and other carditids, with more
than 100 million years of separation (from the Late
Triassic to the Late Cretaceous). There are some pos-
sible records of Carditidae in the Early Jurassic of
Argentina (Ferrari & Bessone 2015) and France
(Gervais 1853) and the Late Jurassic of Germany
(Paulsen 1964) and Portugal (Werner 1986), but these
have been referred to the genera Venericardia or
Cardita without a more precise assignment. Marwick
(1953) assigned an Early Jurassic carditid from New
Zealand to Palaeocardita. Scarlato & Starobogatov
(1979) considered Palaeocarditinae related to
Carditamerinae, Venericardiinae and Carditesinae.
The origin of Carditidae is unclear and different
authors proposed that this family could be related to the
fossil Permophoridae (Chavan 1969) or
Archaeocardiidae, Eodonidae, Aenigmoconchidae,
Cardiniidae and Myophoricardiidae (Scarlato &
Starobogatov 1979; Carter et al. 2011). The genus
Gujocardita Nakazawa & Newell, 1965 (late Permian,
Japan) was considered by Nakazawa & Newell (1965)
as a transitional form between Permophoriidae and
Palaeocardita. The hinge teeth and the displacement of
muscle scars of Gujocardita are present in
Palaeocarditinae and other Archiheterodonta, and the
shell outline and orientation of umbones are similar to
species of Permophoriidae. The genus Septocardia (Late
Triassic of South America, North America and Russia)
is often considered an intermediate form between
Cardiidae and Carditidae (Keen 1969; Schneider &
Carter 2001; Damborenea & Mance~nido 2012). The
presence in Septocardia of an escutcheon and the dis-
placement towards the anterior margin of the muscle
scar resemble Palaeocardita and other archiheterodonts,
and the external sculpture is more similar to that of car-
ditids than cardiids.
Bouchet et al. (2010) and Carter et al. (2011) consid-
ered Palaeocarditinae as a separate family
(Palaeocarditidae). The relationships between
Palaeocarditinae and the remaining carditids need more
revision, including the revision of other Mesozoic
records of the family. The phylogenetic and morpho-
logical disparity analyses allow consideration of the
relationships between Palaeocarditinae and other archi-
heterodonts. Bouchet et al. (2010) considered the group
within Archiheterodonta (Carditida in their classifica-
tion), but by contrast Carter et al. (2011) placed
Palaeocarditidae within Euheterodonta and close to
Cardiidae. Morris & Eagar (1978) and Morris et al.
(1991) placed the Palaeocarditinae close to
Permophoridae and the latter within the
22 D. E. P
erez
Anomalodesmata (in the Euheterodonta group). The
phylogenetic placement of the Palaeocarditinae needs
more research. If the group is placed with the Cardiidae
or outside Archiheterodonta, the temporal gap indicated
by the results recovered here is no longer valid and the
complete origin of the Carditidae should be looked for
in some Mesozoic bivalve group.
As mentioned above, the study of some possible
Palaeocarditinae (Septocardia, Gujocardita) may shed
light on the relationship of this clade with the remaining
carditids and, possibly, on the origin of Carditidae.
More research on other Mesozoic carditids not included
in these analyses (e.g. Fenestricardita, Trapezicardita,
Maghrebella, and others mentioned later in this paper),
could help to interpret the phylogenetic relationships
and characteristics of early carditids. Future phylogen-
etic analyses should look into the unique and shared fea-
tures of Palaeocarditinae, Mesozoic carditids and post-
Cretaceous carditids.
Homoplasy and phylogenetic value of characters
Analysis of the phylogenetic signal highlighted the high
systematic value of escutcheon characters. These fea-
tures represent an important hallmark for the separation
of Carditidae from other related families. Other charac-
ters with high phylogenetic signal have the same sys-
tematic pattern (e.g. the presence of internal resilifer,
laminar lateral teeth, presence of radial ribs, presence of
inner crenulations). These characters agree with those
commonly used to diagnose Carditidae (Morton et al.
1998; Coan et al. 2000).
Among the internal phylogenetic relationships, char-
acters such as the development of the dorsal margin,
relative position of the umbones, development of the
right hinge teeth (particularly anterior tooth), extension
of right ventral edge of hinge, development of left anter-
ior tooth, type of radial ribs, interspaces, and features of
the comarginal nodes and muscle scars are the most use-
ful for reconstructing relationships. The right hinge and
comarginal sculpture features are the most valuable.
Chavan (1969) considered mainly outline and hinge
characters (especially the right hinge) for the subfamiliar
scheme and made few mentions of sculptural characters,
but these results indicate a different scenario. By con-
trast, outline characters have a high degree
of homoplasy.
Some characters with high values of phylogenetic sig-
nal are synapomorphies of different clades. Dorsal mar-
gin characters and the direction of umbones are
important to separate outgroups and Palaeocarditinae
from post-Triassic carditids. The presence of an incuba-
tory chamber is a non-homoplasic feature only present
in Thecaliinae, as indicated by Dall (1902, 1903). A
serrated posterior margin is recovered here as a key fea-
ture of true Carditamerinae, a character first mentioned
by Vermeij (2013). The lateral teeth are important char-
acters present in Palaeocarditinae and species of
Carditamera. The presence and development of a right
anterior tooth allow separation of the following lineages:
Carditinae, Cardites, Cyclocardia, Thecaliinae,
Venericardiinae and North American planicostates.
Some features of the right middle tooth are also import-
ant for some lineages, such as Venericardiinae and pla-
nicostates. Some features of the right posterior tooth are
remarkable characters for Carditinae, among other
clades, as illustrated by Lamy (1922). The projection
adjacent to the nymph margin, a character first men-
tioned by P
erez et al. (2017a), is a non-homoplasic
character of planicostate carditids. The presence of pro-
socline radial ribs is a key feature of Carditamera spp.
that was not described before. The types of comarginal
nodes and separation between them are important for
distinguishing different genera.
Systematic implications
A new classification is proposed herein (Figs 9, 10,
Table 2), based on the results of the phylogenetic, dis-
parity and phylogenetic signal analyses. Firstly,
Carditidae is separated into two major clades:
Palaeocarditinae and Eucarditidae (new clade). This sep-
aration reflects their morphological differences and the
long temporal gap between Palaeocarditinae and remain-
ing carditids. If Palaeocarditinae is no longer a member
of Archiheterodonta, as it has been considered to be by
some authors (Morris & Eagar 1978; Morris et al. 1991;
Carter et al. 2011), Carditidae and Eucarditidae are syn-
onyms and the latter becomes obsolete. Molecular
phylogenetic analyses of living carditids (Gonzalez &
Giribet 2015) estimated the divergence of the group in
the Early Jurassic. Because this topic remains unsolved,
Eucarditidae is proposed herein. Some possible
Mesozoic carditids are mentioned below
(Fenestricardita, Trapezicardita, Izumicardia and
Xenocardita), but their relationships with the rest of the
family are still unclear. The characters present in these
genera separate these taxa from other true carditids (see
below). These taxa could be placed as successive sister
clades to Eucarditidae and fill the morphological gap
obtained in the analysis presented herein. More studies
about Mesozoic carditids are needed to explain the dif-
ferences between Palaeocarditidae and Eucarditidae.
In addition to Palaeocarditinae, five of the original
subfamilies proposed for Carditidae (Dall 1902; Chavan
1969) are maintained, but with some modifications.
Thecaliinae comprises the closely related taxa Thecalia
and Powellina. The subfamily Carditinae incorporates
 Phylogeny of Carditidae 23

Figure 10. Reduced version of the tree including only genus-level taxa showing the subfamiliar scheme and clades proposed in this
paper. Different colours indicate the different subfamilies (or tribes within Venericardiinae).

the Cretaceous genus Birkelundita, interpreted as closely
related to Cardita and Beguina by Heinberg (1993).
Carditamerinae is modified to include only three taxa:
Glans, Centrocardita and Carditamera. The clade com-
posed of Carditamerinae, Carditinae and Thecaliinae is
recovered in all of the phylogenetic searches and is
named here as Carditida (new clade) to highlight these
relationships. The Miodomeridinae is now composed of
Miodomeris, Pteromeris and Kolmeris. The Pliocene
taxa Kolmeris has been considered closely related to
Pteromeris by P
erez & del R
ıo (2017b). A new subfam-
ily is erected herein, Scalaricarditinae (new subfamily),
containing Scalaricardita, Vimentum and Coripia. This
new subfamily, Miodomeridinae and the genera
Cyclocardia and Pleuromeris share a particular sector of
the phylomorphospace and this group is obtained in sev-
eral searches. This new lineage is named herein as
Cyclocardida (new clade). The Venericardiinae is the
most diverse subfamily, including two lineages formal-
ized herein. The first lineage, the Venicorini (new tribe:
formerly planicostates carditids), includes Bathycardita,
Megacardita, Neovenericor, Pacificor, Leuroactis and
Venericor. A close relationship between Megacardita
and planicostates was previously mentioned by Stewart
(1930), Verastegui (1953), Glibert and Van de Poel
(1970) and P
erez et al. (2017a). Bathycardita is for the
first time included in this subfamily. The second lin-
eage, the Venericardiini (new tribe: formerly alticostate
carditids), includes Purpurocardia, Darwinicardia,
Kalelia, Venericardia, Cardiocardita, Paraglans,
Arcturellina, Baluchicardia, Claibornicardia,
Rotundicardia, Glyptoactis and Fasciculicardia. The
genus Cardites may be included in Venericardiini but
its correct placement among Venericardiinae is doubtful.
Huber (2010) included Cardiocardita and
Purpurocardia within Venericardiinae and Beu &
Maxwell (1990) included in this subfamily the genera
Fasciculicardia and Purpurocardia. The phylogenetic
positions of Kalelia and Darwinicardia agree with those
found by P
erez & del R
ıo (2017a, b). All of the taxa
previously considered within Carditesinae are placed in
this subfamily.
There are no previous phylogenetic analyses for
Carditidae as a whole, but the molecular phylogenetic
analysis of Gonzalez & Giribet (2015) sampled several
extant carditid species. They recovered a strong
24 D. E. P
erez

Table 2. Composition of subfamilies proposed here.

Subfamily Genera included
Palaeocarditinae Palaeocardita, Schizocardita
Carditinae Cardita, Beguina, Birkelundita
Carditamerinae Carditamera, Glans, Centrocardita
Miodomeridinae Miodomeris, Pteromeris, Kolmeris
Venericardiinae Venericardia, Venericor, Leuroactis, Pacificor, Megacardita, Glyptoactis, Claibornicardia,
Baluchicardia, Paraglans, Cardites, Arcturellina, Cardiocardita, Bathycardita,
Fasciculicardia, Purpurocardia, Rotundicardia, Kalelia, Darwinicardia, Neovenericor
Scalaricarditinae Scalaricardita, Vimentum, Coripia
Thecaliinae Thecalia, Powellina

Table 3. Possible placement of genera not included in the remains monophyletic although it is included within
phylogenetic analysis. Carditinae. The different placement of Centrocardita
Genus Possible phylogenetic position and Carditamera contradicts the new composition of the
Ainicardita Cyclocardida subfamily Carditamerinae but more research is needed
Akardita Scalaricarditinae? in this topic, including on more living representatives of
Carditomantea Carditidae? the group. A consequence of the phylogenetic relation-
Choniocardia Scalaricarditinae or Cyclocardida ships obtained by Gonzalez & Giribet (2015) is the
Cossmannella Venericardiini
Crassicardia Cyclocardida
partial contradiction with the clade Carditida obtained in
Cretocardia Venericardiini the analyses performed herein. The genera
Cycloglans Venericardiinae Centrocardita, Cardita and Thecalia may correspond to
Fenestricardita Stem-Carditidae? the Carditida, but Carditamera is placed among
Goossensia Carditinae? Venericardiinae (Megacardita and Cardites). The
Gujocardita Stem-Carditidae? or Permophoridae
Izumicardia Stem-Eucarditidae or Stem-Carditida absence of fossil taxa and the reduced sample of the
Lazariella Venericardiini molecular analysis could explain these differences.
Lunulicardita Cyclocardida Cyclocardida, the other higher level clade proposed with
Maghrebella Venericardiinae? living representatives, is not contradicted because there
Malarossia Scalaricarditinae are no members of this group included in the molecu-
Milneria Uncertain
Miodontiscus Miodomeridinae lar analysis.
Pseudocardia Eucarditidae? (some species) Phylogenetic definitions (de Queiroz & Gauthier
Septocardia Stem-Carditidae? or Cardiidae 1990, 1992; Sereno 2005) and unambiguous synapomor-
Subvenericardia Venericardiinae? phies for all the clades discussed here are provided in
Strophocardia Venericardiinae
the Systematic palaeontology section, below.
Trapezicardita Stem-Carditidae?
Xenocardita Uncertain
Phylogenetic placement of some carditid with
monophyletic relationship between Carditamera and uncertain affinities
Megacardita, a placement of Thecalia within the species Beyond the 38 carditid genera included in this analysis,
of Cardita, and a close relationship between 24 other genera were mentioned in the literature as
Centrocardita and Cardita, but without a close relation- members of this family. Some of these taxa were
ship between the first and Carditamera. The first result discussed in previous sections (Carditomantea,
of Gonzalez & Giribet (2015) is not recovered in this Septocardia, Gujocardita, Milneria, Crassicardia,
analysis and contradicts the distant phylogenetic position Lunulicardita and Ainicardita). A list of these genera
of the subfamilies Carditamerinae and Venericardiinae. with their possible placements is provided in Table 3.
This discrepancy may be a result of the absence of fos- The Indian and Mozambican taxon Cretocardia is
sil taxa between both lineages in the molecular analysis. one of the few Cretaceous carditids (Stoliczka 1871;
The placement of Thecalia within Carditinae is recov- Gliozzi & Malatesta 1983) and has a strongly inflated
ered in some searches and the subfamily Thecaliinae shell sculptured with tripartite radial ribs that resemble
may become obsolete as a result. This possibility is dis- those of Baluchicardia. This similarity was already
cussed above but the maintenance of a Thecaliinae clade mentioned by Gliozzi & Malatesta (1983). Cretocardia
is preferred in this work because the nested position is is a Venericardiini carditid, closely related to
not obtained in all of the searches and the clade always Baluchicardia and Claibornicardia.
 Phylogeny of Carditidae
The Cretaceous taxa Fenestricardita (UK, Japan and
Spain) (Casey 1961; Tashiro & Kozai 1982; Garc
ıa
Garmilla & Calzada 1984) and Trapezicardita (UK)
(Casey 1961) are probably closely related to each other
because of their trapezoidal outline and similar sculp-
ture, which consists of marked comarginal ribs and
weak radial ribs. Their phylogenetic placement among
carditids is unknown and are mentioned only in the sub-
familiar arrangement of Chavan (1969). The phylogen-
etic affinities of these genera are puzzling and their
outlines resemble those of Palaeocardita, but the
absence of an escutcheon and slightly prosogyrous
umbones do not allow them to be placed within
Palaeocarditinae. A strong comarginal sculpture is not
present in other carditids, so they could be basal carditid
or stem-carditid taxa. More research about these taxa
may clarify their position within the Carditidae.
The genera Subvenericardia Freneix, 1972 and
Maghrebella Freneix, 1972 were erected for species
from the Cretaceous of west Africa. Subvenericardia
was related to Venericardiinae by its original author,
specifically with Venericor. These taxa share characters
with Venericardiini (small sized shells and strongly
sculptured radial ribs), or Venericorini (high hinge teeth
and strongly marked muscle scars). Maghrebella
presents a hinge and sculpture similar to species of
Venericardiini, but its ventrally projected outline is not
shared with other members of this tribe.
Subvenericardia and Maghrebella could be placed
within Venericardiinae.
Xenocardita (Cretaceous, Lebanon and Japan) (Vokes
1946; Tanaka et al. 1996) presents a particular hinge
morphology and spinose radial ribs with sharp comargi-
nal lamellae in the posterior side. Vokes (1946) inter-
preted the presence of only two right cardinal teeth and
one left anterior lateral tooth but Chavan (1969) indi-
cated three right cardinals. The interpretation of Chavan
(1969) is more accurate and this taxon shares some
characters with Venericardiinae, but the features of the
right hinge resemble those of Pleuromeris and other
Cyclocardida species. The phylogenetic position of
Xenocardita remains uncertain.
The hinge of Izumicardia (Late Cretaceous, Japan)
(Ichikawa & Maeda 1963) shows anterior and posterior
laminar teeth in both valves and a broad triangular right
middle tooth. The shells of Izumicardia also present
prosogyrous umbones, an escutcheon and numerous
strong radial ribs. Ichikawa & Maeda (1963) related
Izumicardia to the Glans group and the poorly known
genus Cycloglans Gorodiski & Freneix, 1959 (in
Freneix 1959) (Eocene, Senegal). Izumicardia shares a
strong similarity in the hinge configuration with that of
Cycloglans, but it differs in its outline and external
25
sculpture. The presence of an escutcheon separates this
genus from Eucarditidae, but the remaining characters
resemble those of species of Carditida.
The confusingly named genus Pseudocardia Conrad,
1866 (Late Cretaceous, Europe and North America)
(Scott 1977; Keen 1980) was originally included in
Carditidae, but Chavan (1969) did not assign it to any
subfamily. Conrad (1868) considered this genus as part
of the Astartidae, but Stoliczka (1871) and Cox (1946)
concluded that Pseudocardia is a carditid related to
Venericardia. Some European species of Pseudocardia,
such as P. tenuicosta (J. de C. Sowerby in Fitton 1836),
may correspond to Cardiidae because of their external
sculpture and escutcheon, and the North American, e.g.
P. arivechensis (Heilprin, 1891), and other European
species, e.g. P. dupiniana (d’Orbigny, 1843), could be
included in Carditidae. The presence in these species of
strong radial ribs, umbones prosogyrous, the configur-
ation of hinge teeth and the absence of an escutcheon
may place Pseudocardia in Eucarditidae, but a more
precise placement is not possible. More studies about
this poorly known genus are needed.
Choniocardia, from the Eocene of the Paris Basin
(Cossmann 1905), has a unique pseudoescutcheon, an
enlarged lunule and a particular hinge configuration that
hides its synapomorphies. Glibert & Van de Poel (1970)
and Pacaud & Le Renard (1995) considered
Choniocardia as a subgenus of Pleuromeris. The small
size of the shells and some particularities of the teeth
and sculpture (including small comarginal nodes), could
relate this taxon with the new clade Cyclocardida, and
possibly with the new subfamily Scalaricarditinae.
Choniocardia could be placed as a basal
Scalaricarditinae with some features shared with other
Cyclocardida species.
Another Eocene genus from the Paris Basin,
Goossensia (Cossmann 1885; Pacaud & Le Renard
1995), has a very variable subtrapezoidal and elongate
outline. The external sculpture of Goossensia includes
irregular comarginal lines above radial ribs. Juvenile
shells show a mytiliform outline, very similar to that of
Thecalia or Cardita species, and strong comarginal
lamellae. Goossensia may represent an apomorphic
taxon derived from a carditine ancestor.
Malarossia, from the Eocene of Ukraine (Berezovsky
1999), was originally included in the subfamily
Venericardiinae. Berezovsky (1999) indicated similar-
ities of this genus with Venericardia and Cyclocardia,
but the size, outline, external sculpture and hinge fea-
tures strongly resemble those of Scalaricardita.
Malarossia could be included in the subfamily
Scalaricarditinae.
26 D. E. P
erez
Cossmannella, an Eocene taxon from Egypt (Mayer-
Eymar 1896), has an elongated outline and a few sculp-
tured radial ribs. This taxon resembles Lazariella and
the Nigerian genera described by Eames (1957):
Amekiglans, Bendeglans and Divergidens (all invalid
taxa according to Chavan 1969). Thiele (1935) and
Chavan (1969) placed Lazariella as a subgenus of
Carditamera, but Huber (2010) discussed this and indi-
cated that Lazariella is much closer to Cardiocardita.
Chavan (1938) also related Cossmannella with
Cardiocardita. Eames (1957) regarded his taxa as sub-
genera of Glans. All of these genera could be closely
related to each other and are probably related to
Cardiocardita because of their similar tripartite radial
ribs and some features of the hinge teeth and umbones.
This group could represent an African lineage of the
Venericardiini tribe, closer to Venericardia and other
European taxa.
Cycloglans is another obscure genus based on speci-
mens from the Eocene of Senegal (Freneix 1959) with
anterior and posterior lateral teeth in both hinges and
very few strong radial ribs covered with subcircular
nodes. Freneix (1959) considered this taxon as a sub-
genus of Glans and Chavan (1969) synonymized it with
Pleuromeris. In many aspects, the characters of
Cycloglans can be considered juvenile states (high radial
ribs strongly sculptured, persistence of lateral teeth and
short cardinal teeth) and as a result this genus could be
considered as paedomorphic. The external sculpture and
outline remembers those of Darwinicardia. This genus
could be placed close to Venericardiinae.
The small-sized genus Miodontiscus
(Miocene–Recent, western North America and Japan)
(Moore 1992; Coan et al. 2000) has low and smooth
radial ribs, with conspicuous comarginal lines, and an
obtuse-triangular outline with a slightly inclined poster-
ior margin. The latter character is shared with
Pteromeris and Miodomeris species and allows inclusion
of Miodontiscus within the subfamily Miodomeridinae.
Dall (1903) already mentioned the similarity between
Miodontiscus and Pteromeris.
The recently defined genus Akardita La Perna,
Brunetti & Della Bella, 2018 (Pliocene–Recent, Europe
and West Africa) was not included in any of Chavan’s
(1969) subfamilies, but it was compared with several
genera. According to these authors, Akardita may
belong to the subfamilies Carditesinae or
Venericardiinae. The species assigned to this genus pos-
sess some similarities in their external sculpture and
hinge teeth with those of Cyclocardia and the subfamily
Scalaricarditinae. Some features of the genus could be
peramorphic, such as the low and obsolete radial ribs
towards the venter. Akardita may be a member of the
new subfamily Scalaricarditinae, but this position needs
more research.
The genus Strophocardia Olsson, 1961 was erected to
separate a group of Recent and fossil species from the
western coast of America (Olsson 1961). This genus was
originally related to species of Cardites and Bernard
(1983) included this genus in the subfamily Carditesinae.
Other authors considered it as a synonym of Cyclocardia
(Chavan 1969) or Lunulicardita (Coan & Valentich-Scott
2012). Strophocardia has large shells with a very triangu-
lar outline, strongly recurved umbones, a very high hinge
plate with long teeth and wide radial ribs. Some of these
characters are shared with planicostate (Venericorini) car-
ditids but could be the result of convergence. Right hinge
teeth with a strongly reduced anterior tooth and a strong
and curved middle tooth, recurved umbones and nodose
radial ribs resemble those of Cardites and may indicate a
close relationship between both genera. Because of this,
Strophocardia can be placed in the subfamily
Venericardiinae.
Comments on heterochronic processes in the
macroevolutionary history of the Carditidae
The new phylogenetic context presented herein allows
discussion of macroevolutionary trends within the fam-
ily, including those proposed by previous authors
(Yonge 1969; Stanley 1972; Heinberg 1993) as well as
considering new potential trends. This section is a first
attempt to summarize the possible heterochronic proc-
esses present in the group and to establish a starting
point for future research on these topics.
Life habits. The main life habit among carditids is
infaunal burrowing, especially in Cenozoic species.
Members of Carditinae, Thecaliinae and Carditamerinae
attach to different substrates via endo- or epibyssate
means. Heinberg (1993) proposed that the Carditidae radi-
ation is pendulate evolution with swings between these
two types of life habits. The presence of a byssus in the
adult stage, a prerequisite for a byssate mode of life, has
generally been regarded as a neotenic condition (Heinberg
1993). Stanley (1972) postulated it as the ancestral condi-
tion for all carditids, also considering neoteny as the evo-
lutionary mechanism, whereas Yonge (1969) considered
the infaunal burrowing type as the basalmost condition.
Neoteny is the reduced rate of morphological development
during juvenile stages resulting in a morphologically
retarded adult (McNamara 1986). Stanley (1972) included
in his postulate the presence of endobyssate species
among Triassic carditids (e.g. Palaeocardita crenata),
infaunal burrowers in the Cretaceous (e.g. Pseudocardia
tenuicosta – a possible Cardiidae instead of Carditidae)
and in the Cenozoic (e.g. Cyclocardia spp.). Another
 Phylogeny of Carditidae
lineage of byssate carditids is represented by Carditamera
spp. in the Pleistocene–Recent and is characterized by
elongated outlines. The ancestral condition is represented
by most Palaeocarditinae, while the infaunal burrowers
include many of the Venericardiinae and other subfamilies
from the Cretaceous–Recent. Some possible returns to a
byssate life habit are seen in Carditinae and
Carditamerinae species.
The most ancient carditids, members of the subfamily
Palaeocarditinae, represent a lineage of possible endo-
or epibyssate carditids. Many characters of this group
are shared with astartid or crassatellid infaunal bur-
rowers, but they are also present in the bivalve family
Permophoridae, a putative ancestral group for all cardi-
tids, composed of many byssate species (Stanley 1972).
Nevertheless, these evolutionary pathways depend on
the phylogenetic placement of Palaeocarditinae (see
above). More studies including possible Mesozoic cardi-
tids could contribute to this topic.
The possible common origin between Carditinae
(including Thecaliinae) and Carditamerinae taxa
(Carditida clade) proposed here includes the two lineages
of byssate eucarditids. The most ancient Carditinae,
Birkelundita, is a shallow burrower (Heinberg 1993).
According to the latter author, this taxon has many fea-
tures related to a byssate mode of life, reflecting a pos-
sible shift from this habit to a burrowing one in the
evolutionary history of the lineage. The phylogenetic pos-
ition of Birkelundita allows consideration of the proposal
of Stanley (1972) of an ancestral byssate condition (with
a transition in Birkelundita to an infaunal burrower mode
of life), but the unsolved position of Carditinae among
other carditids forces us to also consider a secondary tran-
sition to a byssate habit as proposed by Yonge (1969).
Peramorphosis in the evolution of Carditidae.
Peramorphosis is developmental change in which an
ancestral adult morphology is seen in a descendant
juvenile stage of development (McNamara 1986). This
is a heterochronic process that produces parallels
between ontogeny and phylogeny (Gould 1977).
Peramorphosis could have played an important role in
the evolution of Carditidae. For example, in juvenile
shells of planicostate venericardiids (Tribe
Venericoriini) the radial ribs are sharp and their sculp-
ture resembles alticostate ribs (Tribe Venericardiini).
This similarity suggests that the evolution of smooth
radial ribs in Venericoriini taxa could be a result of per-
amorphosis (see also McClure & Lockwood 2015). A
similar case is reported for some Cardiidae bivalves,
Cardium plicatum and Cardium fittoni, from the late
Miocene of Russia (Nevesskaya 1967; Gould 1977).
Other possible peramorphic characters, such as the pres-
ence of obsolete radial ribs and very thick shells
27
towards the ventral margin, can be found in some
Venericoriini taxa and in some species of Cyclocardia.
Both features are related to gerontic variation in
bivalves (Alvarez & P
erez 2016) and could be the result
of peramorphic change.
Paedomorphosis in the evolution of Carditidae.
Paedomorphosis is the phenomenon of phylogenetic
change in which an ancestral juvenile character is
retained in the descendant adult phase (McNamara
1986). As mentioned above, Stanley (1972) and
Heinberg (1993) proposed that the retention of a byssus
in the adult stage of Carditamera spp. could be a case
of paedomorphosis (Gould 1977). Paedomorphic proc-
esses could have driven the evolution of byssate cardi-
tids in the subfamilies Carditamerinae, Carditinae and
Thecaliinae. The tiny size of some Thecaliinae may be
another paedomorphic character.
Systematic palaeontology
Carditidae F
erussac, 1822
Definition. The most recent common ancestor of
Palaeocardita, Cardita and Venericardia and all of its
descendants (node-based).
Synapomorphies. Visible umbones outline in internal
view (c. 28); right anterior laminar teeth absent (c. 46);
left anterior laminar teeth absent (c. 73); radial sculpture
present (c. 90); undulations absent (c. 95); interspaces
present (c. 111); ventral margin crenulations correspond-
ing to interspaces (c. 139).
Temporal range. Late Triassic to Recent (excluding
Carditomantea).
Remarks. Other character present in this group is the
absence of an internal resilifer, apex of right anterior
teeth not resting on middle tooth side, apex of middle
tooth reaching the dorsal extreme of hinge, left teeth
without hook-like outline, and absence of a flat zone
between anterior muscle scar and anterior margin. All
these characters are shared with the Astartidae but are
present in the Crassatellidae.
Palaeocarditinae Chavan, 1969
Definition. The most recent common ancestor of
Palaeocardita and Schizocardita and all of its descend-
ants (node-based).
Synapomorphies. Angle between posterior and dorsal
margins strongly marked (c. 12); dorsal margin elongation
present (c. 14); notch in posterior margin present (c. 17);
strongly rounded umbones in external view (c. 27).
Taxa included. Palaeocardita Conrad, 1867;
Schizocardita K€orner, 1937.
28 D. E. P
erez
Temporal range. Late Triassic to Early Jurassic.
Remarks. Clade 1 in the text. Some of the synapomor-
phies of this subfamily are also present in species of
Carditamerinae and Carditinae. These similarities could
be the result of convergent evolution related to elon-
gated outlines.
Eucarditidae New clade
Definition. The most inclusive clade containing Cardita
and Venericardia, but not Palaeocardita, and all of its
descendants (stem-based).
Synapomorphies. Prosogyrous umbones; escutcheon
absent; presence of comarginal nodes; anterior adductor
muscle scar not displaced towards hinge margin.
Temporal range. Late Cretaceous (by Birkelundita tur-
oniense) to Recent.
Remarks. Other characters present in this clade are a
right anterior tooth not limited to a small sheet adjacent
to lunular margin and at least prominent, left posterior
tooth inclined less than 45 , comarginal lines absent
(except in Miodomeris) and adductor muscle scars
slightly marked (except in species of Miodomeris and
Venericorini). This new clade is proposed to highlight
the separation between Palaeocarditinae and the
remaining carditids obtained from the results.
Carditinae F
erussac, 1822
Definition. The most recent common ancestor of
Cardita, Birkelundita and Beguina and all of its
descendants (node-based).
Synapomorphies. Right anterior tooth vertical.
Taxa included. Cardita Bruguiere, 1792; Beguina
R€
oding, 1798; Birkelundita Heinberg, 1993.
Temporal range. Late Cretaceous (Birkelundita
turoniense)–Recent.
Remarks. Clade 3 in the text. Other characters present
in this clade include: the left anterior tooth inclined
between 0 and 50 (except in Beguina where it is hori-
zontal); and V-shaped cross section of interspaces
in adult shells (except in Birkelundita). In some
searches, Thecaliinae is nested within this subfamily
and could be synonymous. The inclusion of
thecaliines would increase considerably the synapomor-
phies of this subfamily (see Supplementary material,
Appendix 5).
Thecaliinae Dall, 1902
Definition. The most recent common ancestor of Thecalia
and Powellina and all of its descendants (node-based).
Synapomorphies. Mean size of adult shells near to
10 mm; presence of an incubatory chamber in females;
right anterior tooth inclined backwards; left anterior
tooth vertical and subrectangular; U-shaped cross section
of interspaces in adult shells.
Taxa included. Thecalia Dall, 1902; Powellina
Huber, 2010.
Temporal range. Recent.
Remarks. Clade 11 in the text. The close relationships
of Thecaliinae with Carditinae are already mentioned
and some of their features are present in members
of Carditinae.
Carditamerinae Chavan, 1969
Definition. The most recent common ancestor of
Carditamera, Glans and Centrocardita and all of its
descendants (node-based).
Synapomorphies. Angle between posterior and dorsal
margins strongly marked; angle between posterior and
ventral margins rounded; serrated posterior margin pre-
sent; recurved umbones; lunule subrounded; right
anterior tooth not reduced; left anterior lateral tooth
pustular present; left anterior tooth as high as wide;
equally heighted teeth in left hinge; extreme tips of
radial ribs pocking out between crenulations on
internal view.
Taxa included. Carditamera Conrad, 1838; Glans
Megerle, 1811; Centrocardita Sacco, 1899.
Temporal range. Early Miocene–Recent.
Remarks. Clade 2 in the text. The presence of less than
21 radial ribs is another character of this subfamily.
Carditida New clade
Definition. The most inclusive clade containing
Cardita, Thecalia and Carditamera, but not
Venericardia or Cyclocardia, and all of its descendants
(stem-based).
Synapomorphies. Umbones very anteriorly placed;
hinge very reduced and short; right anterior tooth div-
ided (shark tooth-like); right middle tooth with anterior
side convex; ventral extreme of right posterior tooth
protruding from ventral edge of hinge; left posterior
tooth nearly horizontal.
Temporal range. Late Cretaceous (Birkelundita
turoniense)–Recent.
Remarks. Other characters of this group are: right
middle tooth nearly horizontal; comarginal nodes as
scales (without comarginal nodes in Carditamera and
subrounded nodes in Beguina); and crenulations gentle
 Phylogeny of Carditidae
extended towards valve inside (limited to ventral edge
in Carditamera floridana and C. arata). This new
clade is proposed to highlight the relationships
between Thecaliinae, Carditinae and Carditamerinae.
Miodomeridinae Chavan, 1969
Definition. The most recent common ancestor of
Miodomeris, Pteromeris and Kolmeris and all of its
descendants (node-based).
Synapomorphies. Shell with obtuse triangular outline;
dorsal margin curved and descendent; umbonal area
with very low convexity (nearly flat shells); left poster-
ior tooth inclined between 35 and 45 ; subrectangular
cross section of radial ribs in juvenile shells; radial ribs
slightly elevated; smooth radial ribs.
Taxa included. Miodomeris Chavan, 1936; Pteromeris
Conrad, 1867; Kolmeris P
erez & del R
ıo, 2017b.
Temporal range. Middle Eocene (Miodomeris
cossmanni)–Recent.
Remarks. Clade 5 in the text. Other characters of
Miodomeridinae are: the posterodorsal margin with
slight to pronounced inclination; subrectangular cross
section of radial ribs in adult shells (except in
Miodomeris); and inverted trapezoidal cross section of
interspaces in adult shell (except in Miodomeris).
Scalaricarditinae New subfamily
Definition. The most recent common ancestor of
Scalaricardita, Coripia and Vimentum and all of its
descendants (node-based).
Synapomorphies. Slightly truncated posterior margin;
slightly marked lunule; right posterior tooth inclined
more than 35 ; radial ribs wider towards anterior side;
subovate cross section of radial ribs in juvenile shells;
small size of comarginal nodes.
Taxa included. Scalaricardita Sacco, 1899; Vimentum
Iredale, 1925; Coripia de Gregorio, 1885.
Temporal range. Middle Miocene (Scalaricardita
camaronesia)–Recent.
Remarks. Clade 4 in the text. Other characters of this
subfamily are: shell size less than 12 mm (excepting
Scalaricardita laciarina); subcentrally placed umbones
(excepting S. camaronesia); and pallial line placed at a
quarter of total valve height (excepting
Vimentum dilectum).
Cyclocardida New clade
Definition. The most inclusive clade containing
Cyclocardia, Pleuromeris, Scalaricardita and Miodomeris,
29
but not Venericardia, Carditamera or Cardita and all of
its descendants (stem-based).
Synapomorphies. Lunule sculptured only by comarginal
lines; pallial line as a uniform curve.
Taxa included. Cyclocardia Conrad, 1867; Pleuromeris
Conrad, 1867; Scalaricarditinae; Miodomeridinae.
Temporal range. Eocene–Recent.
Remarks. Clade 12 in the text. Other characters of this
group are: V-shaped cross section of interspaces in adult
shells (except in Miodomeridinae, Cyclocardia dalek
and C. nortensis). This clade is only recovered in the
analyses using discrete characters. This new clade is
proposed to highlight the relationships between
Cyclocardia, Pleuromeris, Miodomeridinae and
Scalaricarditinae as obtained in the phylogenetic and
phylomorphospace results.
Venericardiinae Chavan, 1969
Definition. The most recent common ancestor of
Venericardia and Venericor and all of its descendants
(node-based).
Synapomorphies. There are no unambiguous
synapomorphies.
Taxa included. Cardiocardita Anton, 1838; Arcturellina
Chavan, 1951; Bathycardita Iredale, 1925; Venericardia
Lamarck, 1801; Leuroactis Stewart, 1930; Pacificor
Verastegui, 1953; Venericor Stewart, 1930; Megacardita
Sacco, 1899; Neovenericor Rossi de Garc
ıa, Levy &
Franchi, 1980; Cardites Link, 1807; Glyptoactis
Stewart, 1930; Baluchicardia Rutsch & Schenk, 1943;
Claibornicardia Stenzel & Krause, 1957; Paraglans
Chavan, 1941; Fasciculicardia Maxwell, 1969;
Rotundicardia Heaslip, 1968; Purpurocardia Maxwell,
1969; Kalelia P
erez & del R
ıo, 2017a; Darwinicardia
P
erez & del R
ıo, 2017b.
Temporal range. Late Cretaceous (Baluchicardia
beaumonti)–Recent.
Remarks. Clade 13 in the text. This subfamily is sup-
ported by ambiguous apomorphies only. These are:
convex lunule (except in Kalelia multicostata, K. pec-
tuncularis, Leuroactis pilsbryi, Venericor aposmithii
and V. bashiplata); right middle tooth with anterior
side concave (except in Bathycardita raouli and
Cardiocardita ajar); right posterior tooth curved
(excepting Rotundicardia mariobrosorum); and poster-
ior area defined by less thick radial ribs than the
remaining (except Purpurocardia leonensis,
Venericardia imbricata, Cardiocardita ajar,
Arcturellina asperula, Paraglans calcitrapoides and
Rotundicardia spp.).
30 D. E. P
erez
Venericorini New tribe
Definition. The most inclusive clade containing
Venericor, Megacardita and Bathycardita, but not
Venericardia, and all of its descendants (stem-based).
Synapomorphies. Shells size more than 35 mm; slight
projected posteroventral margin; strong thickness of
shells; left posterior tooth inclined more than 20 ;
smooth radial ribs; strongly marked and very large
adductor muscle scars; pallial line inclined towards pos-
terior margin; crenulations extended below posterior
adductor muscle scar.
Temporal range. Paleocene–Recent.
Remarks. Clade 8 in the text. Members of this clade
have been referred to as ‘planicostate carditids’ in the
literature. Other characters include: presence of nymph
margin (except in Bathycardita raouli); left posterior
tooth strongly curved (except in Megacardita jouannetti
and B. raouli); subrectangular cross section of radial
ribs in adult shells (except in M. jouannetti, B. raouli
and Neovenericor carrerensis); and slightly elevated
radial ribs (except in Neovenericor spp.).
Venericardiini New tribe
Definition. The most inclusive clade containing
Venericardia, Claibornicardia and Purpurocardia, but
not Venericor or Megacardita, and all of its descendants
(stem-based).
Synapomorphies. There are no unambiguous
synapomorphies.
Temporal range. Late Cretaceous (Baluchicardia
beaumonti)–Recent.
Remarks. Clade 9 in the text. This subfamily is sup-
ported only by ambiguous apomorphies only. Members
of this clade are called ‘alticostate carditids’ in the lit-
erature. In some searches, the genus Cardites is included
in this clade and the Carditesinae of Chavan (1969)
would be a synonym as a consequence. Ambiguous syn-
apomorphies of the clade (including Cardites) are: regu-
lar convexity of umbonal area (except in Cardites spp.,
Glyptoactis hadra, Baluchicardia beaumonti and
Darwinicardia patagonica); slightly marked lunule
(except in D. patagonica); symmetric and non-projected
lunule (except in G. hadra, B. beaumonti, Rotundicardia
spp. and Claibornicardia spp.); and presence of paracos-
tal ribs (except in Cardites spp., Rotundicardia spp. and
Fasciculicardia sp.). When Cardites is excluded, new
ambiguous synapomorphies are added, including: regular
convexity of umbonal area (except in Glyptoactis hadra,
Baluchicardia beaumonti and Darwinicardia patagon-
ica); left anterior tooth higher towards dorsal (except
Cardiocardita ajar and F. sp.); and left posterior tooth
higher towards dorsal (except in Darwinicardia species).
Conclusions
A new classification for the family Carditidae is pro-
posed herein based on the results of phylogenetic, mor-
phological disparity and phylogenetic signal analyses.
This scheme includes seven subfamilies –
Palaeocarditinae, Carditinae, Thecaliinae,
Miodomeridinae, Carditamerinae, Venericardiinae and
the new subfamily Scalaricarditinae – and rejects the
previously proposed Carditesinae. The generic compos-
ition of all of the subfamilies is re-evaluated and
Palaeocarditinae is the only family recovered in its ori-
ginal sense. Other clades above or below the subfamily
level are proposed to accommodate the new relation-
ships obtained in these analyses. A distinct separation
between Palaeocarditinae and the remaining carditids is
discussed, and the Eucarditidae (all carditids excluding
Palaeocarditinae) is defined as a consequence. The
Carditamerinae and Thecaliinae are closely related to
Carditinae and constitute the new clade Carditida. A
new carditid lineage is recognized, Cyclocardida
(Miodomeridinae, Scalaricarditinae, Pleuromeris and
Cyclocardia). The monophyly of alticostate and plani-
costate venericardiids are discussed in light of the
phylogenetic results and both groups form clearly sepa-
rated monophyletic lineages: the new tribes
Venericoriini (formerly planicostate venericardiids) and
Venericardiini (formerly alticostate venericardiids) are
proposed to reflect this result. Phylogenetic definitions
are provided for all of the clades and the placements of
the other carditid genera that were not included in the
analyses are discussed.
Kalelia is confirmed as an alticostate taxon with con-
vergent planicostate characters and Venericardia is a
Venericardiini grouped with Cardiocardita, Paraglans
and Arcturellina. The monophyly of the genera
Pleuromeris and Cyclocardia is questioned and further
studies on these large and diverse taxa are needed. The
shared characters between Palaeocarditinae and other
archiheterodont bivalves are discussed in the context of
their phylogenetic and stratigraphic position Carditidae.
Finally, macroevolutionary trends in the history of
Carditidae, including the role of peramorphic and paedo-
morphic phenomena, are discussed. Changes in life hab-
its have driven the evolution of some lineages, including
the possible return to a byssate mode of life in
Carditinae and Carditamerinae (Carditida clade). Some
peramorphic (smooth radial ribs of Venericorinii and
thick shells towards ventral margin in Venericorinii and
 Phylogeny of Carditidae
Cyclocardia) and paedomorphic (retention of byssus in
Carditida clade and tiny size in Thecaliinae) characters
are recognized among carditids.
Acknowledgements
The author is indebted to those curators and researchers
who facilitated access to collections: M. Longobucco
and C. del R
ıo (MACN-Pi and CIRGEO-PI, Buenos
Aires), M. Tanuz (CPBA, Buenos Aires), A. Tablado
(MACN-In, Buenos Aires), A. Riccardi (MLP), L. P
erez
(MLP), M. De la Fuente and I. Maniel (MHNSR.Pi, San
Rafael), M. Rodr
ıguez (SGN, Buenos Aires), C. Salazar
(SGO.Pi, Santiago) and C. Fraz
en (PZ-NMR.Mo,
Stockholm). I wish to thank the following for providing
photographic material: T. Waller and M. Florence
(USNM.Mo, Washington, DC); M. Munt (NHMUK,
London); J-M. Pacaud (MNHM, Paris); P. Schuchert
and L. Cavin (GMNH, Geneve); J. Sime and P.
Callomon (ANSP, Philadelphia); J. Simes (TM, GNS
Science, Wellington); and T. Trnski, W. Blom and H.
Schlumpf (AK, Auckland). I thank M. Griffin (MLP)
and S. Popov (RAS, Moscow) for sharing bibliographic
data, W. Svagelj for help with the GLM analysis, M.
Ram
ırez (MACN-Ar) for help with terminal taxon
selection and theoretical design of the phylogenetic
analyses, and C. del R
ıo (MACN-Pi) for guidance
during my PhD thesis. I thank O. Lehmann (CPBA) for
the script for multiple implied weighting searching, C.
Mengoni Go~ nalons for help with art work, M. Alvarez
and M. B. Santelli (MACN-Pi) for their valuable
suggestions and assistance with figures. I also thank I.
Maniel (MHNSR), P. Milla Carmona (CPBA), O.
Lehmann (CPBA) and M. Ezcurra (MACN) for their
useful suggestions, help with different methodologies
and valuable comments, and I. Maniel, M. Ezcurra and
P. Picasso for their valuable help with language. Thanks
to S. Schneider (CASP) for his constructive comments
as a reviewer. CONICET is acknowledged for the post-
graduate grants given to the author and the use of TNT
software is facilitated by the Willi Hennig Society.
Supplemental data
Supplementary material for this article can be accessed
here: http://dx.doi.org/10.1080/14772019.2018.1532463
ORCID
Dami
an Eduardo P
erez http://orcid.org/0000-0001-
6771-4938
31
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Appendix 1

List of taxa revised for the phylogenetic analyses indicating source of information used
(specimens or bibliography). Numbers in italics indicate type specimens (holotypes,
paratypes or lectotypes). Citation in italics indicates illustration of type specimens:

Taxon Source of information

Arcturellina asperula MACN-Pi 955, 959
Astarte sulcata MNHN FR50738; NMR 25682, 40316

Baluchicardia beaumonti MNHN FJ06620; Heaslip 1968

Bathycardita raouli NMR 63303; Iredale 1925

Beguina semiorbiculata MACN-In 17486; NMR 19165; Heinberg 1993

Birkelundita turoniense Heinberg 1993

Cardiocardita ajar NMR 16051; Chavan 1938

Cardita crassa CPBA 3098; MACN-Pi 2353

Cardita variegata NMR 20001; Lamy 1922; Heinberg 1993

Carditamera affinis NHM 196377011; MACN-In 6027; NMR 16042

Carditamera arata UF 80045, 247405, 207359; NMR 67033; YPM IP

560023
Carditamera floridana MACN-In 41173; UF 5508; YPM IZ 050966

Carditamera gracilis MACN-In 2623

Cardites antiquatus MACN-In 1163, 28256; NMR 35663

Cardites feruglioi SGN 9379, 9381, 9395; MACN-Pi 4120, 4197,

5748, 5755, 5763–5765, 5767; CPBA 6411
Cardites partschii CPBA 1111, 3088
Claibornicardia acuticostata MACN-Pi 957; MNHN FJ07564; NMR 64570
Claibornicardia alticostata ANSP-IP 30562; YPM IP 025752

Claibornicardia paleopatagonica MACN-Pi 360, 4289, 5197; CPBA 17286, 17288,

21027; MLP 1475, 3731, 3732, 3734
Centrocardita aculeata MLP 5240, 5243; NMR 26649; Beu 2006

Coripia unidentata MNHN FA49875, FA49876, FJ14611

Crassatella ponderosa MACN-Pi 983; 3012; NMR 65305; GMNH 45970

Cyclocardia awamoensis MACN-Pi 5804; Maxwell 1969; Beu & Maxwell
1990
Cyclocardia cannada MACN-Pi 344, 345–348, 5772, 5778, 5779, 5780;
CPBA 9373; PRI.66848
Cyclocardia compressa
Cyclocardia borealis
Cyclocardia dalek
Cyclocardia mesembria
Cyclocardia nortensis
Cyclocardia ventricosa
Darwinicardia angusticostata
Darwinicardia patagonica
Fasciculicardia acanthodes
Fasciculicardia subintermedia
Fasciculicardia sp.
Glans trapezia
Glyptoactis hadra
Kalelia burmeisteri
Kalelia multicostata
Kalelia pectuncularis
Kolmeris tehuelchana
Laevastarte basteroti
Leuroactis pilsbryi
Megacardita jouanetti
Miodomeris cossmanni
Neovenericor austroplata
NHMUK 1967582; MACN-Pi 363, 364; MACN-In
22758
YPM IZ 033525; NMR 16389
MACN-Pi 5770, 5771, 355; PRI.66396, 66397,
66399, 66407
USNM.Mo 441626, 441627; PRI.59696, 63136
CPBA 13242, 13247, 13246, 1322 –13241, 13243–
13245, 13248–13273; 9326; MLP 1883
USNM.Mo 3373; NMR 19624, 20026
MACN-Pi 958; MNHN A25071, FJ07566; NMR
65304
NHMUK L27955-6; SGO.PI 469; MACN-Pi 357,
358, 359, 362, 365, 3578, 3583, 3602, 3611, 5768,
5769, 5781, 5782; CIRGEO-PI 2856, 2358, 2370,
2831, 2888; CPBA 6543, 8639, 8640, 9328, 9364,
9373; CPBA 15946–15949, 15952–15960; MLP
1837, 1839, 10507, 18334; MACN-Pi 5803;
PRI.66324, 66402, 66403, 66405, 66406, 66408–
66410, 66702
Maxwell 1969; Beu & Maxwell 1990
Maxwell 1969; Beu & Maxwell 1990
MACN-Pi 361b
MACN-Pi 2351, 2352; NMR 35717; Beu 2006
UF 116940; Stewart 1930; Heaslip 1968
MLP 15294a, 15294f, 1876, 1878, 1882, 1897,
4098, 4475, 5517, 7286, 9108, 9416, 9724, 9725,
9768, 10504, 10505; CPBA 6402, 6403, 6407,
8338; MACN-Pi 341–343, 5357, 5747, 5751–5753
MNHN A07711, FJ07559; MACN-Pi 3004
GMNH 46030; CPBA 328
MACN-Pi 371, 353, 2214, 6334; PRI.66711
MACN-Pi 3011; MNHN FR50693; NMR 80518M
Pouwer 2010
Stewart 1930; Gardner & Bowles 1939
MACN-Pi 2175, 2350, 2355; La Perna et al. 2017
MNHN FJ09318
USNM.Mo 327924; CPBA 14321–14327, 15764–
15772, 6576, 7459, 7460, 8738, 11042, 11043,
21861–21870; MACN-Pi 4718, 4719, 4805;

Neovenericor carrerensis
Neovenericor paranensis
Neovenericor ponderosa
Pacificor mulleri
Palaeocardita crenata
Palaeocardita stoecklini
Paraglans calcitrapoides
Pleuromeris decemcostata
Pleuromeris fueguina
Pleuromeris sulcolunularis
Pleuromeris tridentata
Pleuromeris zelandica
Powellina brookesi
Pteromeris perplana
Purpurocardia elegantoides
Purpurocardia leonensis
Purpurocardia purpurata
Rotundicardia diversidentata
Rotundicardia mariobrosorum
Rotundicardia rotunda
Scalaricardita camaronesia
Scalaricardita laciarina
Scalaricardita scalaris
Schizocardita cristata
Spissatella trailli
Venericardia imbricata
Venericor aposmithii
MHNSR.Pi 2785, 2789
MLP 22934, 14188, 14201, 22916–22949, 22950–
22968; MACN-Pi 2252, 2253, 2264–2267, 2269,
2315; CIRGEO-PI 128, 159–162, 2253, 3002
MACN-Pi 2534, 351, 352, 361a, 2632, 5358; PZ-
NRM Mo. 118089; MAS-Pi 4289; DMT-Pi 258,
505; MLP 7278
Beu & Maxwell 1990
Verastegui 1953
CPBA 599, 1108; MACN-Pi 3007
Hautmann 2001
MNHN FJ04948, FJ05266; NMR 64969; MACN-
Pi 2056
Gardner 1943
PZ-NHRM Mo. 117903; CIRGEO-PI 2382, 2877;
MACN-Pi 2619, 3747, 3748; CPBA 9349
MACN-Pi 367, 366, 368, 369, 370, 5773–5777;
PRI.66398
UF 189442; NMR 19837
NMR 71137
AK 70135; AUK 131098; NMR 16419
UF 49642; Gardner 1943
PRI.66394, 66395; CIRGEO-PI 2276; MLP 26811
CPBA 11651, 11652, 12921, 12926, 11653–11656,
12922–12925, 12927–12929, 13583–13585
MACN-In 28255; NMR 18866; Beu 2006
PRI.4283, 4284; Heaslip 1968
CPBA 17276; MACN-Pi 4180, 5758, 5759–5762,
5766
ANSP-IP 5267, 5268; PRI.642–644
MACN-Pi 349
MACN-Pi 2213
NMR 25647; Janssen & Moerdijk 2004
Körner 1937
TM 2859, 2860
GMNH 46023–46025; CPBA 330; MACN-Pi 960;
NMR 65308, MNHN FJ07562, FA25072
Gardner & Bowles 1939
Venericor bashiplata Gardner & Bowles 1939
Venericor planicosta CPBA 329; 3097; MACN-Pi 961; NMR 14599

Vimentum dilectum AMS C18124, C26099, C40720

Thecalia concamerata MACN-In 13778; NMR 16667, 82092

Additional references

Gardner, J. A. 1943. Mollusca from the Miocene and lower Pliocene of Virginia and

North Carolina. Part I. Pelecypoda. United States Geological Survey, Professional

Paper, 199-A, 1–169.

Pouwer, R. 2010. The identity of Isocrassina, Laevastarte and Ashtarotha (Mollusca,

Bivalvia, Astartidae) and their representatives from beaches and estuaries in The

Netherlands and Pliocene strata in Belgium. Cainozoic Research, 7(1–2), 27–67.

Appendix 2

Characters

Below, is provided a full and detailed discussion about characters defined and used in

the phylogenetic analyses.

General shell-outline and shell-shape (characters 2–22). (Fig. 2) Among these

characters was included shell-outline from juvenile and adult shells (c. 3–4), a very

important feature for Chavan (1969a), and the presence of anterior-posterior elongate

shells as in Cardita, Beguina or Carditamera species (c. 5) (Fig. 2D–F, H–I). The rest

of them described particularities of margins and its angles (c. 6–18). The pronounced

inclination of posterior-dorsal margin is a particular feature present in Miodomeris and

Pteromeris species, giving a distinctive outline to these genera (c. 6) (Fig. 2M, O). The

strong and prominent anterior-dorsal corner is present in Birkelundita turoniense (c. 11)

(Heinberg 1993) and the Triassic Schizocardita cristata has a unique radial ribs

extended from the angle between posterior and ventral margins (c. 16). Some carditids

(e.g. Glans, Centrocardita, Palaeocardita and Carditamera species, and Cardita

crassa) show a notch in posterior margin (c. 17) and others (especially

mytiliform/modioliform outlined carditids as Beguina or Thecalia species) have a

ventral gape for byssus protrusion in some ontogenetic stage (byssal gape) (Yonge

1969) (c. 18). Two sexual dimorphic characters included the presence of a ventral

incubatory chamber determined by the invagination of an interspace (as described

Yonge 1969) (c. 19) (Fig. 4D) and the existence of two morphotypes with different
convexity between males and females (Heaslip 1968; Pérez et al. 2017b) (c. 20). The

posterior margin of some carditid species can be dentate by the protrusion of inner

crenulations (Vermeij 2013) (c. 21) (Fig. 3J, L–M). Strong thickening of shells is

present for example in taxa traditionally included in planicostate lineage (e.g. Venericor

and Pacificor) and also in Palaeocardita species, crassatelid and astartid outgroup taxa

(Gardner & Bowles 1939; Verastegui 1953) (c. 22). La Perna et al. (2017) highlighted

this feature as a key character of Megacardita species. Average size reached by adults

was treated as continuous in some searches and discretized in others (c. 2) (comparisons

of different sizes are showed in Figure 2).

Umbones (characters 23–29). (Fig. 2) Orientation, relative position, convexity,

recurved condition and outline of umbones were considered (c. 23–28). In astartid,

crassatellid, Palaeocardita species and Schizocardita cristata the umbones are slightly

prosogyrate, and in Cardiocardita ajar it is minute and slightly medial orientated

(orthogyrate) (c. 23). The relative position of umbones varies from very anterior (e.g.

Cardita variegata) to nearly centrally-placed (e.g. Pleuromeris tridentata) (c. 24).

Distance between umbones can be minimum or there could be almost in contact (this

can be detected by an eroded surface in both valves, e.g. Venericor planicosta) (c. 29).

Escutcheon (characters 30–33). (Fig. 4A–C) This is an important difference between

outgroups and ingroup, and according to Chavan (1969a, b) a difference between

Crassatelloidea (Astartidae + Crassatellidae) and Carditoidea. Astartids and crassatellids

have escutcheon but neither carditid, except for Palaeocardita species and

Schizocardita cristata that present this structure (c. 30) – Chavan (1969a) did not

mention the presence of escutcheon in Palaeocardita and Schizocardita). Remaining

characters refer to relative size, definition and sculpture of escutcheon (c. 31–33).
Lunule (characters 34–42). (Fig. 4A–C) With the exception of Birkelundita

turoniense, Beguina semiorbiculata and Rotundicardia mariobrosorum, all carditids

present lunule (c. 34), but this feature can be more or less deep or defined (c. 35–36),

and it could has different sculpture, shape and concavity. The size of lunule is large in

some carditid (e.g. Pleuromeris, Venericor, Scalaricardita and some Cyclocardia

species) (c. 40).

Hinge (characters 43–88). (Fig. 3) Hinge is subdivided into three subgroups: Hinge

generalities (c. 43–45), Right Hinge (c. 46–72) and Left Hinge (c. 73–88). This is the

most important bivalve structure for many authors, and the historically more used in

systematic arrangements (Bernard 1895; Cox 1969). Carditidae has a particular type of

heterodont hinge named diagenodont (Verastegui 1953) or venericardioid (Heaslip

1968), which consists in two left cardinal teeth (anterior, 2, and posterior, 4b) and three

right cardinal teeth (3a, anterior, 3b, middle and 5, posterior) according to Bernard’s

(1895) nomenclature (Fig. 1B–C). Lateral teeth is not present in most carditids, and they

lack internal resilifer (this structure is present in Crassatellidae) (Fig. 3A–B). A

particular character noted in some carditid species is the presence of finely striations on

surface of cardinal teeth (c. 43) (La Perna et al. 2017) (Fig. 4E). The whole size of

hinge differs significantly from narrow (e.g. Vimentum) to very broad (e.g. Leuroactis)

(c. 44). On right valve, crassatellids and astartids present two laminar lateral anterior

teeth but Carditamera species shows pustular lateral anterior and posterior teeth (c. 46

and c. 72) (Fig. 3A–D, I–M). On left valve, a laminar anterior tooth is presents in

crassatellids and astartids but a pustular one in Thecalia concamerata, Carditamera

species, Glans trapezia and Centrocardita aculeata, while laminar posterior are visible

in crassatellids, Palaeocardita species and Schizocardita cristata and pustular posterior

in Carditamera species (c. 73–74, c. 87–88). Right anterior cardinal tooth is reduced in
most carditids, but in crassatellids, astartids, Palaeocardita species and Schizocardita

cristata is limited to small laminae adjacent to lunular margin, and in Cardites species is

absent (c. 48) (Fig. 3V). Right middle cardinal tooth is ventrally-divided that confer a

‘shark tooth-like’ shape in some species (e.g. Thecalia concamerata, Beguina

semiorbiculata, and in species of Cardita, Carditamera and Purpurocardia) (c. 53)

(Fig. 3E, H, J, K, M). This tooth is fused with anterior tooth in Birkelundita turoniense,

a possible neotenic state (c. 60) (Heinberg 1993). Right posterior tooth shows a

particular projection adjacent to nymph margin (Pérez et al. 2017a) in some large

carditids (Neovenericor, Venericor and Megacardita species) (c. 70) (Fig. 3R, U). On

left hinge, Glyptoactis hadra has a little knob, an structure not related to the hinge,

placed below ventral extreme of lunular margin (Heaslip 1968) (c. 85) (Fig. 4G). This

knob is also present in Thecalia concamerata, Powellina brookesi, Beguina

semiorbiculata and Cardita species. The remaining hinge characters describe

orientation, shape, relative size and height, and curvature of hinge cardinal teeth (c. 49–

68, c. 75–84). Ventral hinge edge is very variable in carditids, with concave, straight,

sinuous or inverted ‘V’-shaped outlines (c. 69).

Sculpture generalities (characters 89–91). (Fig. 4F, H–Q) Three characters described

the sculpture present and its dominance. Among crassatellid and astartids, comarginal

sculpture is dominant (Chavan 1969b) but in carditids radial sculpture is more

important. In Miodomeris cossmanni radial sculpture is very faint (c. 91).

Comarginal sculpture (characters 92–96). (Fig. 4H) This type of sculpture is absent

in most carditids but astartids have two type of comarginal sculpture: comarginal lines

and undulations (Pouwer 2010) (wavy concentric ridges superimposed to comarginal

lines) (c. 95–96).

Radial sculpture (characters 1, 97–119) (Fig. 4). This type of sculpture is a key

feature among carditids according to many authors and is established by strong radial

ribs (Harris 1919; Verastegui 1953; Chavan 1969a; Maxwell 1969; Morton et al. 1998;

Coan et al. 2000). Ribs can become obsolete or vanish in gerontic stages of valves, a

character-state present in some Cyclocardia or Venericor species, and in Leuroactis

pilsbryi (c. 106) (Fig. 4I). These ribs can be high or low (c. 105) and become weak in

different section of valve (c. 107). They can have different cross-sections (rounded,

rectangular, triangular, etc.) (c. 103–104). Some Carditamera species have radial ribs

with posteriorly inclined cross-section (c. 108) (Fig. 4J) and Kolmeris tehuelchana have

stacked ribs on flanks (c. 110) (Fig. 4F). Radial ribs are from two different types: entire

radial ribs or tripartite radial ribs (c. 97) (Fig. 4K–N). The tripartite radial rib has a

central cord flanked by two paracostal ribs on each side (Stewart 1930; Rutsch &

Schenck 1941; Heaslip 1968) and was also named terraced rib (Harris 1919) or

fasciculate (Maxwell 1969). Paracostal ribs show variable development with respect to

central cord (c. 100–102). Instead paracostal ribs, Cardita variegata shows serrations on

rib flanks (c. 109). Interspaces can have different sections (‘V’-shaped, ‘U’-shaped,

rectangular), be restricted to a fine line (c. 103–104), or be extremely reduced (c. 111).

Palaeocardita species and Schizocardita cristata have posterior-ventral prosocline

radial ribs (c. 118). The number of radial ribs remains virtually fixed throughout

lifespan of an individual, and this number shows little intraspecific variation. To test the

correspondence of number of ribs with specific identity, an analysis of Generalized

Lineal Model (GLM) was carried out using R (R Development Core Team 2017). This

analysis is described in Appendix 4. After results, the existence of a significant

correspondence between number of radial ribs and species identity can be determined.
Number of radial ribs was considered as continuous or discrete character in different

search approaches (c. 1).

Comarginal nodes (characters 119–128). (Fig. 4M–Q) Among carditids, radial ribs

are usually covered by nodes with a comarginal arrangement. The terms used by Harris

(1919) to describe different types of ribs (i.e. ‘funginate’, ‘funiculate’ or ‘flabellate’)

actually refers to node morphologies instead ribs. Nodes can be circular (Fig. 4M),

rectangular (Fig. 4Q), scales, funginate (resembling bracket fungi) (Fig. 4O) or with a

saw-like aspect (Fig. 4P) (Heaslip 1968; Pérez & del Río 2017b) (c. 120). Node

development changes with ontogeny, some taxa have nodes only on juvenile section of

valves (e.g. Venericor species) (c. 125) and other have nodes only around lunule

(Neovenericor paranensis and N. ponderosa) (c. 126). In Bathycardita raouli this

lunular nodes are thorn-like (c. 127). Fasciculicardia species have scaly nodes with

tubular extremes in some regions of valve (c. 128).

Internal morphology (characters 130–137). (Fig. 3) Internal morphology of carditids

includes muscle scars and pallial line. The first is generally involved in palaeoecological

interpretations (Stanley 1970, 1972; Heinberg 1993), but these structures received little

attention on descriptions of carditid species. Crassatellids, Cyclocardia and Venericor

species have a flat zone between anterior adductor muscle scar and anterior margin (c.

130) and this scar is displaced towards hinge margin in crassatellids, astartids,

Schizocardita cristata and Palaeocardita species (c. 133) (Fig. 3A–D). Muscle scars are

exceptionally large in some species (e.g. Leuroactis pilsbryi, Venericor planicosta,

Neovenericor species) (c. 132) and occasionally have asymmetrical sizes (c. 134) (a

situation always presents in modioliform/mytiliform carditids) (Fig. 3F, H). Relative

position of pallial line and its point of contact with anterior adductor muscle scar are

variable (c. 135–136).

Ventral margin crenulations (characters 138–144). (Fig. 3) Inner crenulations are a

hallmark of Carditidae (Morton et al. 1998; Coan et al. 2000) and they are also

frequently present in Astartidae. Crassatellidae species may not have crenulations (c.

138). Crenulations of carditids correspond to interspaces, but in astartids and

crassatellids this is not true (c. 139), and both structures are not homologous (Fig. 3B,

D). Some Carditamera species, Centrocardita aculeata, Glans trapezia and Cardita

species have crenulations smoothly prolonged towards valve inside (c. 142) (Fig. 3F, H,

L). Crenulations can be extended by whole valve margin or vanish at some point of

posterior margin (c. 143). In some carditid species, extreme tips of radial ribs poke out

among crenulations in internal view (c. 144) (Fig. 4R).

List of characters used in the phylogenetic analysis, grouped by characters-group

mentioned in the text. Continuous characters of cont+disc approach are mentioned at the

beginning and ordered characters are indicated.

Continuous characters

1. Number of radial ribs.

2. Mean size of adult shells (Large)

Discrete characters

1. Number of radial ribs. 0= without radial ribs, 1= up to 15, 2= from 16 to 20, 3= from
21 to 25, 4= from 26 to 30, 5= from 31 to 37, 6= more than 37 [ordered]

General shell-outline and shell-shape

2. Mean size of adult shells (Large in mm). 0= up to 12, 1= from 12 to 22, 2= from 22
to 32, 3= from 32 to 60, 4= from 60 to 90, 5= more than 90 [ordered]
3. Outline in adult shells. 0= mytiliform/modioliform, 1=obtuse triangle, 2= quadrate,
3=subtrigonal, 4= subrectangular, 5= subrounded (rotund for Heaslip, 1968), 6=
rectangular

4. Outline in juvenile shells. 0= subrectangular, 1= subquadrate, 2= subtrigonal, 3=

subrounded

5. Elongation of outline in anterior-posterior direction. 0= present, 1= absent [ordered]

6. Posterior-dorsal margin inclination. 0= absent, 1= slight inclination (as in Coripia),

2= pronounced inclination (as in Miodomeris and Pteromeris) [ordered]

7. Posterior-ventral margin projection. 0=strongly projected, 1=absent, 2=slight

projected

8. Anterior-ventral margin projection. 0= absent, 1= present

9. Posterior margin outline. 0= rounded, 1=acuminated (acute with marked angle),

2=slightly truncated, 3= vertically truncated

10. Anterior margin outline. 0= rounded, 1= straight

11. Anterior-dorsal corner. 0= softly marked, 1=prominent (as in Birkelundita)

12. Angle between posterior and dorsal margins. 0= softly marked, 1=strongly marked,
2= absent

13. Dorsal margin development. 0= curved and descended, 1= descended, 2=regular, 3=

straight

14. Dorsal margin elongation. 0= absent, 1= present

15. Angle between posterior and ventral margins. 0= absent (rounded angle), 1=present,
2= strongly marked

16. Radial rib projected between posterior and ventral margins angle. 0= absent, 1=
present (as in Schizocardita)

17. Notch in posterior margin. 0= absent, 1=present (as in Glans)

18. Byssal notch in ventral margin. 0= present (as in Beguina), 1=absent

19. Presence of an incubatory chamber in females. 0= absent, 1=present

20. Presence of two morphotypes with different convexity (possible sexual

dimorphism). 0= absent, 1= present

21. Serrated posterior margin. 0= absent, 1=present (as in Glans)

22. Shell thickness. 0=slight, 1=strong

Umbones

23. Umbones direction. 0=prosogyrous, 1=slightly prosogyrous, 2=orthogyrous

[ordered]

24. Umbones placement. 0= very anteriorly placed, 1= at anterior third, 2= subcentrally

placed [ordered]

25. Umbonal area convexity. 0= regular, 1= very low (nearly flat shells), 2= very high

26. Degree of recurve umbones. 0= slightly recurved, 1= recurved, 2=strongly recurved

27. Umbones outline in external view. 0= pointed, 1= slightly rounded, 2= strongly

rounded

28. Umbones outline in internal view. 0= not visible umbones, 1= visible umbones

29. Umbones distance. 0= in contact, 1= minimum spacing [ordered]

Escutcheon

30. Presence of escutcheon. 0= absent, 1= present

31. Escutcheon width. 0= narrow, 1=wide 2= escutcheon absent

32. Escutcheon demarcation. 0= slightly marked, 1= strongly marked, 2= escutcheon

absent

33. Sculpture of escutcheon. 0= absent, 1= present, 2= escutcheon absent

Lunule

34. Presence of lunule. 0= present, 1= absent

35. Lunule depth. 0= shallow, 1=very deep

36. Lunule demarcation. 0= strongly marked by a groove, 1=slightly marked

37. Sculpture of lunule. 0= smooth, 1=only comarginal lines

38. Lunule outline. 0= subovate, 1=subrounded, 2=as an inverted tear

39. Lunule convexity. 0= flat, 1= concave, 2= convex, 3= sinuous

40. Lunule size. 0= small, 1=regular, 2= large

41. Lunule symmetry. 0= symmetric lunule, 1= right lunule larger

42. Lunule projection. 0= absent, 1= right lunule edge projected as a sheet

Hinge generalities

43. Striated teeth. 0= absent, 1= present

44. Hinge size (Height). 0= very reduced and short, 1= regular, 2= extended towards
venter

45. Internal resiliferum. 0= present, 1= absent

Right hinge

46. Anterior lateral teeth laminar. 0= absent, 1= two teeth

47. Anterior lateral teeth pustular. 0= absent, 1=present

48. Presence of anterior tooth (3a). 0= very reduced, 1= present, 2= absent, 3= limited to
a small sheet adjacent to lunular margin

49. Orientation of anterior tooth (3a). 0= vertical, 1= anterior, 2= posterior, 3= absent

50. Prominence of anterior tooth (3a). 0= little prominent, 1= conspicuous, 2= very

developed, 3= absent, 4= very little prominent

51. Shape of anterior tooth (3a). 0= elongate rectangular, 1= pustular, 2= curved

rectangular, 3= absent

52. Apex of anterior tooth (3a). 0= resting on anterior side of middle tooth, 1= not
resting on anterior side of middle tooth

53. Division of anterior tooth (3a). 0= divided (shark tooth-like), 1= not divided

54. Posterior-ventral corner of middle tooth (3b) extension. 0= absent, 1=present (as in
Beguina and Cardita)

55. Shape of middle tooth (3b). 0= triangular with broad base, 1= triangular with narrow
base, 2= scimitar-like

56. Curvature of middle tooth (3b). 0= anterior side concave, 1= anterior side convex,
2= anterior side straight

57. Orientation of middle tooth (3b) (in degrees). 0= 0, 1= from 0 to 30, 2= from 30 to
35, 3= from 35 to 45, 4= from 45 to 90, 5= 90 [ordered]

58. Elevation of middle tooth (3b). 0= constant, 1= higher towards dorsal, 2= higher
towards ventral, 3= higher at centre
59. Apex of middle tooth (3b). 0= not reaching the dorsal extreme of hinge, 1= reaching
the dorsal extreme of hinge

60. Fusion between anterior tooth (3a) and middle tooth (3b). 0= absent, 1= present (as
in Birkelundita)

61. Shape of posterior tooth (5). 0= straight, 1= curved

62. Orientation of posterior tooth (5) (in degrees). 0= more than 35, 1= from 20 to 35,
2= from 0 to 20 [ordered]

63. Elevation of posterior tooth (5). 0= lower than other teeth, 1= equals to other teeth

64. Apex of posterior tooth (5). 0= resting on posterior side of middle tooth, 1= not
resting on posterior side of middle tooth

65. Spacing between ventral extreme of posterior tooth (5) and hinge margin. 0=
present, 1= absent

66. Ventral extreme of posterior tooth (5). 0= protruding from ventral edge of hinge,
1=not protruding

67. Extension of posterior tooth (5). 0= not reaching the apex of middle tooth, 1=
reaching the apex of middle tooth, 2= reaching the middle point of middle tooth

68. Higher tooth in the hinge. 0= 3b, 1=3a, 3= 5

69. Ventral edge of hinge. 0= invaginated to dorsal, 1= concave, 2= straight, 3= sinuous,

4= wide and inverted ‘V’-shaped, 5= irregular

70. Projection of nymph margin. 0=absent, 1= present, 2= large [ordered]

71. Posterior lateral teeth laminar. 0= present, 1= absent

72. Posterior lateral teeth pustular. 0= present, 1=absent

Left hinge

73. Anterior lateral teeth laminar. 0= present, 1= absent

74. Anterior lateral teeth pustular. 0= absent, 1=present

75. Orientation of anterior tooth (2) (in degrees). 0= 0, 1= from 0 to 50, 2= from 50 to
80, 3= from 80 to 90, 4= more than 90 [ordered]

76. Size relations of anterior tooth (2). 0= higher than wide, 2= as high as wide, 3=
wider than high
77. Elevation of anterior tooth (2). 0= higher towards ventral, 1= higher towards dorsal,
2= higher at centre, 3= constant

78. Size of anterior tooth (2). 0= large, 1= very reduced

79. Shape of anterior tooth (2). 0= subquadrate, 1=subrectangular, 2= subtriangular

80. Lateral profile of anterior tooth (2). 0= hook-like outline, 1= without hook-like
outline

81. Orientation of posterior tooth (4b) (in degrees). 0= from 0 to 20, 1= from 20 to 30,
2= from 30 to 35, 3= from 35 to 45, 4= more than 45

82. Elevation of posterior tooth (4b). 0= higher towards ventral, 1= higher towards
dorsal, 2= higher at centre

83. Curvature of posterior tooth (4b). 0= slightly curved, 1= strongly curved, 2= straight

84. Higher tooth in the hinge. 0= 4b, 1=2, 2=equal height

85. Knob placed below ventral extreme of lunular margin. 0= present, 1= absent

86. Lateral profile of posterior tooth (4b). 0= hook-like outline, 1= without hook-like
outline

87. Posterior lateral teeth laminar. 0= present, 1= absent

88. Posterior lateral teeth pustular. 0= absent, 1=present

Sculpture generalities

89. Comarginal sculpture. 0= present, 1= absent

90. Radial sculpture. 0= absent, 1= present

91. Dominant sculpture. 0= comarginal, 1= radial

Comarginal sculpture

92. Comarginal lines. 0= present, 1= absent

93. Demarcation of comarginal lines. 0= more marked anteriorly, 1= equally marked in

whole valve, 2= comarginal lines absent

94. Elevation of comarginal lines. 0= low, 1= elevated, 2= comarginal lines absent

95. Undulations. 0= present, 1= absent

96. Undulations in juvenile shell. 0= limited to juvenile shell, 1= not limited, 2=
undulations absent

Radial sculpture

97. Type of radial sculpture. 0= Entire radial ribs, 1= fine radial ribs, 2= radial ribs
absent

98. Radial ribs width in adult shells. 0= very wide radial ribs in adult shells, 1= not very
wide radial ribs in adult shells, 2= radial ribs absent

99. Radial ribs width along valve surface. 0= wider towards posterior, 1= constant width
along whole valve surface, 2= wider towards anterior, 3= radial ribs absent

100. Presence of paracostal ribs. 0= absent, 1= present, 2= radial ribs absent

101. Paracostal ribs development. 0= absent, 1= present in all radial ribs, 2= less
developed in anterior radial ribs, 3= more developed in anterior radial ribs, 4= radial
ribs absent

102. Paracostal ribs proportions towards centre of valves. 0= absent, 1= much smaller
than central rib, 2= more than middle width of central rib, 3= radial ribs absent

103. Cross-section of radial ribs in adult shell. 0= subovate, 1= subrectangular, 2=

subtriangular, 3= radial ribs absent

104. Cross-section of radial ribs in juvenile shell. 0= subovate, 1= subrectangular, 2=

subtriangular, 3= radial ribs absent

105. Height of radial ribs. 0= very low, 1= slightly elevated, 2= elevated, 3= high, 4=
radial ribs absent [ordered]

106. Radial ribs in gerontic valve surface. 0= persistent, 1= obsolete, 2= radial ribs
absent

107. Smoothed radial ribs in some region of valve surface. 0= never smoothed, 1=
anteriorly smoothed, 2= posteriorly smoothed, 3= radial ribs absent

108. Radial ribs posteriorly inclined in cross-section. 0= absent, 1= present (as in

Carditamera radiata)

109. Serrated flanks of radial ribs. 0= absent, 1= present (as in Cardita variegata)

110. Stacked radial ribs on flanks. 0= absent, 1= present (as in Kolmeris tehuelchana)

111. Interspaces. 0= present, 1= absent, 2= radial ribs absent

112. Interspaces width. 0= narrower than radial ribs, 1= as wide as radial ribs, 2= wider
than radial ribs, 3= interspaces absent

113. Cross-section of interspaces in adult shell. 0= V-shaped, 1= U-shaped, 2=

subrectangular, 3= inverted trapezoid, 4= fine line, 5= interspaces absent

114. Cross-section of interspaces in juvenile shell. 0= V-shaped, 1= U-shaped, 2=

subrectangular, 3= inverted trapezoid, 4= fine line, 5= interspaces absent

115. Seventh or eighth anterior radial rib more marked than the remaining. 0= absent,
1= present, 2= radial ribs absent

116. Posterior area. 0= defined by thicker radial ribs than the remaining, 1= defined by
less thick radial ribs than the remaining, 2= defined by a convexity change, 3= absent

117. Fifth or Sixth posterior radial rib. 0= stronger than previous ones, 1= equally
developed than previous ones, 3= radial ribs absent

118. Posterior-ventral prosocline radial ribs. 0= absent, 1= present, 2= radial ribs absent

119. Presence of umbonal carinae towards posterior margin. 0= absent, 1= present

Comarginal nodes

120. Presence of comarginal nodes. 0= absent, 1= present

121. Type of comarginal nodes in adult shell. 0= scales, 1= smooth radial ribs, 2=
subrectangular nodes, 3= reticulate valve surface, 4= subrounded nodes, 5=
funginated nodes, 6= saw-like nodes, 7= comarginal nodes absent

122. Size of comarginal nodes. 0= large, 1= small, 2= comarginal nodes absent

123. Separation between comarginal nodes along radial rib. 0= regular, 1= very close,
2= without separation, 3= very spaced, 4= absent, 5= comarginal nodes absent

124. Cancelled aspect given by comarginal nodes and radial sculpture. 0= absent, 1=
present, 2= comarginal nodes absent, 3= incipient, 4=smooth radial ribs

125. Development of comarginal nodes in post-juvenile shell. 0= equally developed in

whole valve surface, 1= posteriorly marked, 2= anteriorly marked, 3= comarginal
nodes absent

126. Presence of comarginal nodes in juvenile shell. 0= present, 1= absent, 2=

comarginal nodes absent

127. Comarginal nodes more marked around lunule. 0= absent, 1= present, 2=

comarginal nodes absent
128. Spiny nodes in juvenile shell. 0= absent, 1= present in all umbonal region, 2= more
developed towards anterior side of umbonal region, 3= comarginal nodes absent

129. Tubular nodes in posterior surface of valve. 0= absent, 1= present, 2= comarginal

nodes absent

Internal morphology

130. Flat zone between anterior adductor muscle scar and anterior margin. 0= absent, 1=
present

131. Prominence of adductor muscle scars. 0= slightly marked, 1= marked, 2= strongly

marked

132. Size of adductor muscle scars. 0= regular, 1= very large

133. Placement of anterior adductor muscle scar. 0= displaced towards hinge margin, 1=
not displaced

134. Size relation between both adductor muscle scars. 0= posterior larger, 1= anterior
larger, 2= equal

135. Height of pallial line. 0= less than a fifth of total valve height, 1= a fifth of total
valve height, 2= a quarter of total valve height [ordered]

136. Intersection between pallial line and anterior adductor muscle scar. 0= at middle of
anterior adductor muscle scar, 1= at posterior side of anterior adductor muscle scar

137. Shape of pallial line. 0= inclined towards posterior margin, 1= uniform curve

Ventral margin crenulations

138. Presence of ventral margin crenulations. 0= absent, 1= present

139. Correspondence of ventral margin crenulations. 0= not corresponding to

interspaces, 1= corresponding to interspaces

140. Shape of crenulations. 0= subrectangular, 1= subtriangular, 2= crenulations absent

141. Extreme tip of crenulations. 0= truncated, 1= rounded, 2= crenulations absent

142. Extensions of crenulations toward valve inside. 0= gentle extended towards valve
inside, 1= limited to ventral edge, 2= crenulations absent

143. Extension of crenulations on margins. 0= to dorsal side of posterior adductor

muscle scar, 1= reaching posterior adductor muscle scar, 2= below posterior
 adductor muscle scar, 3= to point of contact between posterior and ventral margins,
4= crenulations absent

144. Radial ribs between crenulations on internal view. 0= extreme tips of radial ribs
not visible, 1= extreme tips of radial ribs poking out on internal view, 2=
crenulations absent

Additional references

Chavan, A. 1969a. Superfamily Carditacea Fleming, 1920. Pp. 543–561 in R. C. Moore

(ed.) Treatise on Invertebrate Paleontology, Part N, Mollusca 6, Bivalvia, volume 2.

Geological Society of America & The University of Kansas, Lawrence.

Chavan, A. 1969b. Superfamily Crassatellacea Férussac, 1822. Pp. 562–582 in R. C.

Moore, & C. Teichert (eds). Treatise on Invertebrate Paleontology. Part N2,

Mollusca 6. Bivalvia, volume 2. The Geological Society of America & The

University of Kansas, Lawrence.

Pouwer, R. 2010. The identity of Isocrassina, Laevastarte and Ashtarotha (Mollusca,

Bivalvia, Astartidae) and their representatives from beaches and estuaries in The

Netherlands and Pliocene strata in Belgium. Cainozoic Research, 7(1–2), 27–67.

Appendix 3A

Character codifications for each terminal. Question mark indicates missing data and en-
dash indicates not applicable data. Polymorphic codifications are indicated with the
following notation: [state A state B]. Codification of continuous characters is listed at
first and discrete characters at second. An alternative visualization of this data is
provided in Appendix 3B (text file ready to run in TNT or similar software).

Continuous characters codifications

1. Number of radial ribs.

2. Mean size of adult shells (Large)

Crassatella ponderosa 0 62.57

Spissatella trailli 0 38.86

Astarte sulcata 0 17.87

Laevastarte basteroti 0 33.17

Palaeocardita stoecklini 22-23 21.88

Palaeocardita crenata 22-24 23

Schizocardita cristata 35 36.71

Birkelundita turoniense 34 8.7

Thecalia concamerata 24-25 12.58

Powellina brookesi 21 13.32

Beguina semiorbiculata 46 66.75

Cardita variegata 25 25

Cardita crassa 18-21 24.01

Claibornicardia paleopatagonica 30-33 37.15

Claibornicardia alticostata 30-33 44.94

Claibornicardia acuticostata 30-33 33.13

Rotundicardia mariobrosorum 25-27 28.97

Rotundicardia rotunda 25-26 24.92

Rotundicardia diversidentata 25-26 19.4

Kalelia burmeisteri 25 43.52

Kalelia pectuncularis 26-28 78.89

Kalelia multicostata 28-30 50.49

Glyptoactis hadra 19-20 33.6

Baluchicardia beaumonti 20-22 26.5

Venericardia imbricata 28-30 31.01

Arcturellina asperula 28-30 13.6

Fasciculicardia subintermedia 28-32 68.21

Fasciculicardia acanthodes 24-26 51.58

Fasciculicardia sp. 25 34.6

Purpurocardia purpurata 22-26 27.17

Purpurocardia elegantoides 22-23 14.64

Purpurocardia leonensis 24-26 29.05

Cardites antiquatus 21-22 26.8

Cardites partschii 22-23 18.23

Cardites feruglioi 28-29 41.85

Cardiocardita ajar 23-26 20

Coripia unidentata 25-26 5.26

Miodomeris cossmanni 18-20 3.04

Scalaricardita scalaris 23-25 12

Scalaricardita camaronesia 23-25 10.36

Scalaricardita laciarina 25-27 14.42

Vimentum dilectum 27-29 8

Pleuromeris tridentata 17-18 6.15

Pleuromeris decemcostata 11-12 5.84

Pleuromeris fueguina 18-21 15.34

Pleuromeris sulcolunularis 18-20 16.6

Pleuromeris zelandica 16-19 8

Pteromeris perplana 10-11 3.5

Cyclocardia borealis 20-23 40

Cyclocardia awamoensis 20-24 17.95

Cyclocardia ventricosa 20-22 15.28

Cyclocardia cannada 23-27 15.03

Cyclocardia mesembria 21-22 12.2

Cyclocardia dalek 21-24 14.74

Cyclocardia compressa 15-17 14.33

Cyclocardia nortensis 21-22 28.66

Darwinicardia patagonica 23-26 30.33

Darwinicardia angusticostata 22-23 19.72

Venericor planicosta 31-34 69.64

Venericor aposmithii 31-33 93

Venericor bashiplata 28-30 63

Bathycardita raouli 14-16 34

Megacardita jouanetti 19-21 53.28

Neovenericor paranensis 23-25 73.6

Neovenericor ponderosa 25-28 73.97

Neovenericor austroplata 21-22 99.31

Neovenericor carrerensis 24-26 87.15

Leuroactis pilsbryi 30-33 93

Pacificor mulleri 30-34 63

Glans trapezia 18-21 10.48

Centrocardita aculeata 20-22 24.6

Carditamera arata 20 41.16

Carditamera floridana 17-19 26.97

Carditamera affinis 17-18 62.27

Carditamera gracilis 17-19 25.24

Paraglans calcitrapoides 23-24 7.76

Kolmeris tehuelchana 13-15 12.57

Discrete characters codifications

Crassatella ponderosa

0441101030003010010001122000111110100032110201032400100153000200012050
0100403020402010000000000022324333423000235522220072523223212102000100
0132

Spissatella trailli

0341002030003110010001112000111110100032110201032400101253000200012050
0100403010402010000000000022324333423000235522220072523223212102000002
2242

Astarte sulcata

0122101000003000010101121000110000010012110111032401100243100001010030
0100403021400111100000100122324333423000235523220072523223201102101100
0112

Laevastarte basteroti

0322101000003000010101121000110100010012110111032401100243100001010030
0100403021400111100000100122324333423000235523220072523223201102101100
0112

Palaeocardita stoecklini

3141002030013100110001112021110000000012110110032401100253100200012050
0110403021402111000110111201100022200000010002010070523223202102000110
0111

Palaeocardita crenata
3241002030013100110001112021110000000012110110032401100253100200012050
0110403021402111000110111201100022200000011002010070523223202102000110
0111

Schizocardita cristata

5341002030013121110001112021110000000022110110032401100253100200012050
0110403021402111000110101201100022200000001002011070523223202102000110
0111

Birkelundita turoniense

532?100001101000010000000011102221-------
1010000011010000110001000000111010010100001110111122120010000010000001
10000001000000000000100001100010

Thecalia concamerata

3100000000100100001000000011102220000000000010002011010000100001000000
1111400011000001101111221200000000100000011000000100000000000010000110
0000

Powellina brookesi

3100000000100100001000000011102220100000000010002011010000100001000000
1110400011000001101111221200000000100000001000000100000000000010000110
0001

Beguina semiorbiculata

6400000000100100000000000001102221-------
0010000001010000100101100000111000000100000110111122121000000000000000
00031001401020000000100001100010

Cardita variegata

3200000000100100000000000011102220000000000010000011010000100001000000
1110100001000001101111221200000000100010000000000100000000000010000110
0000

Cardita crassa

2200000000100100100000000011102220000000000010000011010010100001000000
1110100001000001101111221200000000100000000000000100000000000010000110
0000

Claibornicardia paleopatagonica
5341002010002010010100010111102220011120111110001001100001101201010030
1110111021210011101111221201111222201000004401000140112000001011101111
0100

Claibornicardia alticostata

5341002020002010010100010111102220011120111110001001100001101201011030
1110021021210011101111221201111222201000004401000140112000001011101110
0100

Claibornicardia acuticostata

5321102030002010010000020111102220010120110110001001100001101201011030
1110021021210011101111221201111222201000004401000140112000001011101110
0100

Rotundicardia mariobrosorum

42[25]1101000002010010000010011102221-------
1110000001100001100201010010111002102121001110111122120110001220000001
01020001501120000010111011110101

Rotundicardia rotunda

4251101000002010010000010011102220011120111110000001100001101201010010
1110021021212011101111221201100012200000021102000150112000001011101111
0101

Rotundicardia diversidentata

4151101000002000010100010111102220011120111110001001100001101201010010
1110021021210011101111221201100012200000011102000150112000001011101111
0101

Kalelia burmeisteri

3352102030011000010001020111102220111121001110001011100011101201011020
1110001021110011101111221200213112102000001101000140012000001011101110
0100

Kalelia pectuncularis

4452102030011000010001020011102220111101001110011011100021101201011021
1110001021110011101111221200213112102000001101000120012000001011101110
0100

Kalelia multicostata
4352102030011000010001020011102220111101001110011011100021101201011021
1110001021210011101111221200213112102000001101000140012000001011101110
0100

Glyptoactis hadra

2341002020002010010000012221102220111120111110002011100001101201010040
1110011121111001101111221201111112201000001101000120112000001011101110
0100

Baluchicardia beaumonti

3221101020002000010001012211102220010120110110001011100001101201011000
1110021021210011101111221201111222200000000001000160010000001011101110
0100

Venericardia imbricate

4221101030012000010000020011102220011120000110001001101011101201011020
1110001021112011101111221201111112200000011102100120010000001011101111
1101

Arcturellina asperula

4121101030012000010000020011102220010022000010002001100011101201011020
1110311121112011101111221201111112200000011102100120010000001011101111
1101

Fasciculicardia subintermedia

4441102020002000010000010021102220010120000110002011100001101201010010
1110211121111011101111221201111122202000001101000140112000101011101110
0110

Fasciculicardia acanthodes

3341102020002000010000010021102220010120000110002011100001101201010011
1110211121111011101111221201111122202000001101000140112000101011101110
0110

Fasciculicardia sp.

33411020200020000100000100211022200101200011100????????????0???????0??111
0113121110011101111221201100?222?2000001101000160112000101011???1100?1?

Purpurocardia purpurata

3241001000002000010000010021102220110022000110010101000031101101011030
1110301021210211101111221200111112102000001101000120110000000011211110
0110
Purpurocardia elegantoides

3141001000002000010000010011102220110021000110010101000031101101011030
1110301021210211101111221200111112102000001101100120110000000011111110
0100

Purpurocardia leonensis

3241001000002000010000010011102220110022000110011101000031101101011030
1110101021210211101111221200111112202000001102000120110000000011111110
0110

Cardites antiquatus

3242002020002010010000012211102220010120001110023331100011101201011031
1110113021200011101111221201200011202000000001000120012000001011101110
0110

Cardites partschii

3142002020002010010000012211102220010120001110023331100011101201011030
1110113021200011101111221201200011202000000001000120012000001011101110
0100

Cardites feruglioi

4342002020002010010000012211102220010120001110023331100011101201011030
1110112021200011101111221201200011202000000001000120012000001011101110
0120

Cardiocardita ajar

3152001010003010010010220001102220010120000010012001100231101201011020
1110303021210011101111221201111132200000011102100120010000000011001110
0101

Coripia unidentata

40[13]31110200010000100000200011022200110021101100020011002311000010110
1011103030212220111011112212010000010000000000011001211220000000112011
110110

Miodomeris cossmanni

2012121000000000010000021001102220101002110110002001100031100001011010
1110301021300011100100101201100001000000001003100110?40000000111101110
0110

Scalaricardita scalaris
3033101020021000010000020001102220011002110110002001100231100001011010
1110303021222011101111221201000001000000000003100121122000000011201111
0110

Scalaricardita camaronesia

3033101020021000010000010001102220011002110110002001100231100001011010
1110403021222011101111221201000001000000000003100121122000000011201111
0110

Scalaricardita laciarina

4133101120021000010000020001102220011001110110002001100231100001011010
1110303021222011101111221201000001000000000003100121122000000011201111
0110

Vimentum dilectum

4043101020022000010000020011102220011002110110002001100231100001011010
1110303021221011101111221201000001000000000001100121122000000011101111
0110

Pleuromeris tridentata

2032101010001000010000022001102220101002110110002001100231100001011010
1110401011202011101111221201100022200000000003100120010000000011101111
0110

Pleuromeris decemcostata

1032101010001000010000022001102220101002110110002001100231100001011010
1110401021202011101111221201100022200000000003100120010000000011101110
0110

Pleuromeris fueguina

2132101000001000010000022101102220101002110110000001100231101001011010
1110301021200011101111221201100022200000010003100120110000000011101110
0110

Pleuromeris sulcolunularis

2132101000001000010000022101102220101012110110000001100231101001011010
1110301021200011101111221201100022200000010003100120110000000011101110
0110

Pleuromeris zelandica
2032101010001000010000022001102220101002110110002001100231100001011010
1110401021202011101111221201100022200000010003100120110000000011101110
0110

Pteromeris perplana

1012121000000000010000021001102220101002110110000001101231100001011010
1110301021300011101111221201100011000000003403100110?40000000011101110
0110

Cyclocardia borealis

33531010000220000100000100111022201010020001100010111002311001010110[13
]01110103021221011101111221200100002012000000001100110?0200001001110111
10110

Cyclocardia awamoensis

31531010000220000100000100111022201011000001100010011012211001010110[13
]0111010301122001110111122120110000001200000000110011010200001001110111
10110

Cyclocardia ventricosa

3153101000022000010000010011102220101102000110001011100231100101011030
1110103021222011101111221200100002012000000001100120102000010011101111
0110

Cyclocardia cannada

3153101000022000010000010011102220001002000110002011100231100101011010
11104030212200111011112212001000020120000000011001[14]01020000100111011
100111

Cyclocardia mesembria

3153101020022000010000010011102220001000000110001011100231100101011010
1110?????1220011101111221200100002012000000001100110?020000100111011110
110

Cyclocardia dalek

3153101000022000010000010011102220001002000110002011100231100101011010
11104030212200111011112212001000020120000020011001[24]01020000100111011
100111

Cyclocardia compressa
2153101000022000010000010011102220101002000110001011100231100101011010
1110103021221011101111221200100002012000000001100110?02000010011101111
0110

Cyclocardia nortensis

3253101000022000010000010011102220101100000110001001101221100101011030
11101030112200111011112212011000000120000020011001[12]01020000100111011
100110

Darwinicardia patagonica

32[35]310[12]02000200001010001[02]01110222000012100111000002110002110120
1011040111010101122001110111122120111212220000001000110014011200000001
11011100101

Darwinicardia angusticostata

3153101020002000010000010111102220010120001110000021100021101201011040
1110111021220011101111221201112122200000010001100140112000000011101110
0111

Venericor planicosta

54311020[02]00010000100010122210022200000221102100111011020211010010110
321110101011221011101111221201100011012000004201100110?401000121111001
100120

Venericor aposmithii

55311020[12]00010000100010122210022200002021102100111011020211010010110
311110101011321011101111221201100011012000004201100110?401000121111001
100120

Venericor bashiplata

44311020[02]00010000100010122210022200002021102100111011020311010010110
321110101011321011101111221201100011012000003201100110?401000121111001
100120

Bathycardita raouli

1361002010023010010001000011102220100021111110010001100231101101011030
1110200021100011101111221201200002012000012201100110?40001012110100110
0120

Megacardita jouanetti
2361002010023010010001002121102220000021111110001001100011101101011031
1110100011100011101111221201200012012000004201100110?40000002110100110
0120

Neovenericor paranensis

3441002010021010010001002211102220000020111110001001100011101101011031
1110110001121011101111221201200011102000013201100110?40010002111100110
0120

Neovenericor ponderosa

4441002010021010010001002211102220000021111110001001100011101101011031
1110110001121011101111221201200011102000013201100110?40010002111100110
0120

Neovenericor austroplata

3531002010021010010001002211102220000020111110001001100011101101011031
1110101001121011101111221201200011102000013201100110?40100002111100110
0120

Neovenericor carrerensis

3441002010021010010001002221102220000020110110001001100011101101011031
1110101001121011101111221201200010102000013201100110?40100002111100110
0120

Leuroactis pilsbryi

55311020[12]00010100100010121110022200002021102100102010020311010011110
311110001011321011101111221201100011012000004401100110?401000121112001
100120

Pacificor mulleri

5421102020002000010001012111002220000022110210011101102021101001011031
1110101011221011101111221201100011012000004401100110?40100012111100110
0120

Glans trapezia

2021002030013010110010002121102220010101000010012001000011101201001040
1111413021000211101111221201100000200000001102100100030000000012000110
0011

Centrocardita aculeata
3221002030013010010010002111102220010101000010012001000011101211001040
1111413021000211101111221201100000200000001102100100031000000012000110
0011

Carditamera arata

2340002010013110110010002121102220000101110010112001000011101201001040
1011413021000211111111221201100011200000012202000120032000000011110110
0111

Carditamera floridana

2240002010013110110010002121102220000101110010112001000011101201001040
1011413021000211111111221201100011200000012202000120002000000011010110
0111

Carditamera affinis

2440002010013110110010000121102220000100110010112001000011100201001040
1011413021002211111111221201000011102100002201000120031000000012010110
0011

Carditamera gracilis

2240002010013110110010000121102220000100110010112001000001100201001040
1011413021002211111111221201000011101100002200000120031000000012010110
0011

Paraglans calcitrapoides

3021102030012000010000020011102220010022000010002001100011101201011020
1110311121110011101111221201111112200000011102100120010000001011101110
0101

Kolmeris tehuelchana

1112111000000000010000021001102220101002110110002021100231100001011010
1110301021302011101111221201200011000001003003100110?40100000011201110
0110
 Appendix 3B - Matrix data in TNT format.

nstates cont ;
xread
'Carditidae'144 77

&[continuous]
Crassatella_ponderosa 0 62.57
Spissatella_trailli 0 38.86
Astarte_sulcata 0 17.87
Laevastarte_basteroti 0 33.17
Palaeocardita_stoecklini 22-23 21.88
Palaeocardita_crenata 22-24 23
Schizocardita_cristata 35 36.71
Birkelundita_turoniense 34 8.7
Thecalia_concamerata 24-25 12.58
Powellina_brookesi 21 13.32
Beguina_semiorbiculata 46 66.75
Cardita_variegata 25 25
Cardita_crassa 18-21 24.01
Claibornicardia_paleopatagonica 30-33 37.15
Claibornicardia_alticostata 30-33 44.94
Claibornicardia_acuticostata 30-33 33.13
Rotundicardia_mariobrosorum 25-27 28.97
Rotundicardia_rotunda 25-26 24.92
Rotundicardia_diversidentata 25-26 19.4
Kalelia_burmeisteri 25 43.52
Kalelia_pectuncularis 26-28 78.89
Kalelia_multicostata 28-30 50.49
Glyptoactis_hadra 19-20 33.6
Baluchicardia_beaumonti 20-22 26.5
Venericardia_imbricata 28-30 31.01
Arcturellina_asperula 28-30 13.6
Fasciculicardia_subintermedia 28-32 68.21
Fasciculicardia_acanthodes 24-26 51.58
Fasciculicardia_sp 25 34.6
Purpurocardia_purpurata 22-26 27.17
Purpurocardia_elegantoides 22-23 14.64
Cardites_antiquatus 21-22 26.8
Cardites_partschii 22-23 18.23
Cardites_feruglioi 28-29 41.85
Cardiocardita_ajar 23-26 20
Coripia_unidentata 25-26 5.26
Miodomeris_cossmanni 18-20 3.04
Scalaricardita_scalaris 23-25 12
Vimentum_dilectum 27-29 8
Pleuromeris_tridentata 17-18 6.15
Pleuromeris_decemcostata 11-12 5.84
Pteromeris_perplana 10-11 3.5
Pleuromeris_fueguina 18-21 15.34
Pleuromeris_sulcolunularis 18-20 16.6
Cyclocardia_borealis 20-23 40
Darwinicardia_patagonica 23-26 30.33

file:///D|/DAMIAN/Papers%20mios/tjsp_a_1532463_sm2944.txt[15/01/2019 12:19:59]
 Darwinicardia_angusticostata 22-23 19.72
Venericor_planicosta 31-34 69.64
Bathycardita_raouli 14-16 34
Megacardita_jouanetti 19-21 53.28
Neovenericor_paranensis 23-25 73.6
Neovenericor_ponderosa 25-28 73.97
Venericor_aposmithii 31-33 93
Venericor_bashiplata 28-30 63
Leuroactis_pilsbryi 30-33 93
Pacificor_mulleri 30-34 63
Cyclocardia_awamoensis 20-24 17.95
Cyclocardia_ventricosa 20-22 15.28
Pleuromeris_zelandica 16-19 8
Glans_trapezia 18-21 10.48
Centrocardita_aculeata 20-22 24.6
Carditamera_arata 20 41.16
Carditamera_floridana 17-19 26.97
Carditamera_affinis 17-18 62.27
Carditamera_gracilis 17-19 25.24
Paraglans_calcitrapoides 23-24 7.76
Scalaricardita_camaronesia 23-25 10.36
Cyclocardia_cannada 23-27 15.03
Neovenericor_austroplata 21-22 99.31
Neovenericor_carrerensis 24-26 87.15
Scalaricardita_laciarina 25-27 14.42
Kolmeris_tehuelchana 13-15 12.57
Cyclocardia_mesembria 21-22 12.2
Cyclocardia_dalek 21-24 14.74
Cyclocardia_compressa 15-17 14.33
Cyclocardia_nortensis 21-22 28.66
Purpurocardia_leonensis 24-26 29.05

&[numeric]
Crassatella_ponderosa
411010300030100100011220001111101000321102010324001001530002000120500100403020402010000000000022
324333423000235522220?725232232121020001000132
Spissatella_trailli
410020300031100100011120001111101000321102010324001012530002000120500100403010402010000000000022
3243334230002355222200725232232121020000022242
Astarte_sulcata
221010000030000101011210001100000100121101110324011002431000010100300100403021400111100000100122
3243334230002355232200725232232011021011000112
Laevastarte_basteroti
221010000030000101011210001101000100121101110324011002431000010100300100403021400111100000100122
3243334230002355232200725232232011021011000112
Palaeocardita_stoecklini
410020300131001100011120211100000000121101100324011002531002000120500110403021402111000110111201
1000222000000100020100705232232021020001100111
Palaeocardita_crenata
410020300131001100011120211100000000121101100324011002531002000120500110403021402111000110111201
1000222000000110020100705232232021020001100111
Schizocardita_cristata
410020300131211100011120211100000000221101100324011002531002000120500110403021402111000110101201
1000222000000010020110705232232021020001100111

file:///D|/DAMIAN/Papers%20mios/tjsp_a_1532463_sm2944.txt[15/01/2019 12:19:59]
 Birkelundita_turoniense 2?100001101000010000000011102221????????
101000001101000011000100000011101001010000111011112212001000001000000110000001000000000000100001
100010
Thecalia_concamerata
000000001001000010000000111022200000000000100020110100001000010000001111400011000001101111221200
0000001000000110000001000000000000100001100000
Powellina_brookesi
000000001001000010000000111022201000000000100020110100001000010000001110400011000001101111221200
0000001000000010000001000000000000100001100001
Beguina_semiorbiculata 00000000100100000000000001102221????????
001000000101000010010110000011100000010000011011112212100000000000000000031001401020000000100001
100010
Cardita_variegata
000000001001000000000000111022200000000000100000110100001000010000001110100001000001101111221200
0000001000100000000001000000000000100001100000
Cardita_crassa
000000001001001000000000111022200000000000100000110100101000010000001110100001000001101111221200
0000001000000000000001000000000000100001100000
Claibornicardia_paleopatagonica
410020100020100101000101111022200111201111100010011000011012010100301110111021210011101111221201
1112222010000044010001401120000010111011110100
Claibornicardia_alticostata
410020200020100101000101111022200111201111100010011000011012010110301110021021210011101111221201
1112222010000044010001401120000010111011100100
Claibornicardia_acuticostata
211020300020100100000201111022200101201101100010011000011012010110301110021021210011101111221201
1112222010000044010001401120000010111011100100
Rotundicardia_mariobrosorum [2 5]1101000002010010000010011102221????????
111000000110000110020101001011100210212100111011112212011000122000000101020001501120000010111011
110101
Rotundicardia_rotunda
511010000020100100000100111022200111201111100000011000011012010100101110021021212011101111221201
1000122000000211020001501120000010111011110101
Rotundicardia_diversidentata
511010000020000101000101111022200111201111100010011000011012010100101110021021210011101111221201
1000122000000111020001501120000010111011110101
Kalelia_burmeisteri
521020300110000100010201111022201111210011100010111000111012010110201110001021110011101111221200
2131121020000011010001400120000010111011100100
Kalelia_pectuncularis
521020300110000100010200111022201111010011100110111000211012010110211110001021110011101111221200
2131121020000011010001200120000010111011100100
Kalelia_multicostata
521020300110000100010200111022201111010011100110111000211012010110211110001021210011101111221200
2131121020000011010001400120000010111011100100
Glyptoactis_hadra
410020200020100100000122211022201111201111100020111000011012010100401110011121111001101111221201
1111122010000011010001201120000010111011100100
Baluchicardia_beaumonti
211010200020000100010122111022200101201101100010111000011012010110001110021021210011101111221201
1112222000000000010001600100000010111011100100
Venericardia_imbricata
211010300120000100000200111022200111200001100010011010111012010110201110001021112011101111221201
1111122000000111021001200100000010111011111101

file:///D|/DAMIAN/Papers%20mios/tjsp_a_1532463_sm2944.txt[15/01/2019 12:19:59]
 Arcturellina_asperula
211010300120000100000200111022200100220000100020011000111012010110201110311121112011101111221201
1111122000000111021001200100000010111011111101
Fasciculicardia_subintermedia
411020200020000100000100211022200101200001100020111000011012010100101110211121111011101111221201
1111222020000011010001401120001010111011100110
Fasciculicardia_acanthodes
411020200020000100000100211022200101200001100020111000011012010100111110211121111011101111221201
1111222020000011010001401120001010111011100110
Fasciculicardia_sp 411020200020000100000100211022200101200011100????????????0???????0??
1110113121110011101111221201100?222?2000001101000160112000101011???1100?1?
Purpurocardia_purpurata
410010000020000100000100211022201100220001100101010000311011010110301110301021210211101111221200
1111121020000011010001201100000000112111100110
Purpurocardia_elegantoides
410010000020000100000100111022201100210001100101010000311011010110301110301021210211101111221200
1111121020000011011001201100000000111111100100
Cardites_antiquatus
420020200020100100000122111022200101200011100233311000111012010110311110113021200011101111221201
2000112020000000010001200120000010111011100110
Cardites_partschii
420020200020100100000122111022200101200011100233311000111012010110301110113021200011101111221201
2000112020000000010001200120000010111011100100
Cardites_feruglioi
420020200020100100000122111022200101200011100233311000111012010110301110112021200011101111221201
2000112020000000010001200120000010111011100120
Cardiocardita_ajar
520010100030100100102200011022200101200000100120011002311012010110201110303021210011101111221201
1111322000000111021001200100000000110011100101
Coripia_unidentata [1
3]3111020001000010000020001102220011002110110002001100231100001011010111030302122201110111122120
10000010000000000011001211220000000112011110110
Miodomeris_cossmanni
121210000000000100000210011022201010021101100020011000311000010110101110301021300011100100101201
100001000000001003100110?400000001111011100110
Scalaricardita_scalaris
331010200210000100000200011022200110021101100020011002311000010110101110303021222011101111221201
0000010000000000031001211220000000112011110110
Vimentum_dilectum
431010200220000100000200111022200110021101100020011002311000010110101110303021221011101111221201
0000010000000000011001211220000000111011110110
Pleuromeris_tridentata
321010100010000100000220011022201010021101100020011002311000010110101110401011202011101111221201
1000222000000000031001200100000000111011110110
Pleuromeris_decemcostata
321010100010000100000220011022201010021101100020011002311000010110101110401021202011101111221201
1000222000000000031001200100000000111011100110
Pteromeris_perplana
121210000000000100000210011022201010021101100000011012311000010110101110301021300011101111221201
100011000000003403100110?400000000111011100110
Pleuromeris_fueguina
321010000010000100000221011022201010021101100000011002311010010110101110301021200011101111221201
1000222000000100031001201100000000111011100110
Pleuromeris_sulcolunularis

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 321010000010000100000221011022201010121101100000011002311010010110101110301021200011101111221201
1000222000000100031001201100000000111011100110
Cyclocardia_borealis 531010000220000100000100111022201010020001100010111002311001010110[1
3]01110103021221011101111221200100002012000000001100110?020000100111011110110
Darwinicardia_patagonica [3 5]310[1 2]02000200001010001[0
2]0111022200001210011100000211000211012010110401110101011220011101111221201112122200000010001100
1401120000000111011100101
Darwinicardia_angusticostata
531010200020000100000101111022200101200011100000211000211012010110401110111021220011101111221201
1121222000000100011001401120000000111011100111
Venericor_planicosta 311020[0
2]0001000010001012221002220000022110210011101102021101001011032111010101122101110111122120110001
1012000004201100110?401000121111001100120
Bathycardita_raouli
610020100230100100010000111022201000211111100100011002311011010110301110200021100011101111221201
200002012000012201100110?400010121101001100120
Megacardita_jouanetti
610020100230100100010021211022200000211111100010011000111011010110311110100011100011101111221201
200012012000004201100110?400000021101001100120
Neovenericor_paranensis
410020100210100100010022111022200000201111100010011000111011010110311110110001121011101111221201
200011102000013201100110?400100021111001100120
Neovenericor_ponderosa
410020100210100100010022111022200000211111100010011000111011010110311110110001121011101111221201
200011102000013201100110?400100021111001100120
Venericor_aposmithii 311020[1
2]0001000010001012221002220000202110210011101102021101001011031111010101132101110111122120110001
1012000004201100110?401000121111001100120
Venericor_bashiplata 311020[0
2]0001000010001012221002220000202110210011101102031101001011032111010101132101110111122120110001
1012000003201100110?401000121111001100120
Leuroactis_pilsbryi 311020[1
2]0001010010001012111002220000202110210010201002031101001111031111000101132101110111122120110001
1012000004401100110?401000121112001100120
Pacificor_mulleri
211020200020000100010121110022200000221102100111011020211010010110311110101011221011101111221201
100011012000004401100110?401000121111001100120
Cyclocardia_awamoensis
531010000220000100000100111022201011000001100010011012211001010110[1
3]011101030112200111011112212011000000120000000011001101020000100111011110110
Cyclocardia_ventricosa
531010000220000100000100111022201011020001100010111002311001010110301110103021222011101111221200
1000020120000000011001201020000100111011110110
Pleuromeris_zelandica
321010100010000100000220011022201010021101100020011002311000010110101110401021202011101111221201
1000222000000100031001201100000000111011100110
Glans_trapezia
210020300130101100100021211022200101010000100120010000111012010010401111413021000211101111221201
1000002000000011021001000300000000120001100011
Centrocardita_aculeata
210020300130100100100021111022200101010000100120010000111012110010401111413021000211101111221201
1000002000000011021001000310000000120001100011
Carditamera_arata
400020100131101100100021211022200001011100101120010000111012010010401011413021000211111111221201

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 1000112000000122020001200320000000111101100111
Carditamera_floridana
400020100131101100100021211022200001011100101120010000111012010010401011413021000211111111221201
1000112000000122020001200020000000110101100111
Carditamera_affinis
400020100131101100100001211022200001001100101120010000111002010010401011413021002211111111221201
0000111021000022010001200310000000120101100011
Carditamera_gracilis
400020100131101100100001211022200001001100101120010000011002010010401011413021002211111111221201
0000111011000022000001200310000000120101100011
Paraglans_calcitrapoides
211020300120000100000200111022200100220000100020011000111012010110201110311121110011101111221201
1111122000000111021001200100000010111011100101
Scalaricardita_camaronesia
331010200210000100000100011022200110021101100020011002311000010110101110403021222011101111221201
0000010000000000031001211220000000112011110110
Cyclocardia_cannada
531010000220000100000100111022200010020001100020111002311001010110101110403021220011101111221200
1000020120000000011001[1 4]01020000100111011100111
Neovenericor_austroplata
310020100210100100010022111022200000201111100010011000111011010110311110101001121011101111221201
200011102000013201100110?401000021111001100120
Neovenericor_carrerensis
410020100210100100010022211022200000201101100010011000111011010110311110101001121011101111221201
200010102000013201100110?401000021111001100120
Scalaricardita_laciarina
331011200210000100000200011022200110011101100020011002311000010110101110303021222011101111221201
0000010000000000031001211220000000112011110110
Kolmeris_tehuelchana
121110000000000100000210011022201010021101100020211002311000010110101110301021302011101111221201
200011000001003003100110?401000000112011100110
Cyclocardia_mesembria
531010200220000100000100111022200010000001100010111002311001010110101110?????
1220011101111221200100002012000000001100110?020000100111011110110
Cyclocardia_dalek
531010000220000100000100111022200010020001100020111002311001010110101110403021220011101111221200
1000020120000020011001[2 4]01020000100111011100111
Cyclocardia_compressa
531010000220000100000100111022201010020001100010111002311001010110101110103021221011101111221200
100002012000000001100110?020000100111011110110
Cyclocardia_nortensis
531010000220000100000100111022201011000001100010011012211001010110301110103011220011101111221201
1000000120000020011001[1 2]01020000100111011100110
Purpurocardia_leonensis
410010000020000100000100111022201100220001100111010000311011010110301100101021210211101111221200
1111122020000011020001201100000000111111100110
;

taxname=
ccode + 1 4 5 22 23 28 56 61 69 74 104 134 *;
hold1000;

proc /;
comments 0;

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Appendix 4

Correspondence of number of radial ribs with specific identity in Carditidae

As discussed in Matherials and methods section and Appendix 2, external sculpture is

an important feature in carditid systematics for many authors (e.g. Harris, 1919;

Verastegui, 1953; Heaslip, 1968; Maxwell, 1969), and radial sculpture is the principal

type of sculpture in the family. The number of radial ribs remains virtually fixed

throughout lifespan of an individual and this number may shows little intraspecific

variation according my observations. For test the correspondence between this number

and the species identity of an individual, a Generalized Lineal Model (GLM) (Nelder &

Wedderburn, 1972) analysis was performed using R (R Development Core Team, 2013)

(function: glm).

The question of the analysis is if number of radial ribs is related to species

identity or may be influenced by other variables in carditid bivalves. Radial ribs number

of 304 adult specimens were counted and linear measures of length and height (both in

mm), according to Heaslip (1968) standard measures, were included (Table S1). Three

predictor variables were considered: species identity (categorical), length (continuous),

and shape (considered as height/large) (continuous). Response variable was number of

ribs (discrete) (normal distribution, mean=24.513, median=25) and was modelled with

normal error structure and identity link function. Two initial models were considered,

due to non-independence between length and height. The first one (GLM1) includes

species identity and large as random effects and the second one (GLM2) includes

species identity and shape as random effects. Length is not completely independent

from species identity, so that, a third model (GLM3) including a nesting approach

between these two variables was considered. All models include number of ribs as fixed
effect. A stepwise backwards-elimination procedure was used to obtain the minimal

adequate model. Percentage of variance explained was calculated for each model,

estimated from explained deviance and total deviance of null model.

First model, GLM1, showed a reduced model including species identity and size,

and they explained 65.27% of total variance. Discarding length, species identity

explained 63.52% of total variance (only 1.8% less of anterior number). Second model

(GLM2) concluded in a reduced model that includes only species identity. This model

explained 63.52% of total variance. Third model (GLM3) resulted in a reduced model

including length, species identity and its interaction, and explained 68.08% of total

variance. Results are resumed in Table S2.

According to these results, number of ribs is non-related to size, but it is related

to length and species identity. Shape may be influenced by other variables (e.g. sex,

growth or environment). In contrast, length is not completely independent from species

identity, and the nested model showed the greater value of variance explained. Despite

this, species identity constitutes the variable that most percentage of variance explains.

In conclusion, correspondence of number of radial ribs with species identity is true in

Carditidae.

Table S1

Species identity Length Height Shape Radial Ribs

Arcturellina asperula 14 12,7 1,102362205 26
Arcturellina asperula 13,2 12,5 1,056 28
Cardites antiquatus 28,6 25,2 1,134920635 21
Cardites antiquatus 30,9 30,5 1,013114754 22
Cardites antiquatus 23,1 22,9 1,008733624 21
Cardites antiquatus 23,4 22,6 1,03539823 20
Cardites feruglioi 38,4 37 1,037837838 29
Cardites feruglioi 40,5 38,4 1,0546875 29
Cardites feruglioi 42,5 43,4 0,979262673 28
Cardites feruglioi 41,8 39,5 1,058227848 29
Cardites feruglioi 44,9 42,9 1,046620047 31
Cardites feruglioi 43,1 41,8 1,031100478 29
Cardites feruglioi 41 39,1 1,04859335 29
Cardites feruglioi 40,4 40 1,01 29
Cardites feruglioi 49 46,8 1,047008547 30
Cardites feruglioi 41,9 37,3 1,123324397 28
Cardites feruglioi 40 38,7 1,033591731 29
Cardites feruglioi 38,4 35,9 1,069637883 28
Cardites feruglioi 29,1 28,4 1,024647887 28
Cardites feruglioi 41,9 39,6 1,058080808 28
Cardites feruglioi 52 49,4 1,052631579 28
Cardites feruglioi 44 41,5 1,060240964 28
Cardites feruglioi 33,1 32,2 1,027950311 27
Cardites feruglioi 38,8 35 1,108571429 29
Cardites feruglioi 40,7 37,7 1,079575597 28
Cardites feruglioi 36,5 34,9 1,045845272 25
Cardites feruglioi 41,2 36,2 1,138121547 31
Cardites laevicostata 45,1 35,5 1,270422535 21
Cardites laevicostata 42,1 35,4 1,189265537 22
Cardites laevicostata 37,9 29,8 1,271812081 21
Cardites laticostatus 34,4 27,7 1,241877256 17
Cardites laticostatus 26,1 18,9 1,380952381 18
Cardites laticostatus 26,9 19,7 1,365482234 20
Cardites laticostatus 26,4 19,9 1,326633166 20
Cardites laticostatus 27,7 21,8 1,270642202 20
Cardites laticostatus 35,2 26,1 1,348659004 20
Cardites monilifera 17,7 14,7 1,204081633 20
Cardites monilifera 16,2 12,9 1,255813953 19
Cardites monilifera 13,9 11,6 1,198275862 21
Cardites monilifera 9,7 8,1 1,197530864 17
Cardites partschii 20,5 20,7 0,990338164 22
Cardites partschii 18,1 19 0,952631579 22
Cardites partschii 17,5 16,6 1,054216867 23
Cardites partschii 17,9 18,1 0,988950276 23
Cardites partschii 18,8 18,1 1,038674033 22
Cardites partschii 16,6 17,1 0,970760234 23
Claibornicardia paleopatagonica 42,7 31,8 1,342767296 28
Claibornicardia paleopatagonica 52,7 43,8 1,203196347 25
Claibornicardia paleopatagonica 18,7 17,4 1,074712644 25
Claibornicardia paleopatagonica 42,2 36,1 1,168975069 33
Claibornicardia paleopatagonica 38,4 32,9 1,167173252 33
Claibornicardia paleopatagonica 38,9 33,7 1,154302671 32
Claibornicardia paleopatagonica 37,8 33,3 1,135135135 33
Claibornicardia paleopatagonica 35,9 31,9 1,12539185 30
Claibornicardia paleopatagonica 36,4 31,3 1,162939297 32
Claibornicardia paleopatagonica 31,9 28,7 1,111498258 31
Claibornicardia paleopatagonica 27,1 24,2 1,119834711 29
Claibornicardia paleopatagonica 35,3 30,9 1,142394822 31
Claibornicardia paleopatagonica 32,1 28,7 1,118466899 28
Claibornicardia paleopatagonica 40,1 34 1,179411765 31
Claibornicardia paleopatagonica 38,4 31,4 1,222929936 31
Claibornicardia paleopatagonica 26,8 25,2 1,063492063 25
Claibornicardia paleopatagonica 22,4 21,2 1,056603774 29
Claibornicardia paleopatagonica 13,8 12 1,15 23
Claibornicardia paleopatagonica 17,8 15,7 1,133757962 26
Claibornicardia paleopatagonica 18,5 17,2 1,075581395 22
Cyclocardia cannada 17,9 18,6 0,962365591 24
Cyclocardia cannada 18 19,3 0,932642487 24
Cyclocardia cannada 16,8 16,5 1,018181818 25
Cyclocardia cannada 13,5 12,7 1,062992126 27
Cyclocardia cannada 9,2 9,2 1 23
Cyclocardia mariobrosorum 19,5 19,4 1,005154639 26
Cyclocardia mariobrosorum 19 18,6 1,021505376 26
Cyclocardia mariobrosorum 14,9 15,3 0,973856209 25
Cyclocardia mariobrosorum 12,9 12,2 1,057377049 25
Cyclocardia mariobrosorum 17,2 16,8 1,023809524 27
Cyclocardia mariobrosorum 12,3 11,4 1,078947368 26
Cyclocardia mariobrosorum 18,6 18,9 0,984126984 26
Cyclocardia mariobrosorum 17,7 18,7 0,946524064 26
Cyclocardia mariobrosorum 11,8 11,9 0,991596639 23
Cyclocardia mariobrosorum 15,8 15,9 0,993710692 27
Cyclocardia mariobrosorum 16,6 16,8 0,988095238 27
Cyclocardia mariobrosorum 14,2 13 1,092307692 27
Cyclocardia mariobrosorum 14,4 13,5 1,066666667 28
Cyclocardia mariobrosorum 11 11,2 0,982142857 24
Cyclocardia mariobrosorum 16,2 15,2 1,065789474 27
Cyclocardia mariobrosorum 15,7 14,6 1,075342466 28
Cyclocardia mariobrosorum 12,3 12,2 1,008196721 25
Cyclocardia mariobrosorum 10,3 10 1,03 26
Darwinicardia patagonica 31,5 31,3 1,006389776 25
Darwinicardia patagonica 36,2 34,2 1,058479532 24
Darwinicardia patagonica 47,8 49,8 0,959839357 23
Darwinicardia patagonica 40,5 39,7 1,020151134 24
Darwinicardia patagonica 39,4 36,5 1,079452055 23
Darwinicardia patagonica 30,3 29,7 1,02020202 23
Darwinicardia patagonica 33 31,1 1,061093248 23
Darwinicardia patagonica 30,5 30,8 0,99025974 23
Darwinicardia patagonica 17,2 17,4 0,988505747 20
Darwinicardia patagonica 17,9 16,7 1,071856287 19
Darwinicardia patagonica 13,6 13,3 1,022556391 17
Darwinicardia patagonica 15,7 15,2 1,032894737 19
Darwinicardia patagonica 17,1 16,8 1,017857143 20
Darwinicardia patagonica 41,2 36,7 1,122615804 26
Darwinicardia patagonica 30,8 30,5 1,009836066 22
Darwinicardia patagonica 17,3 16,4 1,054878049 25
Darwinicardia patagonica 34,4 33,7 1,020771513 24
Darwinicardia patagonica 24,5 23,7 1,033755274 25
Darwinicardia patagonica 35,3 36,1 0,977839335 26
Darwinicardia patagonica 34,5 33,8 1,020710059 26
Darwinicardia patagonica 25,8 25 1,032 25
Darwinicardia patagonica 28,9 28,3 1,021201413 25
Darwinicardia patagonica 17,9 17,1 1,046783626 16
Darwinicardia patagonica 37,9 40 0,9475 25
Darwinicardia patagonica 34,9 35,7 0,977591036 25
Darwinicardia patagonica 33,4 34,6 0,965317919 26
Darwinicardia patagonica 27,7 28,6 0,968531469 25
Darwinicardia patagonica 17,8 17,2 1,034883721 16
Darwinicardia patagonica 31,9 31,2 1,022435897 25
Darwinicardia patagonica 38,8 37,7 1,029177719 25
Darwinicardia patagonica 32,9 32,5 1,012307692 25
Darwinicardia patagonica 37,5 36,9 1,016260163 26
Darwinicardia patagonica 35,2 32,3 1,089783282 25
Darwinicardia patagonica 30,8 32,4 0,950617284 25
Darwinicardia patagonica 29,3 30,9 0,948220065 26
Darwinicardia patagonica 15,9 16,2 0,981481481 25
Darwinicardia patagonica 41,3 42,8 0,964953271 24
Darwinicardia patagonica 23,1 23,7 0,974683544 25
Darwinicardia patagonica 24,6 23,7 1,037974684 25
Darwinicardia patagonica 38,7 41,3 0,937046005 27
Darwinicardia patagonica 38,9 39,8 0,977386935 26
Darwinicardia patagonica 39,6 39,1 1,012787724 26
Darwinicardia patagonica 36,2 36,4 0,994505495 25
Darwinicardia patagonica 41,8 39,9 1,047619048 25
Darwinicardia patagonica 20,2 19,3 1,046632124 26
Darwinicardia patagonica 40,3 41,8 0,964114833 25
Darwinicardia patagonica 36,9 39,4 0,936548223 27
Darwinicardia patagonica 35,9 36,8 0,975543478 26
Darwinicardia patagonica 39,4 37,7 1,045092838 25
Darwinicardia patagonica 38,8 38,2 1,015706806 25
Darwinicardia patagonica 28,1 31,1 0,903536977 25
Darwinicardia patagonica 27,2 28,2 0,964539007 25
Darwinicardia patagonica 32,9 34 0,967647059 25
Darwinicardia patagonica 24,5 23,6 1,038135593 24
Darwinicardia patagonica 27,3 27,4 0,996350365 25
Darwinicardia patagonica 29,4 27,4 1,072992701 23
Darwinicardia patagonica 27,8 27,5 1,010909091 25
Darwinicardia patagonica 33,8 32,4 1,043209877 24
Darwinicardia patagonica 38,2 39,5 0,967088608 24
Darwinicardia patagonica 40,4 41,4 0,975845411 23
Darwinicardia patagonica 37 38,7 0,956072351 27
Darwinicardia patagonica 29,1 29,3 0,993174061 25
Darwinicardia patagonica 35,5 34,9 1,017191977 24
Darwinicardia patagonica 33,5 33,4 1,002994012 24
Darwinicardia patagonica 34,4 34,6 0,994219653 24
Darwinicardia patagonica 30,2 29,1 1,037800687 24
Darwinicardia patagonica 26,1 26,2 0,996183206 23
Darwinicardia patagonica 24,4 23,9 1,020920502 25
Darwinicardia patagonica 32,4 33,6 0,964285714 25
Darwinicardia patagonica 33,6 32,8 1,024390244 25
Darwinicardia patagonica 33,1 32,5 1,018461538 24
Darwinicardia patagonica 14,7 13,8 1,065217391 18
Darwinicardia patagonica 46,9 47,8 0,981171548 26
Darwinicardia patagonica 30,3 30,5 0,993442623 24
Darwinicardia patagonica 39,9 39,5 1,010126582 25
Darwinicardia patagonica 21,1 20,9 1,009569378 26
Darwinicardia patagonica 14,9 14,6 1,020547945 25
Darwinicardia patagonica 15,2 15,2 1 25
Darwinicardia patagonica 15,9 15,3 1,039215686 26
Darwinicardia patagonica 12,7 11,7 1,085470085 24
Darwinicardia patagonica 15,2 14,5 1,048275862 26
Darwinicardia patagonica 18,7 17,5 1,068571429 28
Darwinicardia patagonica 12 11,7 1,025641026 28
Darwinicardia patagonica 16,6 15,5 1,070967742 26
Darwinicardia patagonica 45,4 46,9 0,968017058 22
Darwinicardia patagonica 46,1 44 1,047727273 24
Darwinicardia patagonica 48,6 47,9 1,014613779 24
Darwinicardia patagonica 20,4 20,3 1,004926108 23
Darwinicardia patagonica 11,9 12 0,991666667 26
Darwinicardia patagonica 45,4 43,9 1,034168565 24
Darwinicardia patagonica 14,2 14 1,014285714 22
Darwinicardia patagonica 29,2 29 1,006896552 24
Darwinicardia patagonica 33,5 32,8 1,021341463 22
Darwinicardia patagonica 34,7 33,4 1,038922156 23
Darwinicardia patagonica 33,8 34,1 0,991202346 23
Darwinicardia patagonica 21,7 20 1,085 24
Kalelia burmeisteri 48,3 47,4 1,018987342 25
Kalelia burmeisteri 12,8 10,2 1,254901961 20
Kalelia burmeisteri 11,7 10,7 1,093457944 20
Kalelia burmeisteri 13 12 1,083333333 22
Kalelia burmeisteri 10,2 8,7 1,172413793 21
Kalelia burmeisteri 11,6 9,9 1,171717172 20
Kalelia burmeisteri 12,5 11,4 1,096491228 23
Kalelia burmeisteri 7,7 6,6 1,166666667 23
Kalelia burmeisteri 25,4 23,6 1,076271186 22
Kalelia burmeisteri 30,3 31,4 0,964968153 21
Kalelia burmeisteri 41,2 39,1 1,05370844 22
Kalelia burmeisteri 36,5 36,4 1,002747253 22
Kalelia burmeisteri 34,1 33,6 1,014880952 21
Kalelia burmeisteri 27,9 28,1 0,992882562 21
Kalelia burmeisteri 53,8 52,8 1,018939394 24
Kalelia burmeisteri 50 51,4 0,972762646 20
Neovenericor austroplata 118,9 112,7 1,05501331 19
Neovenericor austroplata 82,3 75,1 1,09587217 20
Neovenericor paranensis 87,9 85,3 1,030480657 24
Neovenericor paranensis 46,8 44,3 1,056433409 23
Pleuromeris fueguina 15,2 15,5 0,980645161 21
Pleuromeris fueguina 16 15,6 1,025641026 19
Pleuromeris fueguina 16,6 16,1 1,031055901 18
Pleuromeris fueguina 14,5 15 0,966666667 21
Pleuromeris sulcolunularis 13,4 13,3 1,007518797 17
Pleuromeris sulcolunularis 14,6 14 1,042857143 20
Pleuromeris sulcolunularis 11,5 11,5 1 18
Pleuromeris sulcolunularis 14,2 14,2 1 18
Pleuromeris sulcolunularis 15,9 14,9 1,067114094 20
Pleuromeris sulcolunularis 12,1 11,4 1,061403509 19
Pleuromeris sulcolunularis 17,8 16,8 1,05952381 20
Pleuromeris sulcolunularis 16,7 17,1 0,976608187 21
Pleuromeris sulcolunularis 16,7 16 1,04375 21
Pleuromeris sulcolunularis 15,4 13,8 1,115942029 21
Pleuromeris sulcolunularis 18,1 18,2 0,994505495 18
Pleuromeris sulcolunularis 21,2 20,6 1,029126214 21
Pleuromeris sulcolunularis 13,9 13,1 1,061068702 18
Purpurocardia elegantoides 17,8 15,8 1,126582278 22
Purpurocardia elegantoides 18,2 16,3 1,116564417 23
Purpurocardia elegantoides 11 10,5 1,047619048 22
Rotundicardia mariobrosorum 30,7 28,2 1,088652482 25
Rotundicardia mariobrosorum 27,8 26,4 1,053030303 23
Rotundicardia mariobrosorum 27,8 25,2 1,103174603 23
Rotundicardia mariobrosorum 26,9 27,2 0,988970588 24
Rotundicardia mariobrosorum 30,2 26,9 1,12267658 30
Rotundicardia mariobrosorum 43,1 42,2 1,021327014 29
Rotundicardia mariobrosorum 27,7 26,1 1,061302682 29
Rotundicardia mariobrosorum 24,3 23,7 1,025316456 26
Rotundicardia mariobrosorum 20,9 20,4 1,024509804 25
Rotundicardia mariobrosorum 20,3 19,1 1,062827225 26
Rotundicardia mariobrosorum 15,2 13,5 1,125925926 24
Rotundicardia mariobrosorum 23,9 22,3 1,071748879 28
Rotundicardia mariobrosorum 17,2 15,7 1,095541401 28
Rotundicardia mariobrosorum 35,3 32,6 1,082822086 26
Rotundicardia mariobrosorum 34,5 32,4 1,064814815 29
Rotundicardia mariobrosorum 32,7 29,3 1,116040956 27
Rotundicardia mariobrosorum 25 23,7 1,054852321 23
Rotundicardia mariobrosorum 26,4 26,2 1,007633588 24
Rotundicardia mariobrosorum 33,5 31,2 1,073717949 28
Rotundicardia mariobrosorum 29,5 27,3 1,080586081 26
Rotundicardia mariobrosorum 26,6 25 1,064 31
Rotundicardia mariobrosorum 42,6 39,3 1,083969466 31
Rotundicardia mariobrosorum 32,5 29,6 1,097972973 28
Scalaricardita camaronesia 13,9 14,1 0,985815603 14
Scalaricardita camaronesia 15,8 15,8 1 16
Scalaricardita camaronesia 13,8 13,4 1,029850746 15
Scalaricardita camaronesia 15 15,2 0,986842105 28
Scalaricardita camaronesia 10,5 10,6 0,990566038 16
Scalaricardita camaronesia 12,7 12,2 1,040983607 15
Scalaricardita camaronesia 15,4 14,5 1,062068966 26
Scalaricardita camaronesia 17,2 17,9 0,960893855 26
Scalaricardita camaronesia 12,2 11,7 1,042735043 25
Scalaricardita camaronesia 9,7 9,8 0,989795918 26
Scalaricardita camaronesia 16,9 17 0,994117647 25
Scalaricardita camaronesia 15,9 15,3 1,039215686 25
Scalaricardita camaronesia 14,8 14,7 1,006802721 25
Scalaricardita camaronesia 20,4 19,6 1,040816327 23
Scalaricardita camaronesia 13,1 13,6 0,963235294 22
Scalaricardita camaronesia 13,4 13,4 1 22
Scalaricardita camaronesia 10 10,3 0,970873786 22
Scalaricardita camaronesia 13,8 13,7 1,00729927 25
Scalaricardita camaronesia 10,5 10,1 1,03960396 21
Scalaricardita camaronesia 7,9 7,2 1,097222222 26
Scalaricardita camaronesia 11 11,2 0,982142857 27
Scalaricardita camaronesia 13,5 13,7 0,98540146 25
Scalaricardita camaronesia 12,5 12 1,041666667 24
Scalaricardita camaronesia 21,3 21,9 0,97260274 24
Scalaricardita camaronesia 9,7 9,2 1,054347826 24
Scalaricardita camaronesia 14,6 14,1 1,035460993 27
Scalaricardita camaronesia 8,2 8,3 0,987951807 23
Scalaricardita camaronesia 9,3 9,2 1,010869565 24
Scalaricardita camaronesia 9,4 9,6 0,979166667 22
Scalaricardita camaronesia 9,8 10,1 0,97029703 24
Scalaricardita camaronesia 16,9 17,3 0,976878613 23
Scalaricardita camaronesia 16,5 16,2 1,018518519 23
Scalaricardita camaronesia 12,9 13 0,992307692 24
Scalaricardita camaronesia 9,7 8,8 1,102272727 24
Scalaricardita camaronesia 11,2 11,4 0,98245614 24
Scalaricardita camaronesia 11,2 11,5 0,973913043 22
Scalaricardita camaronesia 10 10,2 0,980392157 25
Scalaricardita camaronesia 8,1 8,5 0,952941176 24
Scalaricardita camaronesia 11,1 11,2 0,991071429 22
Scalaricardita camaronesia 9,8 9,2 1,065217391 23
Scalaricardita camaronesia 8,8 8,6 1,023255814 25
Scalaricardita camaronesia 8,1 7,8 1,038461538 23
Scalaricardita camaronesia 8,3 7,8 1,064102564 25
Scalaricardita camaronesia 6,7 6,5 1,030769231 25
Scalaricardita camaronesia 5,7 5,1 1,117647059 23
Venericardia imbricata 31,2 33 0,945454545 29
Venericardia imbricata 24,2 23,7 1,021097046 29
Venericardia imbricata 22,1 22,3 0,99103139 30
Venericardia imbricata 38,6 36,6 1,054644809 28
Venericardia imbricata 38,3 38,3 1 31
Venericor planicosta 83,5 91,1 0,916575192 34
Venericor planicosta 79,9 87,3 0,915234822 33
Venericor planicosta 75,9 76,6 0,990861619 31
Venericor planicosta 52,8 49,5 1,066666667 34
Venericor planicosta 51,2 48,2 1,062240664 32
Venericor planicosta 37,2 34,6 1,075144509 37

Table S2

Initial Model Reduced Description % Explained

Model

GLM1 Radial_Ribs~Identity+Lenght+Identity:Length

GLM1a Radial_Ribs~Identity+Length* 65.27

GLM2 Radial_Ribs~Identity+Shape+Identity:Shape

GLM2a Radial_Ribs~Identity+Shape

GLM2b Radial_Ribs~Identity* 63.52

GLM3 Radial_Ribs~Identity/Length* 68.08

Appendix 5

Synapomorphies

The family Carditidae are defined by 25 synapomorphies, including the presence

of radial ribs, inner ventral margin crenulations that correspond to radial ribs and

absence of laminar lateral teeth. Clade 1 or Paleocarditinae presents four

synapomorphies as the presence of a very strong angle between posterior and dorsal

margins, enlarged dorsal margin, a notch in posterior margin and strongly rounded

outline of umbones in external view. Clade 2 shows 13 synapomorphies including

posterior protrusion (identation) of radial ribs and subrounded lunule, among others.

Clade 3 (Birkelundita, Beguina and Cardita species) presents three synapomorphies

(right anterior tooth vertical, left anterior tooth nearly horizontal and interspaces in adult

regions with V-section). Clade 4 is supported by 11 synapomorphies, with three

exclusive from all discrete approach and three from continuous and discrete approach,

referred to small size of shells, number of radial ribs, posterior margin slightly

truncated, subcentrally placed umbos, strongly impressed lunule, radial ribs wider

towards posterior side, small nodes and pallial line placed at quarter of total valve

height. Clade 5 also includes 11 synapomorphies (five exclusive of all discrete

approach) as obtuse-triangular outline, posterior margin slightly inclined, dorsal margin

curved and descendent, low convex umbonal area and very low radial ribs without

nodes. Clade 6, genera Pleuromeris, has only one synapomorphy on continuous and

discrete characters approach, radial ribs with subtriangular transverse section on adult

portion of shells. Cardites, or Clade 7, presents 15 synapomorphies (two of it exclusives

of all discrete approach and five of it of cont+disc approach), including subtriangular

outline of juvenile shells, very convex umbonal region, absent or very reduced right
anterior tooth and left anterior tooth as tall as wide. Clade 8 is supported by 19

synapomorphies as absence of marked angle between posterior and dorsal margins,

thick shells, strongly recurved umbones, presence of a nymph margin projection, left

anterior tooth taller towards venter, left posterior tooth very curved, radial ribs smooth

and little raised and large muscle scar strongly pronounced. Clade 9 shows seven

synapomorphies excluding Cardites (all discrete approach) or six including it (cont +

disc approach), as regular convexity of umbonal area, lunule slightly marked and

presence of paracostal ribs. Genus Cyclocardia (Clade 10) are supported by twelve

synapomorphies (one exclusive of cont + disc approach and four exclusive of all disc

approach) including shells with subrounded outline, rounded umbones in external view,

right anterior tooth pustular, wider radial ribs towards venter and obsolete to gerontic

portion, and presence of a flat zone between anterior margin and muscle scar.

Thecaliinae (Clade 11) has six synapomorphies as very low number of radial ribs (less

than 15), presence of an incubatory chamber in females, right anterior tooth inclined

posteriorly and left anterior tooth subrectangular. Clade 12 includes seven

synapomorphies exclusive from all disc approach, as size of shells less than 6 cm with

subtrigonal outlines and interspaces with V-shaped section. Clade 13 is present in both,

all disc and cont+disc approaches, with six synapomorphies in the first and eight in the

second, including convex lunule, hinge teeth with fine striations, right anterior tooth

inclined backwards, right middle tooth vertical, right posterior tooth curved, posterior

area defined by weaker radial ribs. Clade 14 is one of the most supported clades, with

23 synapomorphies (13 exclusive from all disc approach), as more than 29 radial ribs,

posterior margin slightly truncated, gentle marked angle between posterior and dorsal

margins, umbones placed at anterior third of valve length and in contact, lunule large,

hinge extended towards venter, right anterior tooth conspicuous, right middle tooth
scimitar-shaped, left anterior tooth subrectangular-shaped, radial ribs very low and

obsolete in gerontic part of valves and interspaces defined by a fine line. Clade 15 is

rescued only in cont+disc approach and has nine synapomorphies including elongate

outline of shells, angle between posterior and dorsal margins absent, umbones anteriorly

placed, lunule small, right middle tooth nearly vertically oriented, radial ribs wider

towards central part of valves and interspaces as wide as radial ribs. Clade 16 has six

synapomorphies (only one exclusive of all disc approach) as posterior margin truncated,

strongly marked angle between posterior and dorsal margins, subcentrally placed

umbones and right hinge with straight ventral edge. Finally, Clade 17 presents four

synapomorphies including lunule asymmetrical and projected as a sheet and left anterior

tooth wider than tall.

Synapomorphies listed below. They are obtained of both search approaches, all

discrete characters matrix (All Disc) and continuous + discrete characters matrix

(Cont+Disc). Synapomorphies not present in both approaches are indicated. Names of

clades are explained in the Results and Discussion sections. Node numbers followed by

a letter are exclusive from all discrete characters approach.

Spissatella trailli:

Char. 14: 0 --> 1

Char. 55: 0 --> 1

Char. 79: 2 --> 1

Char. 138: 1 --> 0

Char. 140: 0 --> 2

Char. 141: 0 --> 2

Char. 142: 1 --> 2

Astarte sulcata:
 Char. 2: 23 --> 1 (All Disc)

Char. 2: 16.609-22.395 --> 12.065 (Cont+Disc)

Laevastarte basteroti:

No autapomorphies

Palaeocardita stoecklini:

Char. 2: 2 --> 1 (All Disc)

Char. 2: 15.529 --> 14.773 (Cont+Disc)

Char. 113: 1 --> 0

Palaeocardita crenata:

No autapomorphies

Schizocardita cristata:

Char. 1: 23.000 --> 35.000

Char. 2: 16.609-22.395 --> 24.786 (Cont+Disc)

Char. 15: 0 --> 2

Char. 16: 0 --> 1

Char. 39: 1 --> 2

Char. 119: 0 --> 1

Birkelundita turoniense:

Char. 2: 16.609-16.879 --> 5.874 (Cont+Disc)

Char. 3: 0 --> 2 (All Disc)

Char. 5: 0 --> 1 (All Disc)

Char. 10: 0 --> 1

 Char. 13: 0 --> 1 (All Disc)

Char. 14: 1 --> 0 (All Disc)

Char. 18: 0 --> 1 (All Disc)

Char. 43: 0 --> 1

Char. 60: 0 --> 1

Char. 78: 0 --> 1

Char. 85: 0 --> 1 (All Disc)

Char. 99: 0 --> 1 (All Disc)

Char. 112: 0 --> 1

Char. 113: 0 --> 1 (All Disc)

Thecalia concamerata:

Char. 2: 8.993 --> 8.493 (Cont+Disc)

Char. 74: 0 --> 1

Char. 112: 0 --> 1

Powellina brookesi:

Char. 35: 0 --> 1

Char. 144: 0 --> 1

Beguina semiorbiculata:

Char. 1: 25.000-34.000 --> 46.000 (Cont+Disc)

Char. 1: 5 --> 6 (All Disc)

Char. 2: 16.609-16.879 --> 45.068 (Cont+Disc)

Char. 2: 3 --> 4 (All Disc)

Char. 27: 1 --> 0

Char. 51: 1 --> 0 (All Disc)

 Char. 62: 0 --> 1 (All Disc)

Char. 65: 0 --> 1

Char. 75: 1 --> 0

Char. 97: 0 --> 1

Char. 105: 1 --> 0

Char. 116: 0 --> 3

Char. 117: 0 --> 1

Char. 121: 0 --> 4

Char. 123: 0 --> 1

Char. 125: 0 --> 2

Cardita variegata:

Char. 109: 0 --> 1

Cardita crassa:

Char. 1: 3 --> 2 (All Disc)

Char. 17: 0 --> 1

Char. 57: 0 --> 1

Claibornicardia paleopatagonica:

Char. 9: 2 --> 1

Char. 67: 1 --> 0

Char. 75: 0 --> 1

Char. 76: 2 --> 1

Char. 140: 0 --> 1

Claibornicardia alticostata:
 Char. 2: 25.083 --> 30.342 (Cont+Disc)

Claibornicardia acuticostata:

Char. 9: 2 --> 3

Char. 24: 1 --> 2

Rotundicardia mariobrosorum:

Char. 34: 0 --> 1

Char. 61: 1 --> 0

Char. 113: 1 --> 0

Rotundicardia rotunda:

Char. 83: 0 --> 2

Char. 112: 1 --> 2

Rotundicardia diversidentata:

Char. 2: 16.825-19.560 --> 13.098 (Cont+Disc)

Char. 2: 2 --> 1 (All Disc)

Char. 20: 0 --> 1

Char. 26: 0 --> 1

Kalelia burmeisteri:

Char. 26: 0 --> 1

Kalelia pectuncularis:

Char. 2: 34.090 --> 53.265 (Cont+Disc)

Char. 2: 3 --> 4 (All Disc)

 Char. 121: 4 --> 2 (All Disc)

Kalelia multicostata:

Char. 81: 1 --> 2

Glyptoactis hadra:

Char. 1: 25.000-26.000 --> 19.000-20.000 (Cont+Disc)

Char. 1: 3 --> 2 (All Disc)

Char. 5: 1 --> 0

Char. 15: 0 --> 1 (All Disc)

Char. 25: 0 --> 2

Char. 26: 0 --> 2

Char. 35: 0 --> 1

Char. 69: 1 --> 4

Char. 85: 1 --> 0

Char. 121: 4 --> 2 (All Disc)

Baluchicardia beaumonti:

Char. 1: 25.000-30.000 --> 20.000-22.000 (Cont+Disc)

Char. 2: 20.937-22.368 --> 17.892 (Cont+Disc)

Char. 7: 2 --> 1 (Cont+Disc)

Char. 22: 0 --> 1

Char. 25: 0 --> 2

Char. 26: 1 --> 2 (Cont+Disc)

Char. 51: 0 --> 1

Char. 69: 3 --> 0

Char. 107: 1 --> 0 (Cont+Disc)

 Char. 113: 4 --> 0 (Cont+Disc)

Char. 114: 4 --> 0 (Cont+Disc)

Char. 121: 4 --> 6

Char. 123: 1 --> 0

Char. 125: 2 --> 0

Venericardia imbricata:

Char. 1: 25.000-26.000 --> 28.000-30.000 (Cont+Disc)

Char. 1: 3 --> 4 (All Disc)

Char. 55: 0 --> 1

Char. 83: 0 --> 2

Char. 140: 0 --> 1

Char. 141: 0 --> 1

Arcturellina asperula:

Char. 1: 25.000-26.000 --> 28.000-30.000 (Cont+Disc)

Char. 1: 3 --> 4 (All Disc)

Char. 83: 0 --> 2

Char. 140: 0 --> 1

Char. 141: 0 --> 1

Fasciculicardia subintermedia:

Char. 1: 25.000-26.000 --> 28.000-32.000 (Cont+Disc)

Char. 1: 3 --> 4 (All Disc)

Char. 2: 34.826 --> 46.054 (Cont+Disc)

Char. 2: 3 --> 4 (All Disc)

Fasciculicardia acanthodes:

Char. 70: 0 --> 1

Fasciculicardia sp.:

Char. 77: 1 --> 3

Char. 100: 1 --> 0

Char. 101: 1 --> 0

Char. 121: 4 --> 6 (All Disc)

Purpurocardia purpurata:

Char. 27: 1 --> 2

Char. 135: 1 --> 2

Purpurocardia elegantoides:

Char. 2: 18.344 --> 9.884 (Cont+Disc)

Char. 2: 2 --> 1 (All Disc)

Char. 40: 2 --> 1

Char. 117: 0 --> 1 (All Disc)

Char. 143: 1 --> 0

Cardites antiquatus:

Char. 70: 0 --> 1

Cardites partschii:

Char. 2: 18.094 --> 12.308 (Cont+Disc)

Char. 2: 2 --> 1 (All Disc)

Cardites feruglioi:

Char. 1: 23.000-26.000 --> 28.000-29.000 (Cont+Disc)

Char. 1: 3 --> 4 (All Disc)

Char. 2: 19.614-22.368 --> 28.256 (Cont+Disc)

Char. 2: 2 --> 3 (All Disc)

Char. 77: 3 --> 2 (All Disc)

Char. 143: 01 --> 2

Cardiocardita ajar:

Char. 4: 1 --> 2

Char. 5: 1 --> 0

Char. 9: 3 --> 1

Char. 12: 1 --> 0

Char. 13: 2 --> 3

Char. 15: 0 --> 1

Char. 21: 0 --> 1

Char. 23: 0 --> 2

Char. 27: 1 --> 0

Char. 48: 0 --> 1

Char. 56: 0 --> 2

Char. 57: 1 --> 3

Char. 77: 1 --> 3

Char. 81: 1 --> 2

Char. 103: 1 --> 3

Char. 131: 1 --> 0

Char. 135: 1 --> 0

Coripia unidentata:

Char. 2: 5.401-6.994 --> 3.551 (Cont+Disc)

Char. 6: 0 --> 1

Char. 12: 2 --> 0

Miodomeris cossmanni:

Char. 2: 2.363 --> 2.052 (Cont+Disc)

Char. 56: 2 --> 0

Char. 89: 1 --> 0

Char. 91: 1 --> 0

Char. 92: 1 --> 0

Char. 93: 2 --> 1

Char. 94: 2 --> 0

Char. 103: 1 --> 0

Char. 113: 3 --> 1

Char. 132: 0 --> 1

Scalaricardita scalaris:

No autapomorphies

Vimentum dilectum:

Char. 1: 25.000 --> 27.000-29.000 (Cont+Disc)

Char. 3: 3 --> 4 (All Disc)

Char. 13: 1 --> 2 (All Disc)

Char. 27: 0 --> 1 (All Disc)

Char. 83: 2 --> 1 (All Disc)

Char. 135: 2 --> 1 (All Disc)

Pleuromeris tridentata:

Char. 79: 2 --> 1

Char. 140: 0 --> 1

Pleuromeris decemcostata:

Char. 1: 17.000-18.000 --> 11.000-12.000 (Cont+Disc)

Char. 1: 2 --> 1 (All Disc)

Char. 2: 4.152 --> 3.943 (Cont+Disc)

Pteromeris perplana:

Char. 1: 15.000-18.000 --> 10.000-11.000 (Cont+Disc)

Char. 49: 2 --> 0

Char. 55: 0 --> 1

Char. 114: 0 --> 4

Pleuromeris fueguina:

No autapomorphies

Pleuromeris sulcolunularis:

Char. 2: 10.357 --> 11.208 (Cont+Disc)

Char. 39: 0 --> 1

Cyclocardia borealis:

Char. 2: 9.952-10.316 --> 27.007 (Cont+Disc)

Char. 2: 1 --> 3 (All Disc)

Darwinicardia patagonica:

Char. 20: 0 --> 1

Char. 36: 1 --> 0

Char. 40: 0 --> 1

Char. 79: 2 --> 1

Darwinicardia angusticostata:

Char. 2: 19.614-20.478 --> 13.314 (Cont+Disc)

Char. 2: 2 --> 1 (All Disc)

Char. 26: 0 --> 1

Char. 76: 0 --> 1

Venericor planicosta:

Char. 70: 1 --> 2

Bathycardita raouli:

Char. 1: 19.000-21.000 --> 14.000-16.000 (Cont+Disc)

Char. 1: 2 --> 1 (All Disc)

Char. 2: 35.973 --> 22.956 (Cont+Disc)

Char. 25: 2 --> 0

Char. 35: 0 --> 1

Char. 48: 0 --> 1

Char. 49: 1 --> 0

Char. 56: 0 --> 2

Char. 57: 1 --> 3

Char. 70: 1 --> 0 (Cont+Disc)

Char. 75: 1 --> 2

 Char. 103: 1 --> 0

Char. 128: 0 --> 1

Char. 130: 0 --> 1

Megacardita jouanetti:

Char. 26: 0 --> 1 (All Disc)

Char. 27: 1 --> 2

Char. 79: 2 --> 1 (All Disc)

Char. 112: 1 --> 0

Neovenericor paranensis:

No autapomorphies

Neovenericor ponderosa:

Char. 1: 3 --> 4 (All Disc)

Char. 2: 49.693 --> 49.943 (Cont+Disc)

Char. 40: 0 --> 1 (All Disc)

Venericor aposmithii:

No autapomorphies

Venericor bashiplata:

Char. 1: 5 --> 4 (All Disc)

Char. 2: 47.019-62.792 --> 42.536 (Cont+Disc)

Char. 70: 1 --> 2 (Cont+Disc)

Char. 113: 4 --> 3

Leuroactis pilsbryi:

Char. 15: 0 --> 1

Char. 26: 2 --> 1

Char. 27: 2 --> 1

Char. 49: 1 --> 0

Char. 50: 1 --> 2

Char. 53: 1 --> 0

Char. 65: 0 --> 1

Char. 75: 1 --> 0

Char. 114: 2 --> 4

Char. 135: 1 --> 2

Pacificor mulleri:

Char. 13: 1 --> 2 (All Disc)

Char. 26: 2 --> 1

Char. 114: 2 --> 4

Cyclocardia awamoensis:

No autapomorphies

Cyclocardia ventricosa:

Char. 83: 0 --> 2

Char. 121: 1 --> 2

Pleuromeris zelandica:

Char. 112: 0 --> 1 (All Disc)

Glans trapezia:

Char. 2: 16.609 --> 7.075 (Cont+Disc)

Char. 2: 2 --> 0 (All Disc)

Centrocardita aculeata:

Char. 63: 0 --> 1

Carditamera arata:

Char. 2: 18.209 --> 27.790 (Cont+Disc)

Char. 2: 2 --> 3 (All Disc)

Char. 135: 0 --> 1

Carditamera floridana:

Char. 124: 3 --> 0

Carditamera affinis:

Char. 2: 17.041-18.209 --> 42.043 (Cont+Disc)

Char. 2: 2 --> 4 (All Disc)

Carditamera gracilis:

Char. 57: 1 --> 0

Paraglans calcitrapoides:

Char. 1: 25.000-26.000 --> 23.000-24.000 (Cont+Disc)

Char. 2: 9.182 --> 5.239 (Cont+Disc)

Char. 2: 1 --> 0 (All Disc)

Char. 7: 1 --> 2
Scalaricardita camaronesia:

Char. 24: 2 --> 1 (Cont+Disc)

Char. 75: 3 --> 4 (Cont+Disc)

Cyclocardia cannada:

No autapomorphies

Neovenericor austroplata:

Char. 2: 35.973-42.536 --> 1.516 (Cont+Disc)

Char. 2: 4 --> 5 (All Disc)

Char. 3: 4 --> 3 (Cont+Disc)

Neovenericor carrerensis:

Char. 2: 35.973-42.536 --> 58.842 (Cont+Disc)

Char. 27: 1 --> 2 (Cont+Disc)

Char. 104: 1 --> 0

Scalaricardita laciarina:

Char. 2: 8.102 --> 9.736 (Cont+Disc)

Char. 2: 0 --> 1 (All Disc)

Char. 8: 0 --> 1

Char. 40: 2 --> 1

Kolmeris tehuelchana:

Char. 51: 0 --> 2

Char. 83: 0 --> 2 (Cont+Disc)

 Char. 99: 1 --> 2

Char. 110: 0 --> 1

Char. 126: 0 --> 1

Char. 135: 1 --> 2

Cyclocardia mesembria:

Char. 2: 9.952-10.148 --> 8.237 (Cont+Disc)

Char. 9: 0 --> 2 (All Disc)

Char. 40: 2 --> 0

Cyclocardia dalek:

Char. 113: 0 --> 2

Cyclocardia compressa:

Char. 1: 20.000-22.000 --> 15.000-17.000 (Cont+Disc)

Char. 1: 3 --> 2 (All Disc)

Char. 2: 9.952-10.316 --> 9.675 (Cont+Disc)

Cyclocardia nortensis:

Char. 2: 12.119 --> 19.350 (Cont+Disc)

Char. 2: 1 --> 2 (All Disc)

Char. 113: 0 --> 2

Char. 140: 1 --> 0

Purpurocardia leonensis:

Char. 73: 1 --> 0

Char. 116: 1 --> 2

Node 79 - Astartidae (Laevastarte basteroti + Astarte sulcata):

Char. 3: 4 --> 2

Char. 4: 1 --> 2

Char. 20: 0 --> 1

Char. 25: 2 --> 1

Char. 36: 0 --> 1

Char. 62: 12 --> 0

Char. 67: 2 --> 0

Char. 69: 5 --> 3 (Cont+Disc)

Char. 116: 2 --> 3

Char. 131: 2 --> 1

Char. 135: 0 --> 1 (Cont+Disc)

Char. 137: 0 --> 1

Node 80 - Astartidae + Carditidae:

Char. 2: 26.237 --> 16.609-22.395 (Cont+Disc)

Char. 15: 1 --> 0

Char. 31: 1 --> 0

Char. 33: 1 --> 0

Char. 35: 1 --> 0

Char. 37: 3 --> 1

Char. 44: 2 --> 1

Char. 45: 0 --> 1

Char. 52: 0 --> 1

Char. 59: 0 --> 1

Char. 80: 0 --> 1

 Char. 86: 0 --> 1

Char. 93: 0 --> 1

Char. 130: 1 --> 0

Node 81 - Palaeocardita (Palaeocardita stoecklini + Palaeocardita crenata):

Char. 2: 16.609-22.395 --> 15.529 (Cont+Disc)

Char. 94: 0 --> 1

Char. 112: 0 --> 1

Node 82 - Clade 1 (Palaeocarditinae here):

Char. 12: 0 --> 1

Char. 14: 0 --> 1

Char. 17: 0 --> 1

Char. 27: 0 --> 2 (All Disc)

Node 83 - Carditidae:

Char. 1: 0.000 --> 23.000 (Cont+Disc)

Char. 1: 0 --> 3 (All Disc)

Char. 28: 0 --> 1

Char. 46: 1 --> 0

Char. 73: 0 --> 1

Char. 90: 0 --> 1

Char. 91: 0 --> 1

Char. 95: 0 --> 1

Char. 97: 2 --> 0

Char. 98: 2 --> 1

Char. 99: 3 --> 1

 Char. 100: 2 --> 0

Char. 101: 4 --> 0

Char. 102: 3 --> 0

Char. 104: 3 --> 2

Char. 105: 4 --> 2

Char. 106: 2 --> 0

Char. 107: 3 --> 0

Char. 111: 2 --> 0

Char. 112: 3 --> 0

Char. 113: 5 --> 1

Char. 114: 5 --> 0

Char. 115: 2 --> 0

Char. 122: 2 --> 0

Char. 139: 0 --> 1

Node 84 - Clade 3 + Clade 11:

Char. 1: 23.000 --> 25.000-34.000 (Cont+Disc)

Char. 4: 1 --> 0 (All Disc)

Char. 7: 2 --> 0 (All Disc)

Char. 11: 0 --> 1

Char. 13: 3 --> 0 (All Disc)

Char. 14: 0 --> 1 (All Disc)

Char. 18: 1 --> 0 (All Disc)

Char. 25: 2 --> 0

Char. 49: 2 --> 0 (Cont+Disc)

Char. 51: 0 --> 1 (All Disc)

Char. 54: 0 --> 1

 Char. 57: 1 --> 0

Char. 58: 1 --> 0

Char. 62: 12 --> 0 (All Disc)

Char. 67: 1 --> 0

Char. 75: 34 --> 1 (Cont+Disc)

Char. 77: 3 --> 0

Char. 79: 2 --> 0 (Cont+Disc)

Char. 85: 1 --> 0 (All Disc)

Char. 98: 1 --> 0

Char. 99: 1 --> 0 (All Disc)

Char. 105: 2 --> 1 (All Disc)

Char. 116: 2 --> 0

Char. 134: 2 --> 0 (Cont+Disc)

Char. 143: 1 --> 0 (All Disc)

Node 85 – Clade 2 + (Clade 3+ Clade 11) (Carditida here):

Char. 24: 1 --> 0 (Cont+Disc)

Char. 44: 1 --> 0

Char. 53: 1 --> 0

Char. 56: 2 --> 0 (Cont+Disc)

Char. 57: 3 --> 1 (Cont+Disc)

Char. 66: 1 --> 0

Char. 81: 2 --> 0 (All Disc)

Char. 104: 2 --> 0

Char. 121: 2 --> 0 (All Disc)

Char. 142: 1 --> 0

Node 86 – All Carditidae excluding Clade 1 (Eucarditidae here):

Char. 22: 1 --> 0

Char. 23: 1 --> 0

Char. 30: 1 --> 0

Char. 31: 0 --> 2

Char. 33: 0 --> 2

Char. 39: 1 --> 0

Char. 48: 3 --> 0

Char. 50: 4 --> 0

Char. 57: 45 --> 3

Char. 58: 3 --> 1

Char. 67: 2 --> 1

Char. 71: 0 --> 1

Char. 81: 4 --> 2 (All Disc)

Char. 89: 0 --> 1

Char. 92: 0 --> 1

Char. 93: 1 --> 2

Char. 94: 0 --> 2

Char. 120: 0 --> 1

Char. 121: 7 --> 2 (All Disc)

Char. 125: 3 --> 0

Char. 126: 2 --> 0

Char. 127: 2 --> 0

Char. 128: 3 --> 0

Char. 129: 2 --> 0

Char. 131: 2 --> 0

Char. 132: 1 --> 0

 Char. 133: 0 --> 1

Node 87 - Clade 11 (Thecaliinae):

Char. 2: 16.211 --> 8.993 (Cont+Disc)

Char. 2: 2 --> 1 (All Disc)

Char. 19: 0 --> 1

Char. 49: 0 --> 2 (Cont+Disc)

Char. 75: 1 --> 4 (Cont+Disc)

Char. 79: 0 --> 1 (Cont+Disc)

Char. 113: 0 --> 1 (Cont+Disc)

Node 88 - Cardita crassa + Clade 11:

Char. 1: 25.000 --> 21.000-24.000 (Cont+Disc)

Char. 2: 16.609-16.879 --> 16.211 (Cont+Disc)

Node 89 - Cardita variegata (Cardita crassa + Clade 11):

Char. 49: 2 --> 0 (All Disc)

Char. 75: 4 --> 1 (All Disc)

Char. 113: 1 --> 0 (All Disc)

Char. 143: 1 --> 0 (Cont+Disc)

Node 90 - Beguina semiorbiculata + Node 89:

Char. 14: 0 --> 1 (Cont+Disc)

Char. 18: 1 --> 0 (Cont+Disc)

Char. 85: 1 --> 0 (Cont+Disc)

Char. 99: 1 --> 0 (Cont+Disc)

Char. 113: 1 --> 0 (Cont+Disc)

Node 90a - Beguina semiorbiculata + Birkelundita turoniense:

Char. 1: 3 --> 5 (All Disc)

Char. 2: 2 --> 3 (All Disc)

Char. 34: 0 --> 1 (All Disc)

Char. 143: 0 --> 1 (All Disc)

Node 90b - Clade 3 (Cardita variegata + (Cardita crassa + Node 90a)) (Carditinae
here):

Char. 49: 2 --> 0 (All Disc)

Char. 75: 4 --> 1 (All Disc)

Char. 113: 1 --> 0 (All Disc)

Node 91 - Claibornicardia alticostata + Claibornicardia paleopatagonica:

Char. 2: 20.937-22.368 --> 25.083 (Cont+Disc)

Char. 3: 2 --> 4 (All Disc)

Char. 5: 1 --> 0

Char. 20: 0 --> 1

Node 91a - Claibornicardia:

Char. 1: 3 --> 5 (All Disc)

Char. 2: 2 --> 3 (All Disc)

Char. 7: 1 --> 2 (All Disc)

Char. 15: 0 --> 1 (All Disc)

Char. 107: 0 --> 1 (All Disc)

Node 92 - Baluchicardia + Claibornicardia:

 Char. 3: 5 --> 2 (All Disc)

Char. 26: 0 --> 1 (Cont+Disc)

Char. 102: 1 --> 2

Char. 103: 1 --> 2

Char. 113: 1 --> 4 (Cont+Disc)

Char. 114: 1 --> 4 (Cont+Disc)

Node 93 - Clade 17:

Char. 41: 0 --> 1

Char. 42: 0 --> 1

Char. 57: 1 --> 0

Char. 76: 0 --> 2

Node 94 - Clade 16 + Clade 17:

Char. 37: 0 --> 1 (Cont+Disc)

Char. 75: 1 --> 0

Char. 113: 0 --> 1 (Cont+Disc)

Char. 114: 0 --> 1 (Cont+Disc)

Char. 131: 0 --> 1 (All Disc)

Node 95 - Cardites + (Clade 16 + Clade 17):

Char. 7: 1 --> 2 (Cont+Disc)

Char. 57: 2 --> 1 (Cont+Disc)

Char. 131: 0 --> 1 (Cont+Disc)

Node 96 - Darwinicardia + (Cardites + (Clade 16 + Clade 17):

Char. 38: 0 --> 1 (Cont+Disc)

 Char. 40: 2 --> 0 (Cont+Disc)

Char. 43: 0 --> 1 (Cont+Disc)

Char. 62: 1 --> 2 (Cont+Disc)

Char. 125: 0 --> 2 (Cont+Disc)

Node 96a - Darwinicardia + (Clade 16 + Clade 17):

Char. 3: 4 --> 5 (All Disc)

Char. 5: 0 --> 1 (All Disc)

Char. 107: 2 --> 0 (All Disc)

Char. 121: 2 --> 4 (All Disc)

Node 97 - Clade 9 including Cardites:

Char. 25: 2 --> 0 (Cont+Disc)

Char. 36: 0 --> 1 (Cont+Disc)

Char. 41: 1 --> 0 (Cont+Disc)

Char. 42: 1 --> 0 (Cont+Disc)

Char. 100: 0 --> 1 (Cont+Disc)

Char. 102: 0 --> 1 (Cont+Disc)

Node 97a - Clade 9 excluding Cardites (Venericardiini here):

Char. 7: 2 --> 1 (All Disc)

Char. 25: 2 --> 0 (All Disc)

Char. 77: 3 --> 1 (All Disc)

Char. 82: 0 --> 1 (All Disc)

Char. 100: 0 --> 1 (All Disc)

Char. 101: 0 --> 1 (All Disc)

Char. 102: 0 --> 1 (All Disc)

Node 97b - Cardites + Clade 9

Char. 13: 3 --> 2 (All Disc)

Char. 36: 0 --> 1 (All Disc)

Char. 137: 0 --> 1 (All Disc)

Node 98 - Clade 13 (Venericardiinae):

Char. 2: 9.952-16.879 --> 19.614-20.478 (Cont+Disc)

Char. 39: 0 --> 2

Char. 43: 0 --> 1 (All Disc)

Char. 49: 2 --> 1

Char. 56: 2 --> 0 (Cont+Disc)

Char. 61: 0 --> 1 (Cont+Disc)

Char. 69: 1 --> 3 (Cont+Disc)

Char. 75: 3 --> 1

Char. 107: 0 --> 2

Char. 116: 2 --> 1 (All Disc)

Node 99 - (Clade 5 + Clade 6) + Clade 13:

Char. 77: 3 --> 1 (Cont+Disc)

Char. 103: 0 --> 1 (Cont+Disc)

Char. 124: 02 --> 14 (Cont+Disc)

Node 100 – (Clade 4 + Clade 10) + ((Clade 5 + Clade 6) + Clade 13):

Char. 113: 1 --> 0 (Cont+Disc)

Char. 116: 2 --> 1 (Cont+Disc)

Char. 117: 0 --> 1 (Cont+Disc)

 Char. 134: 2 --> 1 (Cont+Disc)

Char. 135: 0 --> 1 (Cont+Disc)

Char. 137: 0 --> 1 (Cont+Disc)

Node 101 - Baluchicardia beaumonti + Claibornicardia acuticostata:

Char. 3: 4 --> 2 (Cont+Disc)

Char. 37: 1 --> 0 (Cont+Disc)

Char. 43: 1 --> 0 (Cont+Disc)

Node 102 - Rotundicardia rotunda + Rotundicardia mariobrosorum:

Char. 15: 0 --> 1 (All Disc)

Char. 49: 1 --> 0

Node 103 - Rotundicardia:

Char. 1: 3 --> 4 (All Disc)

Char. 2: 20.937-22.368 --> 16.825-19.560 (Cont+Disc)

Char. 3: 4 --> 5 (Cont+Disc)

Char. 7: 2 --> 1 (Cont+Disc)

Char. 9: 2 --> 0

Char. 100: 1 --> 0

Char. 101: 1 --> 0

Char. 102: 1 --> 0

Char. 107: 12 --> 0 (Cont+Disc)

Char. 112: 0 --> 1

Char. 116: 1 --> 2

Char. 121: 24 --> 5

Char. 140: 0 --> 1

 Char. 144: 0 --> 1

Node 104 - Rotundicardia + (Glyptoactis + Fasciculicardia):

Char. 67: 1 --> 0

Char. 69: 3 --> 1

Node 105 - Kalelia:

Char. 2: 20.937-22.368 --> 29.384 (Cont+Disc)

Char. 4: 1 --> 2

Char. 7: 1 --> 2 (All Disc)

Char. 13: 2 --> 1

Char. 22: 0 --> 1

Char. 35: 0 --> 1

Char. 40: 0 --> 1

Char. 51: 0 --> 1

Char. 98: 1 --> 0

Char. 99: 1 --> 2

Char. 101: 1 --> 3

Char. 105: 2 --> 1

Char. 107: 0 --> 2 (All Disc)

Node 106 - Clade 16:

Char. 9: 2 --> 3

Char. 12: 0 --> 1

Char. 24: 1 --> 2

Char. 69: 3 --> 2

Char. 81: 2 --> 1

 Char. 123: 1 --> 0 (All Disc)

Node 107 - Kalelia multicostata + Kalelia pectuncularis:

Char. 2: 29.384 --> 34.090 (Cont+Disc)

Char. 39: 2 --> 0

Char. 48: 0 --> 1

Char. 57: 1 --> 2

Char. 70: 0 --> 1

Node 108 - Glyptoactis + Fasciculicardia:

Char. 2: 20.937-22.368 --> 22.686 (Cont+Disc)

Char. 2: 2 --> 3 (All Disc)

Char. 3: 5 --> 4 (All Disc)

Char. 7: 1 --> 2 (All Disc)

Char. 27: 1 --> 2

Char. 49: 1 --> 2

Char. 51: 0 --> 1

Char. 76: 2 --> 1

Char. 78: 0 --> 1

Char. 81: 2 --> 1

Node 109 - Venericardia + (Cardiocardita + (Paraglans + Arcturellina):

Char. 7: 2 --> 1 (Cont+Disc)

Char. 43: 1 --> 0

Char. 107: 2 --> 0 (Cont+Disc)

Char. 112: 0 --> 1

Char. 116: 1 --> 2

 Char. 117: 0 --> 1

Char. 125: 2 --> 0

Char. 144: 0 --> 1

Node 110 - Paraglans + Arcturellina:

Char. 2: 13.503 --> 9.182 (Cont+Disc)

Char. 38: 1 --> 0

Char. 40: 0 --> 2

Char. 76: 0 --> 1

Char. 78: 0 --> 1

Node 111 - Cardiocardita + (Paraglans + Arcturellina):

Char. 2: 20.937 --> 13.503 (Cont+Disc)

Char. 2: 2 --> 1 (All Disc)

Char. 37: 1 --> 0 (Cont+Disc)

Char. 44: 1 --> 0

Char. 49: 1 --> 2

Char. 75: 0 --> 3

Node 112 - Fasciculicardia acanthodes + Fasciculicardia subintermedia:

Char. 2: 23.361 --> 34.826 (Cont+Disc)

Char. 43: 1 --> 0

Char. 75: 1 --> 2

Node 113 - Fasciculicardia:

Char. 2: 22.686 --> 23.361 (Cont+Disc)

Char. 37: 1 --> 0 (Cont+Disc)

 Char. 41: 1 --> 0

Char. 42: 1 --> 0

Char. 75: 0 --> 1

Char. 103: 1 --> 2

Char. 129: 0 --> 1

Char. 143: 0 --> 1

Node 114 - Purpurocardia elegantoides + Purpurocardia purpurata:

Char. 2: 19.614 --> 18.344 (Cont+Disc)

Char. 49: 1 --> 0

Char. 75: 1 --> 3

Char. 105: 2 --> 1 (All Disc)

Node 115 - Purpurocardia:

Char. 5: 1 --> 0 (Cont+Disc)

Char. 35: 0 --> 1

Char. 40: 0 --> 2 (All Disc)

Char. 43: 1 --> 0 (All Disc)

Char. 48: 0 --> 1

Char. 50: 0 --> 1

Char. 51: 1 --> 0

Char. 57: 12 --> 3 (All Disc)

Char. 84: 0 --> 2

Char. 98: 1 --> 0

Char. 113: 0 --> 1 (Cont+Disc)

Char. 114: 0 --> 1 (Cont+Disc)

Char. 136: 0 --> 1

Node 116 - Cardites partschii + Cardites antiquatus:

Char. 2: 19.614-22.368 --> 18.094 (Cont+Disc)

Node 117 - Cardites:

Char. 4: 1 --> 2

Char. 5: 1 --> 0 (Cont+Disc)

Char. 25: 0 --> 2 (Cont+Disc)

Char. 26: 0 --> 2

Char. 48: 0 --> 2

Char. 49: 1 --> 3

Char. 50: 0 --> 3

Char. 51: 0 --> 3

Char. 76: 0 --> 1

Char. 99: 1 --> 2 (Cont+Disc)

Char. 100: 1 --> 0 (Cont+Disc)

Char. 102: 1 --> 0 (Cont+Disc)

Char. 104: 2 --> 1

Char. 113: 1 --> 0 (All Disc)

Char. 131: 0 --> 1 (All Disc)

Node 118 - Coripia + Scalaricardita:

Char. 2: 4 --> 3 (Cont+Disc)

Char. 26: 1 --> 0 (Cont+Disc)

Char. 134: 1 --> 2 (Cont+Disc)

Node 118b - Coripia + Vimentum:

 Char. 116: 3 --> 1 (All Disc)

Node 119 - Clade 4 (Scalaricarditinae here):

Char. 1: 23.000 --> 25.000 (Cont+Disc)

Char. 2: 9.952-10.148 --> 5.401-6.994 (Cont+Disc)

Char. 2: 1 --> 0 (All Disc)

Char. 9: 0 --> 2 (All Disc)

Char. 24: 1 --> 2 (Cont+Disc)

Char. 36: 0 --> 1

Char. 62: 1 --> 0 (Cont+Disc)

Char. 99: 1 --> 0

Char. 104: 2 --> 1

Char. 122: 0 --> 1

Char. 135: 1 --> 2 (All Disc)

Node 120 - Clade 4 + Clade 10:

Char. 4: 1 --> 3 (Cont+Disc)

Char. 12: 0 --> 2

Char. 25: 2 --> 0

Char. 82: 0 --> 2 (All Disc)

Char. 105: 12 --> 0

Char. 125: 0 --> 2

Node 121 - Pteromeris + Miodomeris:

Char. 2: 8.487 --> 2.363 (Cont+Disc)

Char. 2: 1 --> 0 (All Disc)

Char. 6: 1 --> 2
Node 122 - Clade 5 (Miodomeridinae here):

Char. 3: 3 --> 1 (All Disc)

Char. 6: 0 --> 1

Char. 13: 1 --> 0 (All Disc)

Char. 25: 2 --> 1

Char. 81: 2 --> 3

Char. 103: 2 --> 1 (All Disc)

Char. 104: 2 --> 1

Char. 105: 12 --> 0

Char. 113: 0 --> 3

Char. 121: 2 --> 1 (All Disc)

Char. 123: 1 --> 4 (All Disc)

Node 123 - Clade 5 + Clade 6:

Char. 1: 23.000 --> 17.000-18.000 (Cont+Disc)

Char. 1: 3 --> 2 (All Disc)

Char. 4: 1 --> 2 (Cont+Disc)

Char. 24: 1 --> 2

Char. 27: 1 --> 0 (Cont+Disc)

Char. 35: 0 --> 1

Char. 62: 1 --> 0 (Cont+Disc)

Char. 77: 3 --> 1 (All Disc)

Char. 116: 1 --> 3 (Cont+Disc)

Node 124 - Scalaricardita laciarina + Scalaricardita scalaris:

Char. 2: 6.994 --> 8.102 (Cont+Disc)

Node 125 - Scalaricardita:

Char. 116: 1 --> 3 (Cont+Disc)

Node 126 - Pleuromeris decemcostata + Pleuromeris tridentata:

Char. 2: 5.401 --> 4.152 (Cont+Disc)

Char. 123: 1 --> 0

Node 127 - Pleuromeris zelandica + (Pleuromeris decemcostata + Pleuromeris

tridentata):

Char. 2: 8.487-10.357 --> 5.401 (Cont+Disc)

Char. 2: 1 --> 0 (All Disc)

Char. 9: 0 --> 1

Char. 75: 3 --> 4 (Cont+Disc)

Char. 83: 0 --> 2 (Cont+Disc)

Node 128 - Pleuromeris:

Char. 103: 1 --> 2 (Cont+Disc)

Node 129 - Pleuromeris sulcolunularis + Pleuromeris fueguina:

Char. 26: 0 --> 1

Char. 49: 2 --> 0

Char. 61: 0 --> 1

Char. 112: 0 --> 1 (All Disc)

Node 130 - Cyclocardia compressa + Cyclocardia borealis:

Char. 83: 0 --> 1

Node 131 – ((Cyclocardia compressa + Cyclocardia borealis) + Node 146):

Char. 35: 0 --> 1

Node 132 - Cyclocardia mesembria + Node 131:

Char. 1: 23.000 --> 21.000-22.000 (Cont+Disc)

Char. 49: 2 --> 1

Node 133 - Cyclocardia:

Char. 3: 4 --> 5 (Cont+Disc)

Char. 3: 3 --> 5 (All Disc)

Char. 13: 1 --> 2 (All Disc)

Char. 27: 0 --> 1 (All Disc)

Char. 41: 1 --> 0

Char. 42: 1 --> 0

Char. 51: 0 --> 1

Char. 98: 1 --> 0

Char. 106: 0 --> 1

Char. 107: 0 --> 2

Char. 116: 3 --> 1 (All Disc)

Char. 130: 0 --> 1

Node 133b - Clade 12 (Cyclocardida here)

Char. 2: 2 --> 1 (All Disc)

Char. 3: 4 --> 3 (All Disc)

Char. 13: 3 --> 1 (All Disc)

Char. 37: 0 --> 1 (All Disc)

 Char. 113: 1 --> 0 (All Disc)

Char. 116: 2 --> 3 (All Disc)

Char. 137: 0 --> 1 (All Disc)

Node 134 - Darwinicardia:

Char. 3: 4 --> 5 (Cont+Disc)

Char. 4: 1 --> 3

Char. 49: 1 --> 0

Char. 51: 0 --> 2

Char. 69: 3 --> 4

Char. 101: 01 --> 2 (Cont+Disc)

Char. 101: 1 --> 2 (All Disc)

Char. 103: 1 --> 2

Char. 107: 2 --> 0 (Cont+Disc)

Char. 112: 0 --> 1

Char. 113: 1 --> 0 (All Disc)

Char. 117: 0 --> 1 (All Disc)

Char. 144: 0 --> 1

Node 135 - Venericor planicosta + Node 143:

Char. 2: 42.536 --> 47.019 (Cont+Disc)

Char. 27: 1 --> 2 (Cont+Disc)

Node 136 - Clade 14:

Char. 1: 23.000-24.000 --> 30.000-31.000 (Cont+Disc)

Char. 1: 3 --> 5 (All Disc)

Char. 5: 0 --> 1 (All Disc)

 Char. 9: 1 --> 2 (All Disc)

Char. 12: 2 --> 0 (All Disc)

Char. 15: 1 --> 0 (All Disc)

Char. 24: 0 --> 1 (All Disc)

Char. 29: 1 --> 0

Char. 40: 0 --> 2 (All Disc)

Char. 44: 1 --> 2

Char. 48: 0 --> 1

Char. 50: 0 --> 1

Char. 55: 0 --> 2

Char. 57: 1 --> 2 (All Disc)

Char. 62: 1 --> 0

Char. 79: 0 --> 1 (All Disc)

Char. 81: 1 --> 2 (All Disc)

Char. 99: 2 --> 1 (All Disc)

Char. 105: 1 --> 0

Char. 106: 0 --> 1

Char. 112: 1 --> 0 (All Disc)

Char. 113: 3 --> 4 (All Disc)

Char. 130: 0 --> 1

Node 136a – Neovenericor carrerensis + Clade 14:

Char. 43: 1 --> 0

Node 136b – Neovenericor austroplata + Node 136a:

Char. 77: 0 --> 1

Char. 126: 0 --> 1

Node 137 - Clade 8 (Venericorini here):

Char. 2: 19.614-20.478 --> 35.973-42.536 (Cont+Disc)

Char. 2: 2 --> 3 (All Disc)

Char. 7: 1 --> 2 (Cont+Disc)

Char. 12: 0 --> 2 (All Disc)

Char. 22: 0 --> 1

Char. 26: 0 --> 2 (Cont+Disc)

Char. 70: 0 --> 1 (Cont+Disc)

Char. 77: 3 --> 0 (All Disc)

Char. 81: 2 --> 1 (All Disc)

Char. 83: 0 --> 1 (Cont+Disc)

Char. 104: 2 --> 1 (Cont+Disc)

Char. 105: 2 --> 1 (All Disc)

Char. 114: 0 --> 2

Char. 121: 2 --> 1 (All Disc)

Char. 126: 0 --> 1 (Cont+Disc)

Char. 131: 0 --> 2

Char. 132: 0 --> 1

Char. 137: 1 --> 0 (Cont+Disc)

Char. 143: 1 --> 2

Node 138 - Megacardita + Bathycardita:

Char. 1: 22.000-24.000 --> 19.000-21.000 (Cont+Disc)

Char. 1: 3 --> 2 (All Disc)

Char. 3: 4 --> 6

Char. 13: 1 --> 3 (Cont+Disc)

 Char. 40: 0 --> 1 (All Disc)

Char. 82: 2 --> 0 (Cont+Disc)

Char. 83: 1 --> 0 (Cont+Disc)

Char. 104: 1 --> 2 (Cont+Disc)

Char. 105: 1 --> 0

Char. 106: 0 --> 1

Char. 134: 1 --> 0

Node 139 - Node 138 + Node 142:

Char. 77: 1 --> 0 (Cont+Disc)

Char. 126: 1 --> 0 (Cont+Disc)

Node 140 - Neovenericor austroplata + Node 139:

Char. 43: 0 --> 1 (Cont+Disc)

Node 141 - Clade 15:

Char. 5: 1 --> 0 (Cont+Disc)

Char. 12: 0 --> 2 (Cont+Disc)

Char. 15: 0 --> 1 (Cont+Disc)

Char. 24: 1 --> 0 (Cont+Disc)

Char. 40: 2 --> 0 (Cont+Disc)

Char. 57: 2 --> 1 (Cont+Disc)

Char. 81: 2 --> 1 (Cont+Disc)

Char. 99: 1 --> 2 (Cont+Disc)

Char. 112: 0 --> 1 (Cont+Disc)

Node 142 - Neovenericor ponderosa + Neovenericor paranensis:

 Char. 2: 35.973-42.536 --> 49.693 (Cont+Disc)

Char. 76: 0 --> 1

Char. 127: 0 --> 1

Node 143 - Venericor aposmithii + Node 144:

Char. 38: 0 --> 2 (Cont+Disc)

Char. 39: 2 --> 0 (Cont+Disc)

Char. 81: 2 --> 3 (Cont+Disc)

Node 143a - Venericor bashiplata + Node 144a:

Char. 38: 0 --> 2 (All Disc)

Char. 39: 2 --> 0 (All Disc)

Char. 81: 2 --> 3 (All Disc)

Node 144 - Leuroactis pilsbryi + Venericor bashiplata:

Char. 57: 2 --> 3 (Cont+Disc)

Node 144a - Leuroactis pilsbryi + Venericor aposmithii:

Char. 2: 4 --> 5 (All Disc)

Node 145 - Cyclocardia nortensis + Cyclocardia awamoensis:

Char. 2: 10.316 --> 12.119 (Cont+Disc)

Char. 40: 2 --> 0

Char. 51: 1 --> 0

Char. 55: 0 --> 1

Char. 57: 3 --> 2

Char. 79: 2 --> 1

 Char. 98: 0 --> 1

Char. 104: 2 --> 0

Node 146 - Cyclocardia ventricosa + Node 145:

Char. 38: 0 --> 1

Char. 69: 1 --> 3

Node 147 - Centrocardita + Glans:

Char. 36: 0 --> 1

Char. 117: 0 --> 1

Node 148 - Clade 2 (Carditamera + Node 147 or Node 148a) (Carditamerinae here):

Char. 1: 23.000 --> 20.000-21.000 (Cont+Disc)

Char. 12: 0 --> 1

Char. 15: 0 --> 1 (Cont+Disc)

Char. 21: 0 --> 1

Char. 26: 0 --> 1

Char. 38: 0 --> 1

Char. 40: 0 --> 1 (All Disc)

Char. 48: 0 --> 1

Char. 74: 0 --> 1

Char. 76: 0 --> 1

Char. 84: 0 --> 2

Char. 114: 0 --> 1 (All Disc)

Char. 144: 0 --> 1 (All Disc)

Node 148a - Carditamera + Glans:

 Char. 1: 3 --> 2 (All Disc)

Char. 17: 0 --> 1 (All Disc)

Char. 27: 1 --> 2 (All Disc)

Node 149 - Carditamera floridana + Carditamera arata:

Char. 112: 0 --> 1

Char. 134: 2 --> 1

Char. 142: 0 --> 1

Node 150 - Carditamera:

Char. 2: 16.609-16.879 --> 17.041-18.209 (Cont+Disc)

Char. 4: 1 --> 0 (All Disc)

Char. 9: 3 --> 1 (All Disc)

Char. 14: 0 --> 1

Char. 41: 0 --> 1 (All Disc)

Char. 42: 0 --> 1 (All Disc)

Char. 47: 0 --> 1

Char. 72: 1 --> 0

Char. 88: 0 --> 1

Char. 103: 0 --> 1

Char. 104: 0 --> 1

Char. 113: 1 --> 2

Char. 114: 1 --> 2 (All Disc)

Char. 121: 0 --> 2 (All Disc)

Char. 136: 0 --> 1

Node 151 - Carditamera gracilis + Carditamera affinis:

 Char. 25: 2 --> 0

Char. 40: 1 --> 0 (All Disc)

Char. 61: 1 --> 0 (All Disc)

Char. 83: 0 --> 2

Char. 99: 1 --> 0

Char. 105: 2 --> 1 (All Disc)

Char. 108: 0 --> 1

Node 152 - Cyclocardia dalek + Cyclocardia cannada:

Char. 75: 3 --> 4 (Cont+Disc)

Char. 144: 0 --> 1

Appendix 6

Stratigraphic data of taxa used in analyses. Ages expressed in millions years. Zero
numbers indicated extant taxa.

Crassatella ponderosa 41.2 37.8

Spissatella trailli 28.1 15.97

Astarte sulcata 2.58 0

Laevastarte basteroti 3.6 0

Palaeocardita stoecklini 227 201.3

Palaeocardita crenata 221 201.3

Schizocardita cristata 227 208.5

Birkelundita turoniense 93.9 66

Thecalia concamerata 0 0

Powellina brookesi 0 0

Beguina semiorbiculata 0 0

Cardita variegata 2.58 0

Cardita crassa 20.44 13.82

Claibornicardia paleopatagonica 66 61.6

Claibornicardia alticostata 56 37.8

Claibornicardia acuticostata 41.2 37.8

Rotundicardia mariobrosorum 66 61.6

Rotundicardia rotunda 41.2 37.8

Rotundicardia diversidentata 37.8 33.9

Kalelia burmeisteri 66 61.6

Kalelia pectuncularis 59.2 56

Kalelia multicostata 59.2 56

Glyptoactis hadra 20.34 15.97

Baluchicardia beaumonti 72.1 61.6

Venericardia imbricata 41.2 37.8

Arcturellina asperula 41.2 37.8

Fasciculicardia subintermedia 23.03 15.97

Fasciculicardia acanthodes 41.2 37.8

Fasciculicardia sp. 11.63 5.33

Purpurocardia purpurata 3.6 0

Purpurocardia elegantoides 37.8 33.9

Purpurocardia leonensis 11.63 5.33

Cardites antiquatus 23.03 0

Cardites partschii 23.03 11.63

Cardites feruglioi 66 62

Cardiocardita ajar 0 0

Coripia unidentata 5.33 0

Miodomeris cossmanni 41.2 37.8

Scalaricardita scalaris 11.63 2.58

Scalaricardita camaronesia 15.97 13.82

Scalaricardita laciarina 5.33 3.6

Vimentum dilectum 0 0

Pleuromeris tridentata 3.6 0

Pleuromeris decemcostata 5.33 0.781

Pleuromeris zelandica 3.6 0

Pleuromeris fueguina 20.44 15.97

Pleuromeris sulcolunularis 28.1 20.44

Pteromeris perplana 5.33 0

Cyclocardia borealis 2.58 0

Cyclocardia awamoensis 23.03 15.97

Cyclocardia ventricosa 0 0

Cyclocardia cannada 28.1 15.97

Cyclocardia mesembria 47.8 37.8

Cyclocardia dalek 28.1 15.97

Cyclocardia compressa 0.126 0

Cyclocardia nortensis 11.63 5.33

Darwinicardia patagonica 28.1 20.44

Darwinicardia angusticostata 41.2 37.8

Venericor planicosta 41.2 37.8

Venericor aposmithii 59.2 56

Venericor bashiplata 56 47.8

Bathycardita raouli 0 0

Megacardita jouanetti 23.03 5.33

Neovenericor paranensis 11.63 5.33

Neovenericor ponderosa 28.1 23.03

Neovenericor austroplata 15.97 13.82

Neovenericor carrerensis 37.8 33.9

Leuroactis pilsbryi 59.2 56

Pacificor mulleri 37.8 33.9

Glans trapezia 0 0

Centrocardita aculeata 0 0

Carditamera arata 23.03 0

Carditamera floridana 0.781 0

Carditamera affinis 5.33 0

Carditamera gracilis 5.33 0

Paraglans calcitrapoides 41.2 37.8

 Kolmeris tehuelchana 5.33 3.6

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