J. foveolata
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in
Cyamioidea
and that the species is not closely related to
Gomphina undulosa
(type species of
Gomphina
) or other Veneroidea, as previously thought.
Considering the number, morphology and degree of development of cardinal teeth, the presence of an internal and an external ligament, the numerous tentacles at the posterior part of mantle margin, the long byssus groove and the left and right inner demibranchs being posteroventrally fused by a short distance,
J. foveolata
shows greater similarity to
Cyamiidae
than to
Gaimardiidae
. However, it differs clearly from all other
Cyamiidae
studied herein by having both inhalant and exhalant apertures projected as siphons and crowned by tentacles, related to the presence of a well-marked pallial sinus, and by having the series of posterior tentacles arising from the distal part of the middle mantle fold instead of arising from the base of the inner mantle fold as in
Cyamiocardium
,
Cyamium
,
Gaimardia
,
Heteromactra
,
Kidderia
,
Pseudokellya
and
Ptychocardia
. Furthermore, the anterior labial palps are subquadrate, and the foot appears to lack a differentiated stalk, a condition not seen in any other
Cyamiidae
. In contrast, unlike any other
Cyamiidae
and
Gaimardiidae
, the foot of
J. foveolata
is compressed and has a narrow base. The molecular information obtained from the two markers considered herein does not allow us to be conclusive whether
J. foveolata
is a
Cyamiidae
or belongs to a different family in
Cyamioidea
. It is interesting to note the similarity in shell morphology and gross anatomy of this species to
Cyamiomactra problematica
(type species of the genus), although the restricted knowledge of that species does not allow us to be conclusive regarding whether both taxa should be reunited into the same genus.
OTHER TAXA PREVIOUSLY ASSIGNED TO
CYAMIOIDEA
Besides the taxa considered above, 11 other (sub) genera (
Costokidderia
,
Cyamiomactra
,
Cyamionema
, Dicranodesma,
Eugaimardia
,
Legrandina
,
Lutetina
,
Neogaimardia
,
Perrierina, Progaimardia
and
Reloncavia
) were previously mentioned in the literature as belonging to either
Cyamiidae
or
Gaimardiidae
, and seven other families (
Bernardinidae
,
Basterotiidae
,
Galatheavalvidae
,
Juliidae
,
Neoleptonidae
,
Sportellidae
and
Turtoniidae
) were assigned to
Cyamioidea
/Gaimardioidea.
TAXA HERE REGARDED AS PROBABLY BELONGING TO
CYAMIOIDEA
Reloncavia
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(type species:
Kingiella chilenica Soot-Ryen, 1959
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)
The hinge of this genus (figured by Soot-Ryen, 1959: figs 14, 15; and Chavan, 1969: fig. E39, 3 b, 3c) shows great similarity with that of
Cyamium
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. Soot-Ryen (1957, 1959) described the gross anatomy of
Reloncavia chilenica
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, reporting the presence of a large pedal opening and two smaller (inhalant and exhalant) openings, a large foot, small labial palps and small posterior ‘papillae’. Gallardo (1993) described how ‘each embryo is contained in a small capsule attached to a branchial filament by a short peduncle’. The ‘capsule’ and ‘peduncle’ are likely to correspond to the follicular epithelium recognized here as diagnostic for
Cyamioidea
. All morphological and anatomical characters suggest
Reloncavia
View in CoL
to be a
Cyamiidae
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.
Cyamiomactra
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(type species:
Cyamiomactra problematica Bernard, 1897
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)
The description of this genus was not given separately from that of the type species. Bernard (1897) emphasized the similarity of this genus to
Cyamium
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, pointing out as the main differences the shell outline (‘
Cyamium
View in CoL
is much more elongated’) and the presence of posterior cardinal teeth in the left valve (regarded as absent in
Cyamium
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, but as a consequence of an erroneous interpretation of the hinge; see above). After studying the gross anatomy of
Cyamiomactra problematica, Ponder (1971)
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concluded that it ‘agrees closely with
Cyamium antarcticum
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’. In fact, the author considered
Cyamiomactra
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as a subgenus of
Cyamium
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. In its shell outline,
Cyamiomactra
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does not show great morphological differences from
Cyamium
View in CoL
.
Perrierina
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(type species:
Perrierina taxodonta Bernard, 1897
View in CoL
)
The genus has the same number, morphology and arrangement of cardinal teeth as
Cyamiomactra problematica
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. This fact was pointed out in its description ( Bernard, 1897) and ratified by subsequent authors (e.g. Fleming, 1948). The only difference is the presence of several lateral ‘crests’ along the dorsal margin in
Perrierina
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(diversely referred as lateral teeth, marginal denticles, lateral lamellae, taxodontlike laterals or taxodont lamellae). Ponder (1971) studied the gross anatomy of the type species of
Perrierina
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, which, although similar to that of
Cyamium
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and
Cyamiomactra
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, was described as lacking a byssus groove in the foot and tentacles at the posterior end of the mantle margin (two characters present in the other species of
Cyamioidea
studied herein). Despite that, Ponder (1971) concluded that
Perrierina
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‘is clearly derived from a
Cyamium
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or
Cyamiomactra
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- like ancestor’. In addition, he considered
Legrandina
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to be a subgenus of
Perrierina
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. We have no additional evidence to confirm or reject Ponder’s (1971) opinion. Consequently, we follow the family placement proposed by Ponder (1971) for those taxa.
Neogaimardia
(type species:
Kellia rostellata
Tate
, 1889)
Since its description,
Neogaimardia
has been considered a member of the
Gaimardiidae
. Odhler (1924) compared it with
Gaimardia
, and Ponder (1971) and Huber (2010) considered it to be a subgenus of
Gaimardia
. Morton (1979) provided valuable anatomical information of
Neogaimardia finlayi Powell, 1933
, a species very similar to
N. rostellata
. The main differences of
Neogaimardia
(with respect to
Gaimardia
) are the presence of a short internal ligament, well-defined posterior lateral teeth, accessory ‘marginal teeth’ and the presence of only one (the inner) demibranch at each side. Furthermore, judging from Morton’s (1979) figure 5, the laterofrontal cilia of ctenidia in
Neogaimardia
do not form hornlike structures. In all other aspects,
Neogaimardia
appears similar to
Gaimardia
. In fact, both taxa also have similar modes of life, attached to floating algae. Odhler (1924) described how
N. rostellata
brood ‘large eggs … in follicles which are formed as in
Pseudokellya
(cf. Pelseneer, 1903)’. Considering the very peculiar structure described by Pelseneer (1903), it seems clear that Odhner’s description refers to the follicular epithelium considered here as diagnostic for
Cyamioidea
.
Progaimardia
(type species: Modiolarca minutissima Iredale, 1908)
The general shell morphology of the type species closely resembles that of
Gaimardia
. In fact, this genus was originally proposed as a subgenus of
Gaimardia
, from which it was distinguished by the presence of a large, internal ligament and a strong hinge plate, bearing one large tooth in each valve and moderately long anterior and posterior teeth ( Ponder, 1971). No additional anatomical or reproductive information is available for this genus.
Eugaimardia
(type species:
Neogaimardia perplexa Cotton, 1931
)
Cotton (1931a) described a new species of gaimardiid, erecting
Neogaimardia
as a new genus for its placement, but had overlooked that this name was preoccupied by
Neogaimardia Odhler, 1924
. Cotton (1931b) proposed
Eugaimardia
as a replacement name for the former. Cotton (1931a) distinguished his new genus from
Gaimardia
by the absence of a concavity in the ventral margin and by its different ‘dentition of the hinge’, namely the presence of a ‘U-shaped tooth, and a small tooth between the arms of the U’ in the right valve. The morphology of the teeth he described does not appear as discrepant from the one we find in
G. trapesina
. Furthermore, in
G. trapesina
, the concavity of the ventral margin proves to be variable among specimens. Thus, from a morphological point of view, we find no evidence to consider
Eugaimardia
to be distinct from
Gaimardia
. Unfortunately, nothing is known about the anatomy or reproduction of
Eugaimardia perplexa
. Despite that,
Eugaimardia
was considered a valid genus by Huber (2010).
Costokidderia
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(type species:
Kidderia costata Odhler, 1924
View in CoL
)
The type species of this genus has a shell outline that closely resembles that of
Kidderia
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. Odhler (1924) described for that species the presence of strong radial sculpture and strong cardinals. The first of these characters led Finlay (1926) to propose the genus
Costokidderia
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, a name currently regarded as a subgenus of
Kidderia
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(e.g. Ponder, 1971).
Cyamionema
(type species:
Cyamium (Cyamionema) decoratum Melvill & Standen, 1914
)
Despite having originally been compared with
Cyamium
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, the general shell outline and morphology of the hinge teeth of
Cyamium decoratum
appear more similar to those of
K. minuta
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(type species of
Kidderia
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) than to those of
Cyamium antarcticum
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(type species of
Cyamium
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).
Cyamionema decoratum
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differs from
Cyamium antarcticum
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(and from other
Cyamiidae
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) by lacking an internal ligament, by having only one tooth in the left valve and by the presence of thin radial sculpture in the central part of the shell ( Melvill & Standen, 1914). The significance of these differences and the relationship of
Cyamionema
to
Kidderia
View in CoL
deserve further studies. Scarlato & Starobogatov (1979) proposed a new family,
Cyamionematidae
(in
Galeommatoidea
), based on this genus.
Cyamionema
appears listed as a subgenus of
Kidderia
View in CoL
by Huber (2010).
TAXA HERE EXCLUDED FROM
CYAMIOIDEA