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Further comments on the origin of oysters

Article  in  Palaeogeography Palaeoclimatology Palaeoecology · October 2006

DOI: 10.1016/j.palaeo.2006.03.007

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 Palaeogeography, Palaeoclimatology, Palaeoecology 240 (2006) 672 – 674
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Discussion

Further comments on the origin of oysters

Antonio G. Checa a , Antonio Pablo Jiménez-Jiménez a ,
Ana Márquez-Aliaga b,⁎, Hans Hagdorn c
a
Departamento de Estratigrafía y Paleontología, Facultad de Ciencias, Universidad de Granada, Avda. Fuentenueva s/n,
E-18071, Granada, Spain
b
Instituto Cavanilles de Biodiversidad y Biología Evolutiva and Departamento de Geología, Universitat de València,
Dr. Moliner 50, E-46100 Burjassot, València, Spain
c
Muschelkalkmuseum Ingelfingen, Schloss-Straβe 11, D-74653 Ingelfingen, Germany
Received 9 February 2006; received in revised form 22 February 2006; accepted 8 March 2006

Keywords: Oysters; Origin; Bivalves; Prospondylids

In his comment to our recent paper (Márquez-Aliaga of the Japanese and Chinese forms, which we attribute
et al. 2005), Hautmann (2006) raises two interesting to Prospondylidae gen. indet., to be considered the
questions: (a) the ambivalent attachment to the substrate closest to earliest oysters.
recognized in the species cristadifformis Schlotheim, While acknowledging Hautmann for his constructive
1820 and spondyloides Schlotheim, 1820, which we critique, we would like to additionally comment on
include into the Ostreoidae genus Umbrostrea, is in some of them while disagreeing with others:
conflict with the sinistral attachment usually recognized
as an autapomorphy of the group and (b) antimarginal (a) Recognition of autapomorphies permits the clear-
ribs are not valid as a character linking Prospondylus cut definition of clades. Our proposal to include
acinetus Newell and Boyd, 1970 and early oysters (our ambivalent forms in Ostreidae somehow alters the
proposal of derivation), because they appear in several present-day well-defined concept of oysters. We
unrelated families of bivalves. Moreover, Hautmann adopted our position by taking into consideration
(2005), finds additional difficulties in accepting our that it does not affect the accepted monophyly of
hypothesis of descent of oysters from the Prospondy- the clade (but see Malchus, 1990, regarding
lidae because, contrary to the latter group, (c) oysters, in Cenozoic Lophinae) and the lack of knowledge
general, do not develop a pallial line; (d) the resilifer of on the evolution of oysters through the Permian–
the right valve is in high relief and flanked by two Triassic transition. Future progress in this subject
depressions and the ligamentary area of the left valve will allow more formal aspects generic and
impresses such shape; (e) the microstructure of the inner familial assignment for these early oysters.
shell layer of prospondylids (crossed-lamellar) is (b) Antimarginal ribs were recognized in several
different from that of early oysters (probably nacreous); bivalve families, but we also established the
(f) too few data are known from the internal morphology clear morphogenetic difference between the anti-
marginal ribs of oysters, plicatulids and dimyids,
⁎ Corresponding author. which affect the whole shell thickness, and those
E-mail address: marquez@uv.es (A. Márquez-Aliaga). of other bivalves, in which ribs are restricted to the
0031-0182/$ - see front matter © 2006 Elsevier B.V. All rights reserved.
doi:10.1016/j.palaeo.2006.03.007
 Discussion 673

outer shell layers (Checa and Jiménez-Jiménez pallial line in some specimens of Lopha?
2003, p. 147). We determined that ribs of oysters, murakamii (which we instead assign to Prospon-
plicatulids and dimyids basically form by a dylidae and find to be intermediate between P.
process of folding of the mantle margin and that acinetus and oysters). Therefore the pallial
these groups are the only ones among present-day attachment might have already been lost in these
bivalves sharing that morphological, as well as Late Permian forms.
morphogenetic, trait. Differences in the micro- (d) In general this is a highly variable character in
structure and hinge-ligament structure allow us to oysters. In our collection of Cenozoic oysters, the
also differentiate two independent clades (Ostrei- resilifer of the right valve varies from elevated to
dae and Plicatulidae+Dimyidae). Recognition of flat to depressed within single species (Fig. 1A,
an oyster-like ribbing pattern in Prospondylus B). This change in shape also affects the lateral
acinetus and other indeterminate Late Permian elevations of the resilifer. The resilifer of the left
Prospondylidae link these groups with oysters valve is always depressed. Interestingly, good
(see additional data in Márquez-Aliaga et al., examples of right valves with depressed resilifers
2005). Without further explanation, Hautmann can be appreciated in the specimens of Umbros-
(2005) disregards the relevance of antimarginal trea emamii figured by Hautmann (2001, Plate 6,
ribs in bivalve evolution. Fig.13, Plate 8, Figs. 2,4,5,7). In view of this
(c) We thank Hautmann for attracting our attention to highly intraspecific variability, we find the
this point. Certainly the transition from P. acinetus alivincular-arcuate ligamentary area of Hautmann
to early oysters involved loss of the pallial (2004) hardly tenable as an autapomorphy of
attachment. This transition covered a wide time oysters. Additionally, the right resilifer of the
span and there are surely species involved, which recent gryphaeid genus Neopycnodonte is always
are yet to be discovered. Liberation from its low relief (Fig. 1C).
attachment to the shell margin enables the oyster (e) This aspect was discussed at length in our original
mantle to extend or contract to a high degree, paper. In short, the only reliable data on the
which might be useful for, e.g., the periodic microstructure of the Prospondylidae come from
secretion of the extensive lamellae usually found two species of the Triassic genus Newaagia
on both valves. As mentioned by Hautmann, P. (Carter, 1990; Hautmann, 2001). The statement
acinetus still shows a pallial line, but we want to that “prospondylids also differ in the crossed-
stress that it is very internal within the shell (see lamellar layer microstructure of their inner shell
Newell and Boyd, 1970, Fig. 20C). Internalization layer from Triassic ostreids, in which the
of the pallial line in P. acinetus (and in the aragonitic shell layer was probably nacreous”
Prospondylidae, in general) may represent a first Hautmann (2005) is based on the assumptions that
step towards the acquisition of a widely extensible (1) the microstructure of Newaagia represents the
mantle, enabling this and other species of the general condition in Prospondylidae (note that
Prospondylidae to secrete their extensive shell Carter, 1990, related Newaagia with pectinoi-
lamellae. Nakazawa and Newell (1968) recog- deans, rather than with the Pseudomonotidae) and
nized adductor muscle imprints, but no traces of a (2) the same applies to the Carnian specimen UNC

Fig. 1. Examples of depressed resilifers of the right valve in Neogene oysters and gryphaeids. (A) Ostrea lamellosa Brocchi, Lower Pliocene, El
Alquián, Almería, Spain. (B) Hyotissa hyotis (Linnaeus), Lower Messinian, La Mela, Almería, Spain. (C) Neopycnodonte navicularis (Brocchi),
Lower Pliocene, El Alquián, Almería, Spain. Specimens reposited in the Departamento de Estratigrafía y Paleontología, Universidad de Granada;
scale bars = 1 cm.
674 Discussion

13497b of Carter (1990) with regard to early References

oysters. More data are still needed to make this a
useful character in ostreid phylogeny. Carter, J.G., 1990. Evolutionary significance of shell microstructure in
(f) From the exterior of the valves we concluded that the Palaeotaxodonta, Pteriomorphia and Isofilibranchia (Bivalvia:
Mollusca). In: Carter, J.G. (Ed.), Skeletal Biomineralisation:
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described by Nakazawa and Newell (1968) as Reinhold, New York, pp. 135–296.
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classified as Enantiostreon by Xu (1976) were and Plicatuloidea (Bivalvia, Pteriomorphia) and its evolutionary
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postlarval stages and had a typical oyster-like rib tación en bivalvos. Fabricación y significado evolutivo de las
pattern . This alone certifies intermediate place- costillas antimarginales de Ostreoidea y Dimyoidea. Fundación
ment between P. acinetus and oysters. Conjunto Paleontológico de Teruel, Teruel.
Checa, A., Jiménez-Jiménez, A.P., Márquez-Aliaga, A., Hagdorn, H.,
Finally, Hautmann (2005) states that we prefer 2003. Revisión de Enantiostreon e implicaciones sobre el origen
de Plicatulidae y Dimyidae (Plicatuloidea, Pteriomorphia, Bival-
derivation of the Ostreidae, instead of the Plicatulidae, via). Libro Resúmenes XIX Jornadas Soc. Española Paleontol.
from the Prospondylidae. Obviously, the two possibil- Morella, pp. 51–52.
ities are not mutually exclusive. We do not understand Hautmann, M., 2001. Taxonomy and phylogeny of cementing Triassic
why he goes into the origin of the Plicatulidae, because bivalves (families Prospondylidae, Plicatulidae, Dimyidae and
this is briefly alluded to in the introduction of our paper Ostreidae). Palaeontology 44, 339–373.
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with reference to a previous report (Checa et al., 2003) ligaments and its implications for the timing of the “Mesozoic
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