THE SHELL M I C R O S T R U C T U R E ,

) MINERALOGY AND ISOTOPIC C O M P O S I T I O N

OF A M U S S I O P E C T E N B A R A N E N S I S
(PECTINIDAE, BIVALVIA) FROM
THE M I O C E N E OF SPAIN :
A VAI,UABLE P A L E O E N V I R O N M E N T A L TOOL
ISABELZAMARREI~O, JAIMEDE PORTA & ANTONIOV/LZQUEZ

ZAMARREI~O I., DE PORTA J. & V/~ZQUEZ A. 1996. The shell microstructure, mineralogy and isotopic composi-
tion ofAmussiopecten baranensis (Pectinidae, Bivalvia) from the Miocene of Spain : a valuable palaeoenvironmen-
tal tool. [Microstructure, minSralogie et composition isotopique de la coquille d'Amussiopecten baranensis
(Pectinidae, Bivalvia) du Miocene d'Espagne : un outil de valeur pour le pal~oenvironnement]. GEOBIOS, 29, 6 : 707-
724. Villeurbanne, le 31.12.1996.

Manuscrit d~pos~ le 16.11.1994 ; accept~ d~finitivement le 26.06.1995.

ABSTRACT -Amussiopecten baranensis (ALMERA• BOFILL,1896), of the Family Pectinidae, is widespread in Miocene
marine sediments from northeastern and southeastern Spain, ranging in age from Early Langhian to Middle
Messinian. The shell consists of an outer homogeneous and foliated layer composed of low-magnesium calcite, a
middle layer aragonitic, and an inner foliated layer of low-magnesium calcite. Both left and right valves have, in
general, the same microstructure and mineralogy except for an outermost thin polygonal layer in the earliest-for-
med dissoconch of the right valve. The homogeneous structure (dense and granular sublaminae) up to 50 ~m thick
overlies the sublayer of foliated calcite (regular, irregular and crossed foliated) that extends throughout the whole
shell, being thinnest in the dorsal margin and thickening toward the ventral margin. The regular foliated inner
layer extends from the beak, where it attains its maximum thickness, to near the adductor muscle scar. The midd-
le layer is the most complex in structure and distribution, being composed of three sublayers which from exterior to
interior are : 1 ) aragonitic simple crossed lamellar, 2) aragonitic prismatic (myostracal), and 3) aragonitic complex
crossed lamellar. The latter sublayer in the internal shell surface extends to the area interior to the pallial line.
Prominent features of the middle layer are the asymmetrical distribution of the prismatic structure (absent in the
anterior margin), and the thinning of the three sublayers both in radial and antero-posterior directions. The divari-
cate microsculpture extending on the umbonal area and auricles in both valves originates from the granular calci-
te of the homogeneous structure. 5180 values of the outer foliated layer from the ventral margin suggest that A.
baranensis segregates its calcite in isotopic equilibrium with ambient sea-water, and thus affords a valuable tool in
paleoenvironmental studies.

KEYWORDS : SHELL STRUCTURE, STABLE ISOTOPES, AMUSSIOPECTEN BARANENSIS, PALAEOENVIRON-

MENT, MIOCENE, SPAIN.

R]~SUM]~ - Amussiopecten baranensis (ALMERA& BOFILL,1896) de la famille des Pectinidae est tr~s r~pandue dans
les s~diments du Miocene marin du Nord-Est e.t Sud-Est de l'Espagne. La r~partition stratigraphique du taxon est
comprise entre le Langhien infSrieur et le Messinien moyen. La coquille est form~e par une couche externe de calci-
te homog~ne et feuillet~e, une couche moyenne d'aragonite prismatique et lamellaire crois~e et finalement une couche
interne de calcite feuillet6e. Les deux valves d'un m~me specimen montrent en g~n~ral les m~mes microstructure et
min~ralogie ~ l'exception de la jeune dissoconque de la valve droite qui pr~sente ~ la surface du test un agencement
polygonal des cristaux. La structure homog~ne (calcite prismatique et granulaire), atteignant 50 pm d'~paisseur et
qui surmonte la couche de calcite feuillet~e (r~guli~re, irr~guli~re et crois~e), est pr~sente partout dans la coquille.
La structure homog~ne s'amincit vers le bord dorsal tandis qu'elle atteint le maximum d'6paisseur vers le bord ven-
tral. La couche feuillet~e r~guli~re interne est pr~sente depuis le crochet ok elle atteint le maximum d'~paisseur, jus-
qu'~ la r~gion de l'empreinte du muscle adducteur. La couche moyenne est plus complexe rant du point de vue de la
structure que de sa distribution dans le test. De la face externe ~ la face interne du test elle est subdivisSe en trois
zones : 1 ) aragonite lamellaire-crois~e simple ; 2) aragonite prismatique (couche myostracale) ; et 3) aragonite lamel-
laire-croisSe complexe qui recouvre toute la face interne jusqu'~ la ligne pall~ale. On remarquera la distribution
asym6trique de l'aragonite prismatique de la couche moyenne (qui manque dans le bord ant~rieur) ainsi que l'amin-
cissement des trois zones tant dans le sens umbono-pall~al qu'ant~ro-post~rieur. La microsculpture divariqu~e, pr~-
sente dans la rSgion umbonale et sur les oreillettes des deux valves, est form~e par la calcite granulaire de la
 708
structure homog~ne. La calcite feuillet~e du bord ventral chez A. baranensis montre des valeurs du 5180indiquant
qu'elle a pr~cipit~ en ~quilibre isotopique avec le milieu marin et s'av~re donc ~tre un outil important pour les
~tudes pal~o~cologiques.
MOTS-CLES : MICROSTRUCTURES, COQUILLE, ISOTOPES STABLES, AMUSSIOPECTEN BARANENSIS,
PAL]~OENVIRONNEMENT, MIOCt~NE, ESPAGNE.
INTRODUCTION
During a geochemical s t u d y carried out on
Miocene deposits from northeastern and sou-
theastern Spain, foraminifera were proven to be
unsuitable for isotopic and trace element ana-
lyses. Even after breaking the foraminifera tests
it was not possible to clean the chambers of
infilling sediment. F u r t h e r m o r e , taking into
account that specimens of Amussiopecten bara-
nensis (AL~ERA & BOFILL, 1896) were abundant in
shallow-water marine sequences at several locali-
ties covering an extended stratigraphical range
(Langhian to Upper Messinian), they were used
for geochemical analyses instead of foraminifera.
Based on observations from scanning electron
microscopy and X-Ray diffraction analysis, it was
noticed that A. baranensis conserved its original
mineralogy and microstructure. According to our
results Pectinidae shells m a y be a potentially
valuable tool in providing an indicator of the
variations in 51s0 and 513C.
A general description of the shell microstructure
of the Pectinidae has been given among others by
Bcggild (1930), Taylor et al. (1969), Waller
(1972b), Bongrain & Fatton (1982), Schein-Fatton
(1988), and Schein (1989). A noteworthy recent
general review has been made by Carter (1990, p.
260-269 and 379-387) and for deep-sea Pectinidae
by Schein (1989). The microstructure features in
the early growth stages of recent and fossil
Pectinidae have also been documented by Fatton
& Bongrain (1980). Amussiopecten is a member of
the Pectinidae not only according to its shell mor-
phology, b u t also because of its general micro-
structural features. The shell microstructure of A.
baranensis, however has been described only in
broad lines by Freneix et al. (1982).
This paper is mainly concerned with the detailed
description and origin of the microstructure and
microsculpture of A. baranensis and may also
contribute to the knowledge of the affinities bet-
ween Amussiogecten and Flabelligecten genera as
discussed by Freneix et al. (1982).
Finally, mineralogical determinations in conjunc-
tion with microstructural observations have pro-
vided the basis for establishing the distribution
and mineralogical variability of the shell in A.
baranensis.
MATERIAL A N D M E T H O D S
Specimens of A baranensis were collected from
twelve sections at various localities in shallow-
water Miocene basins from northeastern Spain
and from two sections in southeastern Spain for
shell microstructure and mineralogical analyses
(Fig. 1). The specimens were sampled at regular
intervals in marl and sandy marl beds. They are
abundant in most localities b u t it was difficult to
obtain complete specimens because of shell frag-
mentation due to compaction. Most of the shells
p r e s e r v e d in soft marls are d i s a r t i c u l a t e d
although a few have the valves still associated.
Only the shells found in sandy marls exhibit dia-
genetic alteration.
Radial and transverse cross-sections of both left
and right valves were oriented from umbo to ven-
tral and in antero-posterior margin directions res-
pectively. Several oblique sections have also been
studied for additional information concerning the
extension of certain shell sublayers.
The specimens were embedded in epoxy resin
prior to sectioning, polished and etched for 5
seconds in 5% HC1 and then coated with gold. A
few samples were also etched for 30 minutes with
EDTA. Observations of shell microstructure were
undertaken with a Hitachi 239 scanning electron
microscope (SEM) of polished and fractured verti-
cal sections. SEM observations of the external
shell surface permitted their microsculpture to be
described.
Shell mineralogy determined by X-ray diffraction
of powdered specimens supplemented with Feigl
staining and R a m a n spectra of complete speci-
mens as well as polished cross-sections allowed to
discriminate b e t w e e n l o w - m a g n e s i u m calcite
(LMC) and aragonite shell sublayers.
Carbon and oxygen isotopic analyses were perfor-
med on A. baranensis samples removed from the
ventral margin which consists of the outer homo-
geneous and foliated low-magnesium calcite layer.
The samples were ultrasonically cleaned, dried at
60oC and ground. Each powdered sample of 20 mg
in weight was vacuum roasted at 400 ° C for 6
hours in a furnace of t h e Stable Isotope
L a b o r a t o r y of t h e E a r t h Sciences I n s t i t u t e
(C.S.I.C.), Barcelona, in order to remove the orga-
nic components of the shell matrix. C02 for ana-
lyses was extracted by reaction in 100% phospho
 709

FIGURE 1 - M a p s h o w i n g location
of s a m p l e sites of Amussiogecten
baranensis i n n o r t h e a s t e r n a n d
s o u t h e a s t e r n Spain. 1 : C a n Rosell
; 2 : Els Monjos ; 3 : C a n J e i r a ; 4 :
E r m i t a de Ber~ ; 5 : C a s e s Virgili ;
6 : L a N o u de Gai~; 7 : Els M a s o s
de V e s p e l l a ; 8:Vespella ; 9 :
C a t l l a r ; 10 : A l t a f u l l a ; 11 :
Capr~s-E1 Tale ; 12: C a m p o s del
Rio. Carte de localisation des
prdl~vements d'Amussiopecten
b a r a n e n s i s dans le Nord-Est et le
Sud-Est de l'Espagne.

ric acid at 25 ° C and analysed in a VG Sira mass

spectrometer in the Department of Analytical
Chemistry at the University of Barcelona. The
analytical precision of the carbonate standard
(Carrara marble) run was +0.01 per mil for 5~3C
and 0.05 per mil for 5~80.All isotope data are refer-
red to PDB in the international a-notation.

LIST OF LOCALITIES AND

CHRONOSTRATIGRAPHIC
A B
SOURCES
Twelve stratigraphic successions were sampled
for microstructural and isotopic analyses of A.
b a r a n e n s i s (Fig. 1 ). Collection sites and stratigra-
phical positions of samples will be referred to by
letters and numbers in Figures and Tables.
Additional stratigraphical and lithological des-
criptions can be found in the references quoted for
each location.
1. Can Rosell section (CR, 13 samples). Langhian
(N8 Zone). Agusti et al. (1990) ; V~zquez et al. C • D
(1990, 1991).
2. Els Monjos section (MJ, 11 samples). Langhian FIGURE 2 - A m u s s i o p e c t e n baranensis. E x t e r n a l (A) a n d i n t e r -
n a l (B) v i e w s of a r i g h t valve. E x t e r n a l (C) a n d i n t e r n a l (D)
(N8 Zone). v i e w s of a left valve. Catllar. All s p e c i m e n s x 1. Vue externe (A)
3. Can Jeira section (CJ, 14 samples). Langhian et vue interne (B) d'une valve droite. Vue externe (C) et rue
(N8 Zone). V~zquez et al. (1991). interne (D) d'une valve gauche.
4. Ermita de Beret section (BA, 5 samples).
Serravallian (G. mayeri Zone). Porta et al. (1977, 7. Els Masos de Vespella section (MV, 12 samples).
1985). Langhian-Serravallian (G. fohsi peripheroronda-
5. Cases Virgili section (CV, 9 samples). Langhian G. mayeri Zone). Civis et al. (1985).
(N8 Zone). Porta et al. (1985) ; Civis et al. (1985). 8. Vespella section (TG, 8 samples). Langhian-
6. La Nou de Gai~ section (LN, 6 samples). Serravallian (G. fohsi peripheroronda-G, mayeri
Langhian (N9 Zone). Zone). Civis et al. (1985).
 710

9. Catllar section (CC, 4 samples). Serravallian
(G. mayeri Zone). Civis et al. (1985).
10. Altafulla section (AT, 1 sample). Tortonian.
Porta (1972) ; Porta & Civis (1996).
11. Capr6s-EI Tale section, Fortuna Basin (CT, 55
samples). Middle Messinian (G. elongatus Zone).
Sierro et al. (1992) ; Vfizquez et al. (1992).
12. Campos del Rio section, Mula Basin (ML, 162
samples). Middle Messinian (G. elongatus Zone).
All the specimens are housed at the University of
Barcelona (Departament de Geologia Din~mica,
Geofisica i Paleontologia).
SHELL MORPHOLOGY
beds. The right valve with prominent spines in
the hinge margin is more convex than the left
valve. The hinge shows three short cardinal crura
in the right valve and two in the left.
A general revision and detailed description of the
species has recently been made by Freneix et al.
(1982). Several specimens from Can Rosell (Lower
Langhian, N 8 Zone) and Punta de la Mora
(Tortonian) localities attain heights of 56.4 mm
and 63.7 mm respectively. Therefore, they are
higher than the specimens quoted by Freneix et
al. (1982). The greatest dimension (71.5 mm),
however, corresponds to a Middle Messinian spe-
cimen collected at the Campos del Rio section.

Amussiopecten baranensis (AL~ERA & BOFILL,

1896) of the Family Pectinidae is widespread in
Miocene marine sediments in Spain. The species
is characterized by disc-shaped shells with smoo-
th to nearly obsolescent radial external costae
with correspondingly stronger internal radial
ribs, predominantly occurring in pairs (Fig. 2).
Each pair of internal ribs corresponds to an
interval between two external costae. In most
specimens a Camptonectes microsculpture can be
observed in both valves.
In general the specimens are characterized by
thin shells in marly deposits whilst they are thic-
ker in sandy marl and fine-grained sandstone
SHELL M I C R O S T R U C T U R E
Bivalve shell microstructure is rather complex
and several layers can be distinguished. These
layers differ from one another in orientation of
crystals and sometimes in mineralogy.
Since the outstanding paper of Bcggild (1930) des-
cribing the shell microstructure and mineralogy
of shell molluscs, several classifications of mollus-
can shell microstructures have been established,
based largely on B0ggild's terminology. The termi-
nology for microstructures, however, has not rea-
ched a general agreement among specialists (for a

Lv

FIGURE 3 - Composite reconstruction of the distribution of shell layers in Amussiopecten baranensis s h o w n in longitudinal a n d
t r a n s v e r s e sections b a s e d on s c a n n i n g electron microscope observations. T r e n d of first-order lamellae s c h e m a t i c a l l y s h o w n in t h e
outer crossed foliated calcite layer. AM, Adductor muscle scar ; PI, pallial line ; LV, i n t e r n a l view of t h e left valve w i t h location of
t h e cross-section. 1 : H o m o g e n e o u s calcite ; 2 : Crossed foliated calcite ; 3 : Regular foliated calcite ; 4 : I r r e g u l a r foliated calcite ;
5 : C r o s s e d l a m e l l a r aragonite ; 6 : P r i s m a t i c aragonite. Reconstitution longitudinale et transversale des couches de la coquille chez
A. b a r a n e n s i s , basdes sur des observations au MEB. L'agencement des lamellae de premier ordre est indiqud par des traits dans la
couche calcitique feuilletde externe. AM, empreinte du muscle adducteur ; Pl, ligne palldale ; LV, face interne de la valve gauche avec
localisation des coupes. 1 : structure homog~ne calcitique ; 2 : calcite feuilletge-croisd~ ; 3 : calcite feuilletde-rdguli~re ; 4 : calcite
feuilletde-irrgguli~re ; 5 : aragonite lamellaire-croisge ; 6 : aragonite prismatique.
 711

FIGURE 4 - SEM photomicrographs of the homogeneous structure. Scale bars : 25 pm in 1 and 3 ; 5 pm in 2 and 2.5 pm in 4.1, trans-
verse vertical section of the left valve near the crura, showing the calcitic porcellaneous sublamina (a) and granular inner sublami-
na (b) of the homogeneous structure, underlain by the calcitic crossed foliated sublayer (c) of the outer shell layer. Can Rosell (CR-
7/3). 2, closer view of the outermost porcellaneous sublamina in A. Note the faint, vertical structural trend exhibited by this subla-
mina. 3, transverse vertical fracture of the right valve showing the porcellaneous (a) and granular (b) sublaminae of the homogeneous
structure grading inward to the foliated calcite (c). Can Rosell (CR-B). 4, external view of the left valve showing the granular inner
calcitic sublamina of the homogeneous structure. Note the irregular morphology and variable size of crystallites. Can Rosell (CR-53).
Vues au M E B de la structure homog~ne. 1, coupe transversale de la valve gauche pros de la crura montrant les deux zones de la struc-
ture homog~ne : a, calcite et b, calcite granulaire ; c, calcite feuilletge de la couche externe. 2, dgtail de la zone de calcite de la photo 1.
A noter l'agencement vertical. 3, cassure transversale de la valve droite montrant le passage de la calcite homog~ne (a et b) h la calci-
te feuilletge (c). 4, face externe de la valve gauche montrant la couche de calcite granulaire intacte de la structure hornog~ne. On obser-
ve une morphologie irrdguli~re ainsi qu'une variation dans la taille des cristaux.

r e v i e w s e e T a y l o r e t al. 1969 ; K o b a y a s h i 1971 ; T h e dissoconch ofA. baranensis at low m a g n i f i c a -

C a r t e r & C l a r k I I 1985 ; a n d C a r t e r 1990). T h e t i o n a p p e a r s to b e f o r m e d b y t h r e e m a i n c a l c a -
standard descriptive terminology in the Glossary r e o u s l a y e r s , h e r e r e f e r r e d to a s o u t e r , m i d d l e a n d
of S k e l e t a l B i o r n i n e r a l i z a t i o n ( C a r t e r 1990) h a s inner layers. SEM observation shows that the
b e e n u s e d i n this study, m i d d l e l a y e r is t h e m o s t c o m p l e x i n s t r u c t u r e ,
 712

being formed by up to three sublayers. The follo-
wing major structural types have been recognized
in A. b a r a n e n s i s : homogeneous, foliated, prisma-
tic, and crossed lamellar. The outer layer is com-
posed of the homogeneous and foliated structure
and is always calcitic. The middle layer is formed
together with the aragonitic crossed-lamellar and
prismatic structures whilst the inner layer is
made up of calcitic foliated structure. It is worth
mentioning t h a t the whole shell microstructure of
A. b a r a n e n s i s in all specimens studied except at
one locality (Altafulla section) is an original fea-
ture and not diagenetic as indicated by trace ele-
m e n t analyses of the shell (V~zquez et al. 1991 ).
The description of A. b a r a n e n s i s microstructures
is based on SEM observations of radial, transver-
se and oblique sections although the oblique sec-
tions are the most difficult to interpret due to the
t h i n n i n g of the middle sublayer toward the umbo-
nal and ventral margins. The shell microstructu-
re distribution is summarized in Figure 3.
HOMOGENEOUS STRIJCTURE
This calcitic structure constitutes the outermost
and extremely thin layer covering the whole shell
and overlying the foliated structure. Its thickness
varies from 4 to 7.5 pm in the u m b o n a i area and
from 20 to 50 pm in the ventral margin, although
variations could exist between specimens. SEM
observations in numerous specimens from both
valves in either radial or transverse sections show
two distinct portions ; an outer, even dense and
porcellaneous sublamina exhibiting a faint, verti-
cal structural trend (Fig. 4, la-2), ranging from
1.75 pm to 18 pm in thickness ; an inner granular-
like sublamina with more regular a r r a n g e m e n t of
calcite crystallites but without l a t h formation
(Fig. 4, lb). This inner sublamina ranges in thick-
ness between 2.5 and 32 pm. It is made up of
minute, irregular crystallites up to 3 pm in dia-
meter although their size is quite variable (Fig 4,
4). Under crossed polarized light the aggregations
of randomly-distributed calcite crystallites inside
each sublamina exhibit uniform extinction, but
differ in angle of extinction between the outer
sublamina with respect to the inner one.
The distinction of two sublaminae within the
homogen.eous structure has been also pointed out
by Schein (1989), Carter (1990) and Barrera et al.
(1990). Similar arrangements of calcite crystals
have been described by Clark II (1974) during
the early stages of calcification u n d e r the perios-
tracum in shells of the living Pecten diegensis,
suggesting the importance of substrate stability
for orderly shell growth of the calcite l a t h crystals
(foliated outer sublayer) with a m a r k e d layering.
FOLIATED STRUCTURE
This structure is built up of tiny elongate LMC
calcite laths (third-order lamella) arranged in
sheets (second-order lamella) grouped in larger,
lenticular folia (first-order lamella).
According to the architectural p a t t e r n s of first-
order lamella the following microstructural types
have been recognized :

PLATE 90

Fig. 1 - Slightly oblique, radial section through the right valve, showing the outermost extremely thin sublayer of
homogeneous structure (a), underlain by a crossed foliated sublayer (b), grading inward from regular foliated (c)
to irregular foliated structure (arrow). The ventral shell margin is toward the right. S, sediment infilling. Can
Rosell (CR-12). Coupe radiale de la valve droite montrant le passage de la structure homog~ne (a) & la feuilletde-
croisde (b), feuilletge-rdguli~re (c) et feuilletde-irrgguli~re (fl~che). S, remplissage de sddiment.
Fig. 2 - Transverse, vertical section through the outer shell layer, showing low-angle crossed foliated calcite (com-
pare with fig. 1 ). The external surface of the shell is at the top of the figure. Can Rosell (CR-7). Coupe transver-
sale. La calcite feuilletde-croisde prdsente des angles faibles en comparaison avec ceux de la fig. 1. La surface exter-
ne de la coquille est situde dans la partie supdrieure de la photo.
Fig. 3 - Oblique radial, vertical section through the outer foliated calcite layer of the left valve, showing the irregu-
lar foliated structure (a), grading inward to the regular foliated calcite (b). The ventral shell margin is toward the
left. Els Monjos (MJ-15). Coupe radiale de la valve gauche. Passage de la calcite feuilletde-irrdguli~re (a) & la cal-
cite feuilletde-rdguli~re (b). Le bord ventral de la coquille est situd h gauche.
Fig. 4 - Close up view of the internal shell surface near the ventral margin, showing the low-magnesium calcite
laths of the foliated structure. The ventral shell margin is toward the lower left-hand corner. Can Rosell (CR-12).
Ddtail de la surface interne de la coquille pros du bord ventral montrant les lattes de calcite de la structure
feuilletde. Le bord ventral de la coquille se situe en bas et & gauche de la photo.
Fig. 5 - The same section as in fig. 1, showing the concentric arrangement of the regular foliated structure of the
inner shell layer in the beak. M~me coupe que fig. I montrant la disposition concentrique de la structure feuilletde-
rdguli~re de la couche interne pros du crochet.
SEM photomicrographs of the foliated structure. Scale bars : 250 pm in fig. 1 and 5 ; 25 pm in 2 and 3 ; 5 pm in fig. 4.
Geobios P1. 90
n ° 29, fasc. 6 I. Z a m a r r e f l o , J. d e P o r t a & A. V ~ z q u e z
 714

1. Crossed foliated s t r u c t u r e , consisting in r a d i a l a large portion of t h e depositional surface" (P1.90,

section of a l t e r n a t i n g , lensatic to b r a n c h i n g first-
o r d e r l a m e l l a e clearly identifiable u n d e r SEM by
t h e i r v a r i a t i o n in colour w i t h a l t e r n a t i o n of g r e y
a n d w h i t e folia. T h e g r e y folia are in general thin-
n e s t a n d develop slight or g r e a t c u r v a t u r e s w i t h
t h e c o n v e x i t y facing t o w a r d t h e v e n t r a l m a r g i n
(P1.90, fig. lb). T h e calcite crystals in all g r e y folia
display t h e s a m e orientation, different from those
of t h e light folia w h i c h in t u r n p p r e s e n t a u n i q u e
o r i e n t a t i o n of calcite l a t h s (P1. 90, fig. 2).
2. R e g u l a r foliated s t r u c t u r e , defined b y C a r t e r &
C l a r k II (1985) and t h e Glossary of Skeletal
B i o m i n e r a l i z a t i o n (in C a r t e r 1990, p. 656) as "a
l a m i n a r s t r u c t u r e in w h i c h parallel calcitic l a t h s
or blades are a r r a n g e d in sheets dipping at the
s a m e angle a n d in t h e s a m e g e n e r a l direction over
fig. lc).
3) I r r e g u l a r foliated structure. In several areas
w i t h i n the shell highly i r r e g u l a r orientations of
folia occur displaying a dendritic a r r a n g e m e n t (P1.
90, fig. 1, arrow). This s t r u c t u r e is similar to the
common foliated s t r u c t u r e previously described
except for the i r r e g u l a r shape and s m a l l e r size of
the first-order lamella (P1. 90, fig. 3a). T h e r e is a
gradational t r a n s i t i o n from crossed foliated to irre-
gular and to r e g u l a r foliated structures.
T h e foliated m i c r o s t r u c t u r e shows t h e following
distribution w i t h i n t h e shell : 1) a n o u t e r foliated
l a y e r e x t e n d i n g t h r o u g h o u t t h e whole shell beco-
m i n g t h i n n e r in its dorsal m a r g i n a n d t h i c k e n i n g
t o w a r d t h e v e n t r a l one ; a n d 2) an i n n e r foliated

PLATE 91

Fig. 1 - Radial, slightly oblique and bevelled section through the posterior margin of the left valve near the beak,
showing the thin outer layer of regular foliated calcite (a), and the three sublayers of the middle shell layer : b,
aragonitic crossed lamellar ; c, irregular simple prismatic aragonite ; and d, aragonitic complex crossed lamellar
draping the regular foliated calcite (e) of the inner shell layer. The ventral shell margin is toward the right. Can
Rosell (CR-7/1). Coupe radiale, l~g~rement en biseau de la valve gauche, a, calcite feuilletde-r~guli~re ; b, arago-
nite lamellaire-croisde simple ; c, aragonite prismatique ; d, aragonite lamellaire-croisde complexe ; e, calcite
feuilletde-rdguli~re. Le bord ventral de la coquille est orientd vers la droite.
Fig. 2 - Transverse, vertical section through the adductor muscle scar area showing the contact between the arago-
nitic prismatic (below), and the aragonitic complex crossed lamellar structure (above). Note the third-order ara-
gonitic lamellae sloping in two directions inside each first-order lamella (above). Can Rosell (CR-7). Coupe trans-
versale & travers l'impression du muscle adducteur montrant la relation entre l'aragonite prismatique (en bas) et
la structure lamellaire-croisde aragonitique (en haut).
Fig. 3 - Radial, vertical section through the anterior margin of the left valve nearthe beak, showing the aragonitic
crossed lamellar and prismatic aragonite (arrow) of the middle shell layer, overlain by the regular foliated calci-
te of the outer shell layer (a). Note that the outer and inner crossed lamellar sublayers are in contact when the
prismatic sublayer disappears, Note also the branching first-order lamellae of the outer crossed lamellar struc-
ture (b) and the larger and more irregularly shaped first-order lamellae of the inner sublayer (c). Els Monjos (MJ-
10). Coupe radiale & travers le bord antdrieur de la valve gauche pros du crochet. La fl~che signale l'aragonite pris-
matique de la couche moyenne, a, calcite feuilletde-rdguli~re de la couche externe. A noter que les deux zones de la
structure lamellaire-croisde sont en contact lh oh dispara$t l'aragonite prismatique. Les lamellae de premier ordre
de la structure lamellaire-crois~e de la zone supdrieure sont ramifides (b) tandis que celles de la zone infdrieure sont
plus irrdguli~res et larges.
Fig. 4 - Transverse, vertical fracture, through the middle shell layer in the umbonal area of the right valve, displaying
the aragonitic prismatic structure thinning out and iuterfingering within the aragonitic simple crossed lamellar
structure (above) and complex crossed lamellar structure (below). Anterior shell margin toward the right. Internal
shell surface at the bottom (arrow). Can Rosell (CR-B). Cassure transversale dans la zone umbonale de la valve droi-
te montrant la structure aragonitique prismatique qui s'amincit et s'intercale entre la lamellaire-croisde simple (en
haut) et la lamellaire-croisde complexe (en bas). La fl~che indique la surface interne du test.
Fig. 5 - Transverse, vertical fracture through the aragonitic crossed lamellar structure of the left valve near the
beak, showing the characteristic cross pattern of the second-order lamellae. The furrow of calcitic foliated struc-
ture indicative of the appearance of the internal rib is observable (above). Els Monjos (MJ-8B). Cassure trans-
versale de la valve gauche pros du crochet montrant la disposition typique croisde des lamellae de second ordre. La
calcite feuilletde en haut correspond & la c6te interne.
Fig. 6 -Radial, slightly oblique and bevelled section through the posterior margin of the left valve near the beak. The
aragonitic crossed lamellar inner sublayer of the middle shell layer, is underlain by the regular foliated calcitic
structure of the inner shell layer. The sharp contact with the aragonitic prismatic structure can be seen (above).
The ventral shell margin is toward the right. Can Rosell (CR-7/1). Coupe radiale lgg~rement en biseau de la valve
gauche pros du crochet. Le contact entre l'aragonite prismatique (en haut) et l'aragonite lamellaire-croisde sous-
jacente est tr~s net.
 P1. 91
Geobios
n ° 29, f a s c . 6 I. Z a m a r r e S o , J. d e P o r t a & A. V ~ z q u e z
 716

thickness n e a r t h e b e a k (Fig. 2, 3). Only in t h e

FIGURE 5 - Radial, vertical section through the shell anterior
of the left valve near the beak, displaying the gradational
contact between the regular foliated calcite of the outer shell
layer (a) and the regular foliated inner shell layer (c). The
middle shell layer is shown in b. Els Monjos (MJ-10). Scale
bar : 501]m. Coupe radiale & travers la valve gauche pr@s du
crochet montrant la continuitd entre la calcite feuilletde-rdgu-
li~re de la couche externe (a) et celle de la couche interne (c) ;
b, couche moyenne.
l a y e r e x t e n d i n g from t h e b e a k to t h e p r o x i m i t y of
t h e a d d u c t o r muscle scar. It a t t a i n s the m a x i m u m
u m b o n a l a r e a are b o t h l a y e r s in c o n t a c t (Fig. 5).
The outer foliated layer in either radial or transver-
se sections underlies the homogeneous s t r u c t u r e
and exhibits the following distribution. The regular
foliated structure extends from the umbo to the ven-
tral margins and is overlain by the crossed foliated
structure which appears n e a r the umbonal area and
terminate in the ventral margin w h e r e it attains its
m a x i m u m thickness. The irregular foliated structu-
re appears only locally and w h e n it occurs the follo-
wing distribution can be observed in vertical radial
sections. The crossed foliated s t r u c t u r e is gradually
succeeded inwards by the i r r e g u l a r foliated structu-
re t h a t also grades laterally to the regular foliated
structure (P1.90, fig. 1). The irregular foliated struc-
ture contributes to the development of the internal
ribs (Fig. 6). The same distribution is observed in
transverse sections, the only difference being in the
crossed foliated s t r u c t u r e w h e r e the first-order
lamellae display lower angles a m o n g adjacent
lamella (P1.90, fig. 2).
T h e i n n e r foliated l a y e r is p r e d o m i n a n t l y compo-
sed of r e g u l a r foliated s t r u c t u r e . T h e f i r s t - o r d e r
l a m e l l a b e n d close to t h e u m b o n a l region giving
w a y to a concentric, a r r a n g e m e n t (PI. 90, fig. 5).
According to this distribution the a r r a n g e m e n t of
calcite laths in the foliated structure m a y only be
seen in the internal surface of the shell, outside the
pallial line where the outer or inner shell layers are
exposed (P1.90, fig. 4). The calcite laths always inter-
sect the internal valve surface at very low angles.
PRISMATIC STRUCTURE (MYOSTRACAL
LAYER)
In vertical, radial and t r a n s v e r s e sections this
s t r u c t u r e is composed of aragonite p r i s m s showing
a high length/width ratio. The prisms are simple,
have s t r a i g h t or i r r e g u l a r b o u n d a r i e s a n d are elon-

PLATE 92

Fig.1 - External view of the right valve showing the smooth prodissoconch and earliest dissoconch. Els Monjos (MJ-
6/3). Surface externe de la valve droite montrant la prodissoconque et la dissoconque lisses.
Fig. 2 - Closer view of the prodissoconch in fig 1. The arrow indicates the ribbon at the boundary between the pro :
and dissoconch. Ddtail de la prodissoconque de la rue prdc~dente. La fl~che indique le bourrelet sdparant la pro-
dissoconque de la dissoconque.
Fig. 3 - Irregular polygonal pattern occurring in the earliest dissoconch of the specimen in fig. 1. Agencement poly-
gonal irrggulier des prismes dans la jeune dissoconque. M@rne spgcirnen que fig. 1.
Fig. 4 -A detail of fig 3. Ddtail du m~rne spdcimen que fig. 3.
Fig. 5-7 - Non-polygonal pattern on the prodissoconch of the right (fig. 5) and left valve (fig. 6) and the earliest dis-
soconch of the left valve (fig. 7). Els Monjos (MJ-6/6). La microstructure concentrique & la surface de la prodisso-
conque de la valve droite (fig. 5), de la valve gauche (fig. 6) et de la jeune dissoconque de la valve gauche (fig. 7),
ne prdsente pas d'agencernent polygonal des cristaux.
SEM photomicrographs of the external surface. Scale bars : 250 ~m in fig. 1 ; 50 ~m in fig. 2 ; 10 ~m in fig. 3 ; 2.5
gm in fig. 4 and fig. 7 ; 1 ~m in fig. 5 and 6. Vues au M E B de la surface externe.
 P1. 92
Geobios
n ° 29, f a s c . 6 I. Z a m a r r e ~ o , J. d e P o r t a & A. V f i z q u e z
 718

FIGURE 6 - Transverse, vertical section near the ventral shell

margin showing the outermost thin sublayer of homogeneous
structure (a) and the distribution of the foliated calcite in the
internal radial rib. The crossed foliated calcite (b) grades
inward to irregular foliated (c), and then to regular foliated
structure (d). Capr6s-EI Tale (CT-7/8). Scale bar : 250 pro.
Coupe transversale pros du bord ventral de la coquille mon-
trant la structure homog~ne (a) ainsi que la distribution de la
calcite feuillet~e dans la cSte interne, b, feuilletge-croisge ; c,
feuilletge-irrggulikre ; d, feuilletge-rdguli~re interne.

The d i a m e t e r of p r i s m s r a n g e s from one to three

g a t e d in the radial direction w i t h their l e n g t h axes
p e r p e n d i c u l a r to the shell surface (P1. 91, fig. 1-2).
microns with the average being 2.3 p m . T h e longest
crystal l e n g t h m e a s u r e d was 200 1Jm.
T h e i r r e g u l a r s i m p l e p r i s m a t i c s t r u c t u r e pre-
s e n t s a w e d g e - s h a p e d d i s t r i b u t i o n a n d only in
t h e a d d u c t o r m u s c l e s c a r a r e a w h e r e a t t a i n s its
m a x i m u m t h i c k n e s s c a n be o b s e r v e d (Fig. 3).
Hence, t h e m o s t p r o m i n e n t f e a t u r e is t h e i r a s y m -
m e t r i c a l d i s t r i b u t i o n in b o t h valves. I n all direc-
tions o u t s i d e of t h e m u s c l e s c a r a r e a t h e s t r u c t u -
re t h i n s out, i n t e r f i n g e r i n g w i t h t h e c r o s s e d
l a m e l l a r s t r u c t u r e a n d s h o w i n g one (P1.91, fig. 3)
or m o r e d i g i t a t i o n s (P1. 91, fig. 4).
T h e details a n d d i s t r i b u t i o n of this s t r u c t u r e as
well as t h e i r r e l a t i o n s h i p s w i t h t h e crossed lamel-
lar s t r u c t u r e are given f u r t h e r on.

PLATE 93

Fig. 1 - External view of the left valve, showing the smooth prodissoconch and earliest dissoconch, and the external
costae (arrow). Note the divaricate Camptonectes striae, extending on the dissoconch and the two auricles. Can
Rosell (CR-12B). Surface externe de la valve gauche montrant la surface lisse de la prodissoconque et de la jeune
dissoconque ainsi que les c6tes externes. On observe une microsculpture de type Camptonectes.
Fig. 2 - A closer view of the shell as in fig. 1 of the contact between the dissoconch and the posterior auricle (upper
right), displaying the external radial costae (arrow) and the divaricate microsculpture formed by the outermost
porcellaneous sublamina of the homogeneous structure. Ddtail de la microsculpture Camptonectes du m~me spg-
cimen que fig. 1. La flbche indique les c6tes externes.
Fig. 3 - Oblique, radial, vertical section of the left valve through an external radial costae (thick arrow), portraying
the lensatic first-order lamellae (thin arrows) per regular foliated layer. The aragonitic crossed lamellar struc-
ture a base of the figure. Els Monjos (MJ-15). Coupe radiale de la valve gauche montrant une c6te radiale externe
(fl~che dpaisse) et l'arrangement lenticulaire des lamelles de premier ordre (flbche mince) dans la couche de calci-
te feuilletde-rdguli~re. En bas la couche aragonitique lamellaire-croisde.
Fig. 4 - Closer view of two of the lensatic first-order lamellae in fig. 3, showing the different orientation of calcite
laths between adjacent lamellae, and the unique and parallel disposition of laths inside each lensatic first-order
lamella (arrows). Dgtail de la vue prdcgdente montrant les diffdrentes orientations des lattes (flbches).
Fig. 5 - External view of the left valve, in the umbonal area, of an abraded specimen, showing the three sublayers
of the outer shell layer : porcellaneous (a) and granular calcite (b) of the homogeneous structure, underlain by the
regular foliated calcite (c). Note the ridges and furrows in the granular sublayern, originating the divaricate
microsculpture (thin arrow). An external radial costa is shown between thick arrows, cl, aragonitic crossed lamel-
lar structure. Can Rosell (CR-53). Face externe de la rdgion umbonale de la valve gauche chez un spdcimen drodd.
On observe la calcite prismatique (a) et granulaire (b) de la structure homog~ne et la calcite feuilletde-rdguli~re (c)
sous-jacente. La flkche mince indique les stries divariqudes ; une c6te radiale externe est observable entre les deux
grosses fl~ches, cl, aragonite lamellaire-croisde.
Fig, 6 - Higher magnification of the concentric growth scales in fig. 5, originated by the porcellaneous sublamina of
the homogeneous structure. Ddtail de la rue prdcddente.
SEM photomicrographs of the microsculpture. Scale bars : 0.5 mm in fig. 1 ; 100 pm in fig. 2 and 5 ; 25 pm in fig. 3;
10 pm in fig. 4 and 6. Vues au M E B de la microsculpture.
Geobios P1. 93
n ° 29, fasc. 6 I. Z a m a r r e ~ o , J. d e P o r t a & A. V ~ z q u e z
 720

CROSSED LAMELLAR STRUCTURE

It consists of elongate, lensatic to branching first-
order lamellae built up of parallel aragonite crys-
tal aggregations (P1.91, fig. 1). The length axes of
second-order lamellae m a y alternate in two oppo-
site directions in adjacent first-order lamellae
showing a characteristic cross pattern of simple
crossed lamellar 2 structure (P1. 91, fig. 4-5) or
m a y exhibit three or more orientations (complex
crossed lamellar) as in P1. 91, fig. 6. According to
several authors this crystal arrangement is an
obvious adaptation for adding strength to the
shell. The crossed lamellar structure is always
aragonitic. The crossed lamellar structure deve-
lops into two sublayers separated by the aragoni-
tic prismatic structure. The first-order lamellae
are long and regularly shaped in the upper
sublayer, draping the prismatic structure. Bran-
ching is a striking feature of the first-order lamel-
lae in the upper portion of this sublayer (P1. 91,
fig. 1). The inner sublayer beneath the prismatic
structure is composed of complex crossed lamellar
structure, exhibiting shorter and more irregularly
shaped first-order lamellae in comparison with
the outer sublayer.
DISTRIBUTION OF THE PRISMATIC AND
CROSSED LAMELLAR STRUCTURES
The prismatic structure together with the outer
and inner crossed lamellar sublayers constitute
the middle shell layer which is the most complex
in terms of structure distribution. The middle
layer thins both in radial and antero-posterior
directions (Fig. 3). In the umbonal region, between
the upper boundary of the adductor muscle scar
and the beak, the middle layer intertongues bet-
ween the foliated structure of the outer and inner
shell layers (Fig. 5). In this region the complex
crossed lamellar structure forming the inner
sublayer presents an irregular bottom surface
(P1.91, fig. ld), draping the regular foliated struc-
ture of the inner shell layer (P1.91, fig. le).
The crossed lamellar structure constitutes the
innermost, extremely thin internal shell layer
beneath the regular foliated calcite of the outer
shell layer in the region between the adductor
muscle scar and the pallial line (Fig. 3). Because
of its very reduced thickness, in fossil specimens
the innermost crossed lamellar layer is generally
eroded and in the muscle scar area is always
absent. In a few specimens the only trace of the
crossed lamellar layer is a faint, white stripe in
the proximity of the pallial line.
A prominent feature of the outer crossed lamellar
sublayer is its undulating top boundary in contact
with the regular foliated calcite, each furrow cor-
FIGURE 7 - A, transverse, vertical section t h r o u g h the adductor
muscle scar area, showing the calcitic homogeneous (a), cros-
sed foliated (b), regular (c), and irregular foliated (d) structure
of the outer shell layer. The aragonitic crossed lamellar (e),
and prismatic structures (f) of the middle shell layer is also
shown. Note the undulating top boundary of the crossed lamel-
lar sublayer ; each furrow corresponding to the development of
the internal ribs. The i n t e r n a l shell surface is a t the base of
the figure. Can Rosell (CR-7). Scale bar : 250 1Jm. Note in B the
a r r a n g e m e n t in pairs of the furrows (arrows). Scale b a r : 0.5
l~m. A, coupe transversale ~t travers la rdgion du muscle adduc-
teur. a, structure homog~ne ; b, feuilletde-croisde ; c, feuilletde-
rdguli~re; d, feuilletde-irrgguli~re ; e, lamellaire-croisde et f, pris-
matique. A noter la surface ondulde qui sgpare la couche exter.
ne (a, b, c et d) de la couche moyenne (e et 19 ; chaque ddpression
correspond au dgveloppement des cStes internes. B, on observe
l'agencement des ddpressions en deux paires (fl~ches).
responding to the development of the internal ribs
(Fig. 7A, B). This undulating boundary is only
observed in transverse sections since the early
start of rib growth before attaining the internal
shell surface.
The prismatic structure reaches its m a x i m u m
thickness in the adductor muscle scar area. It
thins toward the shell margin and intertongues
within the crossed lamellar structure (Fig. 3 ; P1.
91, fig. 3-4). The outer and inner crossed-lamellar
sublayers are in contact w h e n the prismatic
sublayer disappears (P1. 91, fig. 3) ; nevertheless
the inner crossed-lamellar sublayer thins out
toward the margins of the shell (Fig. 3). In several
specimens, however, the prismatic structure in the
muscle scar area has been eroded, and the 0nly
observed layer is the outer foliated calcite.
In specimens displaying diagenesis the middle
layer is leached and infilled with sparry calcite
(Fig. 8).
 721
FIGURE 8 - Slightly oblique, radial, vertical section of the right
valve, portraying the outer calcitic foliated layer (a), the midd-
le layer (b), and the inner calcitic foliated layer (c). Sparry cal-
cite infilling the middle shell layer in this diagenetically-alte-
red shell. The ventral shell margin is toward the right.
Altafulla (AT-333/334). Scale bar : 50 pro. Coupe radiale, ldg~-
rement oblique de la valve droite montrant la couche externe de
calcite feuilletde (a), la couche moyenne avec remplissage de
calcite (b) et la couche interne de calcite feuilletde (c) dans un
exemplaire affectg p a r la diagen~se.
EXTERNAL SHELL SURFACE
FEATURES AND MICROSCULPTU-
RE
The left and right valves ofA. baranensis exhibit
the s a m e m i c r o s t r u c t u r e and mineralogy,
although both differ in the microstructure of the
outermost layer during their early growth stages.
In only a few specimens, the external surface cor-
responding to these stages can be observed (P1.
92, fig. 1-2).
The ribbon (P1. 92, fig. 2, arrow) and earliest dis-
soconch of the right valve show an 3 irregular
polygonal a r r a n g e m e n t (P1. 92, fig. 3) whereas
the prodissoconch does not display even at high
magnification such polygonal p a t t e r n (P1.92, fig.
5). In the ribbon and earliest dissoconch each
polygon is composed of numerous crystallites (P1.
92, fig. 4). This polygonal p a t t e r n is similar to the
p a v e m e n t simple prismatic structure described
by Fatton & Bongrain (1980) in Pecten and
Chlamys, Schein (1989) in deep sea Pectinidae,
and C a r t e r (1990, p. 379) and Barrera et al.
(1990) in A d a m u s s i u m colbecki. The polygonal
and non-polygonal layers t e r m i n a t e early in
ontogeny attaining only 1.7 m m height from the
origin of growth.
The pro- and earliest dissoconch of the left valve
present similar patterns (P1. 92, fig. 6-7) to those
of the prodissoconch of the right valve. This non-
polygonal layer reaches up to 1.1 m m height from
the origin of growth, hence the smooth external
surface in the left valve is narrower t h a n in the
right valve. In the left and right valves, the outer-
most layer with the polygonal and non-polygonal
a r r a n g e m e n t of crystallites overlies the foliated
structure.
The adult shells ofA. baranensis are also charac-
terized by a well developed external divaricate
ornamentation composed of pairs of fine, widely
divergent Camptonectes striae (Freneix et al.
1982). The divaricate microsculpture extends onto
the umbonal area and auricles in both valves. It
initiates at the boundary of the smooth early
growth stage of the dissoconch where the external
radial costae appear P1.93, fig. 1-2).
The granular sublamina within the homogeneous
structure is responsible for the initiation of the
external radial costae. Toward the ventral margin
the costae are made up by the granular sublami-
na together with the foliated structure (P1.93, fig.
1-2). The thickening of the foliated structure, lea-
ding to stronger costae, is achieved by interton-
guing lenses of foliated calcite (P1.93, fig. 3-4). The
costae gradually disappear toward the ventral
m argin.
The divaricate microsculpture consists of micro-
scopic striae oblique to and divergent from the
external costae (P1.93, fig. 2-5-6) which are com-
posed of the granular sublamina of the homoge-
neous structure (P1. 93, fig. 5). Superimposed on
these divaricate striae are small and concentric
growth scales composed of the outer porcella-
neous sublamina of the homogeneous structure
(P1. 93, fig. 6).
In A. baranensis the Camptonectes microsculptu-
re therefore appears to originate from the homo-
geneous structure, unlike Delectogecten randolphi
and Patinogecten (Mizuhopecten) yessoensis where
it is formed by the foliated calcite sublayer
(Hayami & Okamoto 1986).
SEM observations on external surfaces of partial-
ly eroded specimens allowed the microstructural
composition of the external radial costae and
divaricate microsculpture (P1. 93, fig. 5) to be
illustrated.
PHYLOGENETIC STATUS AND
STRATIGRAPHICAL RANGE IN
SPAIN
Freneix et al. (1982) gave a generic status to
Amussiopecten which was considered as a subge-
nus by other authors. Alternatively it is possible
 722

F I G U R E 9 - 8180 a n d 5 ~ C v a l u e s o f n °
180 13C location age code
t h e v e n t r a l m a r g i n o f A . baranen- samples X max mla X max rain
sis (in %°). (1) G . f o h s i p e r i p h e r o -
r o n d a - G , m a y e r i Z o n e ; (2) G . 13 -1.39 -0.95 -2.44 1.44 1.70 1.21 Can Resell Langhian (N8) CR
14 -0.87 -0.33 -1.89 1.66 2.02 1.15 Can Jeira Langhian (N8) CJ
m a y e r i Zone; (3) G. e l o n g a t u s 11 -0.62 -0.46 -0.93 1.49 1.80 1.21 Els Monjos Langhian (NS) MJ
Z o n e . Valeurs isotopiques du ~73C 9 -1.50 -1.00 -2,04 -0.32 2.24 -1.39 Cases Virgfli Langhian (N8) CV
et du 5t"O de la calcite du bord
palldal chez A. b a r a n e n s i s (en %o). 6 -L29 - 0 , 6 2 -2,12 0.22 1.40 -1.74 La Nou de Gai~i Langhian (N9) LN

82 -2.20 -1.16 -3,59 -0.07 1.28 -1.49 Masos Vespella Langbdan-Serravalian(1) MV

-2.23 -1.44 -3.15 -1.08 -0.20 -3.60 Vespella Langhian-Serravalian (1) TG

5 -2.62 - 0 . 4 6 -3,78 -1.78 1.88 -3.05 Ermita de Berh Serravalian (2) BA

4 -1.63 - 1 , 3 3 -1,98 -0.68 2.70 -2.75 Catllar Serravalian (2) CC

1 -2.75 3.00 AltafuUa Tortonian AT

55 0.34 1.08 -3.05 1.10 1.81 O.00 Capr6s-E1Tale Middle Messinian (3) CT
162 0,08 1.86 -1.07 0.97 1.90 -0.17 Campos del Rio Middle Messinian ML

4.
3" .AT
2,
• CR "CJ.MJ
3 1
0 .LN
,CV
-CC
-1" • TG
• BA
-2
oxygen 18 (PDB)
F I G U R E 1 0 - C r o s s p l o t of 5~sO a n d 813C a v e r a g e v a l u e s f r o m t h e
ventral m a r g i n o f A. baranensis. S e e fig. 9 for c o d e s .
Corrdlation entre les valeurs du 513C et du 5180 de la calcite du
bord pallgal chez A. b a r a n e n s i s . Voir abrdviations de la fig. 9.
that A m u s i u m was phylogenetically derived from
Amussiogecten as has been postulated by Freneix
et al. (1982) and Hayami (1988) among others.
Furthermore Hayami (1988) has pointed out that
A m u s i u m was phylogenetically derived from
Amussiopecten-like weakly costate warm-water
species and not from Lentipecten as was conside-
red by some previous authors.
•According to Freneix et al. (1982), in the Mediter-
ranean area and in the Atlantic (Aquitanian
Basin) A. baranensis occurrences range from
Upper Burdigalian (?) to Lower Langhian. In nor-
theastern Spain it ranges from Langhian (N 8
zone) to Tortonian which corresponds to the last
deposits cropping out in this basin. In the Fortuna
and Mula basins (southeastern Spain), however, it
also occurs in Middle and Upper Messinian marl
successions thus expanding its stratigraphical
range in the Mediterranean area according to our
findings (Zamarrefio et al. 1992) and those of Ben
Moussa et al. (1987).
ISOTOPE DATA
A total of 300 specimens ofA. baranensis collected
at 12 localities in northeastern and southeastern
Spain were analyzed for oxygen and carbon isoto-
pic composition of the outer homogeneous and
foliated low-magnesium calcite layer. The speci-
; .ML"CT mens cover a timespan from the Langhian (N 8
Zone) to the Middle Messinian.
Maximum, minimum and average 5180 and 51~C
values as well as the stratigraphical position of
samples in each locality are reported in Fig. 9.
Average values are plotted on Fig. 10.
Based on 51~0 and 513C values of A. baranensis
from the different sections, several considerations
should be stressed. Fig. 10 shows that in general
the values are relatively tightly clustered. The
observed average range of 5180 is 2.96 %0, varying
from -2.62 to 0.34 %o whilst for 513C it is 3.44 %o
and varies frorn 1.66 %o to -1.78 %0. These values
are very close to the values of the low-magnesium
calcite inorganically precipitated in isotopic equi-
librium with sea-water (Morrison & Brand 1986),
suggesting that the isotope signals ofA. baranen-
sis are not largely affected by vital effects.
Consequently the isotopic signatures of the shells
investigated should represent the original compo-
sitions without significant postdepositional alte-
ration, hence reflecting the paleoenvironmental
and paleoceanographic changes affecting the
water masses.
The isotopic signatures from the Middle Messinian
samples at Capr6s-EI Tale (CT) and Campos del
Rio (ML) sections were used to analyze the main
paleoceanographic changes that occurred in these
basins. In both sections the d180values are enriched
in 180 reflecting the increased evaporation and sali-
nity of sea-water in the Mula and Fortuna Basins
prior to the Messinian salinity crisis (V~zquez et
al. 1992). Moreover, both 51~0 and 513C data display
throughout the sections high frequency variations
 723
(5'80 +1%°, 513C +0.5%0) suggesting, together with
trace-element data, paleoceanographic changes in
primary productivity, salinity, oxygen content and
temperature affecting the bottom waters (Vfizquez
et al. 1992).
A similar paleoceanographic study based on 5180,
513C and trace-element values ofA. baranensis (CR,
CJ, MJ, CV, LN, MV, and CC sections) is at present
under progress in the northeastern Miocene basins
of Spain (Zamarrefio, Porta & Vfizquez).
Preliminary data indicates that in general both
5180 and 51~C tend to be lighter from the N8 Zone
toward the N10 Zone (Fig. 9-10).
The isotopic signature of one specimen from
Altafulla (AT) is slightly shifted to lighter 5180
values whilst it is enriched in 1sC. This variation
of the isotopic signals seems to suggest the
influence of meteoric diagenesis. SEM observation
shows the complete replacement of the middle
aragonitic shell layer by a coarser mosaic of spar-
ry calcite (Fig. 8). In spite of the fact t h a t SEM
observations on thelow-magnesium calcite outer
and inner shell layers of this specimen reveal
noalteration in the microstructure, their isotopic
signature indicates t h a t thedissolution-reprecipi-
tation process by meteoric water near the equili-
brium withatmospheric C02 could also influence
the isotopic signature of the calcite layers. Similar
isotopic and trace-element diagenetic changes
without microstructural alteration ofbiogenic cal-
cites have been previously described by Morrison
& Brand (1986) and Brand (1987).
CONCLUSIONS
The adult shell microstructure of A m u s s i o p e c t e n
baranensis (Pectinidae) is characterized by an
outer LMC calcite homogeneous and foliated
layer, a middle aragonitic layer and an inner LMC
foliated layer. The outer foliated layer maintains
its thickness in both valves.
The early stages of ontogeny are different in both
valves. A non-polygonal layer extends from the
prodissoconch to the earliest dissoconch in the left
valve. In the right valve, layer occurs in the pro-
dissoconch whilst in the earliest dissoconch an
irregular polygonal layer is present.
Three sublayers constitute the middle shell layer:
1) aragonitic crossed lamellar, 2) aragonitic pris-
matic occurring in the muscle scar area, thus
exhibiting an asymmetrical distribution, and 3)
aragonitic complex crossed lamellar.
Divaricate microsculpture, external radial costae
and strong internal radial ribs are common lea-
tures in A. baranensis. The external costae origi-
nate from the granular sublamina of the homoge-
neous structure together with lenses of the folia-
ted structure. The internal ribs are formed by the
irregular foliated structure exhibiting a dendritic
p a t t e r n whilst the divaricate microsculpture
develops from the granular sublamina of the
homogeneous structure.
The 5aO values of the outer homogeneous and
foliated LMC layer in the ventral margin seems to
suggest t h a t the shell precipitated in isotopic
equilibrium with ambient seawater. Therefore,
the isotopic values have proven to be useful in
paleoenvironmental interpretations.
A c k n o w l e d g m e n t s - We are grateful to Dr Craig Docherty for
the revision of the English version and to Dr J. G. Carter for
his suggestions on a n earlier version of the manuscript. We
also t h a n k the Geobios reviewers Drs. A. Lauriat-Rage and E.
Schein for her valuable comments which have improved this
manuscript. Special t h a n k s to Albert Lleal for providing the
photographs reproduced in Fig. 2. This research was supported
by DGICYT grants PB 88-0050 and PB 91-0097.
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I. ZAMARRENO & A. V~kZQUEZ
Instituto de Ciencias de la Tierra (J. Almera) CSIC
c/Martl i Franqu6s S/RUm
08028 Barcelona, Spain
Jo D E P O R T A
Departament de Geologia Din~mica
Geofisica i Paleontologia
Facultat de Geologia
c/Marti i Franqu6s S/RUm
08028 Barcelona, Spain