Rev. bras. paleontol.

17(3):289-306, Setembro/Dezembro 2014

2014 by the Sociedade Brasileira de Paleontologia
doi: 10.4072/rbp.2014.3.02

TAXONOMY AND PALAEOECOLOGY OF PINNA (P.) CRETACEA (SCHLOTHEIM,

1813) FROM THE UPPER CRETACEOUS OF THE SERGIPE BASIN, BRAZIL
PETER BENGTSON

Institut fr Geowissenschaften, Im Neuenheimer Feld 234, 69120 Heidelberg, Germany.

peter.bengtson@geow.uni-heidelberg.de

WAGIH S. AYOUB-HANNAA

Geology Department, Faculty of Science, Minufiya University, El-Minufiya, Shibin El Kom, Egypt. wagih_hannaa@yahoo.com

FRANZ T. FRSICH

GeoZentrum Nordbayern, Fachgruppe PaloUmwelt, Friedrich-Alexander-Universitt Erlangen-Nrnberg, Loewenichstr.

28, 91054 Erlangen, Germany. franz.fuersich@gzn.uni-erlangen.de

MICHAEL HEINZE

GeoZentrum Nordbayern, Fachgruppe PaloUmwelt, Friedrich-Alexander-Universitt Erlangen-Nrnberg,

Loewenichstr. 28, 91054 Erlangen, Germany. michael.heinze@gzn.uni-erlangen.de

ABSTRACT The bivalve species Pinna (Pinna) cretacea (Schlotheim, 1813) is redescribed from Turonian and presumed
lower Coniacian beds of the Sergipe Basin, north-eastern Brazil, with the description of the new subspecies P. (P.) cretacea
brevis from the upper Turonian and/or lower Coniacian. Some of the specimens of P. (P.) cretacea show predation scars of
different types. As these bivalves lived with their posterior end of the valves projecting above the sediment surface, they were
at a constant risk of damage by predators, although able to repair their shells relatively easily. The subspecies P. (P.) cretacea
brevis shows a higher repair frequency than the nominotypical subspecies, which is explained mainly by morphological
features (e.g. size, outline, inflation and ornamentation) and by its ability to repair the damaged shell. Thus, the wide, less
convex, strongly ribbed shells of the subspecies brevis display a higher percentage of predation scars. In addition, different
life habits of the two subspecies may have caused differences in repair frequency. As both subspecies possess thin shells,
shell thickness does not appear to be significantly correlated with repair frequency.
Key words: bivalves, Pinna, taxonomy, palaeoecology, predation, Upper Cretaceous, Sergipe Basin, Brazil.
RESUMO O bivlvio Pinna (Pinna) cretacea (Schlotheim, 1813) redescrito do Turoniano e provvel Coniaciano inferior
da bacia de Sergipe, nordeste do Brasil, enquanto a nova subespcie P. (P.) cretacea brevis descrita do Turoniano superior
e/ou Coniaciano inferior da bacia. Alguns dos espcimes de P. (P.) cretacea mostram marcas de predao. Como estes
bivlvios viveram com a parte posterior das valvas projetando-se acima da superfcie do sedimento, estavam submetidos ao
constante risco de danos por predadores. No entanto, eram capazes de reparar suas conchas de forma relativamente efetiva.
A subespcie P. (P.) cretacea brevis mostra uma frequncia maior de reparo que a subespcie nominotpica, o que se explica
principalmente pelas caractersticas morfolgicas (tamanho, forma, inflao e ornamentao da concha) e pela sua capacidade
de reparar a concha danificada. Assim, as conchas largas, menos convexas e ornamentadas por costelas fortes, caractersticas
da subespcie brevis, mostram uma porcentagem maior de cicatrizes de predao. Alm disso, diferentes modos de vida das
duas subespcies podem ter causado diferenas na frequncia de reparo. Como ambas as subespcies possuem conchas finas,
a espessura da concha no parece ser significativamente correlacionada com a frequncia de reparo.
Palavras-chave: bivlvios, Pinna, taxonomia, paleoecologia, predao, Cretceo Superior, bacia de Sergipe, Brasil.

INTRODUCTION

Formation of Sergipe and may reach into the lower Coniacian.

They are particularly abundant in the uppermost parts of the
succession, previously referred to the Coniacian but now
dominantly upper Turonian (see below). The first report of
Pinna from the Cretaceous of Brazil was by White (1887,
p. 58), who mentioned a fragmentary specimen from the
Albian of Sergipe. However, in her revision of Whites
material, Maury (1937) did not describe or mention this

Cretaceous marine carbonate rocks are well exposed in the

Sergipe Basin in north-eastern Brazil (Figures 1A,B, D). The
succession contains a rich macroinvertebrate fauna dominated
by ammonites, bivalves, gastropods and echinoids.
Well-preserved specimens of the bivalve genus Pinna
occur in the lower to upper Turonian of the Cotinguiba
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specimen. The first systematic description of a species

referred to Pinna was by Maury (1930), from the uppermost
Cretaceous of the Pernambuco-Paraba Basin, although this
species is now referred to the genus Atrina (Muniz, 1993).
From the Albian of the Camamu Basin, Bahia, Brito (1967)
described two specimens as Pinna aff. P. decussata (Goldfuss,
1837), here tentatively referred to P. (P.) cretacea cretacea
(Schlotheim, 1813). From the Sergipe Basin, Duarte (1936,
1938) and Wanderley (1938) reported Pinna from beds
that are now known to be Barremian and late Turonian to
early Coniacian in age. Subsequently, Beurlen (1964) noted
the presence of a species of Pinna in the Aptian-Albian
Riachuelo Formation of Sergipe, and Brito (1981) remarked
on the abundance of the genus in the uppermost part of the
Cenomanian-lower Coniacian Cotinguiba Formation. More
recently, Seeling & Bengtson (2003) studied the taxonomy
and palaeobiogeography of Pinna cretacea from the Sergipe
Basin, on the basis of the material restudied here.
According to Vermeij (1987) and Dietl & Alexander
(2005), the persistence of the Pinnidae since the Carboniferous
in an increasingly hazardous world with numerous new
and powerful shell-crushing predators is possibly the best
testimony of the success of this family to cope with shell
damage. After suffering damage by predators, pinnids are able
to repair their shell rapidly (Yonge, 1953). However, repaired
valves are rarely shaped like the original valves. This results
in great variations in outline and ornamentation (Rosewater,
1982), which may cause problems in identification.
The aims of this paper are (i) to document the pinnids from
the Sergipe Basin in the collections of P. and S.I. Bengtson
(coll. 1971-1972), (ii) to describe the nominotypical subspecies
and the new subspecies Pinna (Pinna) cretacea brevis from
the upper Turonian and/or lower Coniacian of Sergipe, (iii) to
analyse scars of unsuccessful predation, observed in some of
the specimens, (iv) to compare the repair frequency in the two
subspecies, and (v) to elucidate the relationship between repair
frequency and crush-resistant features in pinnids (e.g. shell
thickness, valve size, outline, and ornamentation).

marine carbonate succession (Aptian-Albian Riachuelo and

Cenomanian-Coniacian Cotinguiba formations, Figure 1D)
is well exposed and highly fossiliferous. The Cotinguiba
Formation normally reaches a thickness of c. 200 m, but
locally exceeds 1000 m in the onshore part of the basin
(Bandeira, 1978; Koutsoukos & Bengtson, 1993). The
formation is overlain by the siliciclastic Santonian-Miocene
Calumbi Formation. The Cotinguiba Formation consists
mainly of grey to blue-grey (cream or yellowish when
weathered), massive, bioturbated, fine-grained limestones
with sparse siliciclastic intercalations at the base (Figure 1D).
Bengtson (1983) provided a summary of the literature on the
Cenomanian-Coniacian succession of the basin. More recent
data are found, for example, in Andrade (2005).
Koutsoukos & Bengtson (1993) discussed a number of
problems in establishing a detailed macrofossil zonation for
the Sergipe Basin, which are mainly due to the discontinuous
and small outcrops (usually only a few metres of section) in
relation to the total thickness of the succession (well over
500 m). This, in combination with local shifts in strike and
dip, makes a precise stratigraphic positioning of individual
specimens difficult.
In earlier studies of the Sergipe succession, the TuronianConiacian boundary was generally positioned at the first
appearance of the ammonite genera Barroisiceras, Forresteria
and Prionocycloceras, corresponding to the traditional
boundary position, which led to a considerable part of the upper
Cotinguiba succession being referred to the Coniacian. In later
years, pending ratification of a GSSP, the stage boundary is
generally conceived as marked by the first occurrence of the
inoceramid bivalve Cremnoceramus deformis erectus (Meek,
1877) (Kauffman et al., 1996), which results in a considerable
upward shift of the boundary in Sergipe. Thus, only the topmost
beds of the Cotinguiba Formation, containing the ammonite
Peroniceras, are now undoubtedly referred to the Coniacian
(S.I. Bengtson et al., 2005).

GEOLOGICAL SETTING AND STRATIGRAPHY

The material was previously studied by Seeling & Bengtson

(2003) and comprises 35 specimens of Pinna (P.) cretacea from
seven quarries exposing the lower Turonian to lower Coniacian
of the Cotinguiba Formation, Sergipe Basin (Figure 1C). Two
of their specimens were excluded from our study for reasons
explained in the discussion of the species below. For locality
descriptions, see Bengtson (1983). The material consists of
moderately well-preserved, articulated composite and internal
moulds, with parts of the original shell preserved only in a few
specimens. Some specimens display traces of repaired shell
damage, which occur on both valves and occasionally on a
single valve. Repair scars are identified only in well-preserved
specimens. Repair frequency (f) has been standardized for
articulated specimens only (Vermeij & Dudley, 1985) and is
defined as the number of repairs in a sample divided by the
total number of specimens in the sample.
The quarries, from where the specimens were collected,
expose approximately 5-20 m of the Cotinguiba succession.

The Sergipe Basin (Figures 1A,B) is one of four subbasins

(Cabo, Alagoas, Sergipe and Jacupe subbasins; SouzaLima et al., 2002) composing the Sergipe-Alagoas Basin in
north-eastern Brazil. The Sergipe Basin comprises an area of
approximately 11,000 km2, of which 6,000 km2 are onshore.
Structurally the basin consists of a series of half-grabens with
a regional dip averaging 10-15 to the southeast, resulting
from NE-SW trending normal faults. This regional dip causes
progressively younger rocks to crop out towards the coast.
The geological evolution and the development of the marine
Cretaceous sediments of the basin have been discussed by,
for example, Ojeda & Fugita (1976), Ojeda (1982), Bengtson
(1983), Koutsoukos & Bengtson (1993), Koutsoukos et al.
(1993), Feij (1995) and Souza-Lima et al. (2002).
The Sergipe Basin contains a thick succession of nonmarine and marine rocks. In particular, the Cretaceous

MATERIAL AND METHODS

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291

Figure 1. A-B, location of the Sergipe Basin in north-eastern Brazil. C, locality map. D, simplified Cretaceous stratigraphic framework for the
Sergipe Basin (modified from Bengtson, 1983; Hessel, 1988; Smith & Bengtson, 1991; Feij, 1995; Souza-Lima et al., 2002). Abbreviations:
AL, Alagoas; BA, Bahia; CE, Cear; MA, Maranho; PB, Paraba; PE, Pernambuco; PI, Piau; RN, Rio Grande do Norte; SE, Sergipe.

As most of the specimens were collected by quarry workers

without precise information about the sampling levels, only
a broad stratigraphic assignment to substage is possible.
The uncertain stratigraphic position of the new subspecies
Pinna (P.) cretacea brevis, described below, is due to the fact
that the 20 m high quarry (locality Mucuri 7) straddles the
Turonian-Coniacian boundary, as currently conceived (see
above). For localities and chronostratigraphic assignments,
see Appendix 1.
Linear measurements (taken with a Vernier caliper) are
in millimetres. The orientation of measurements and position
of cross-sections are shown in Figure 2. Abbreviations.
L, shell length; H, shell height; W, width of articulated valves;
A, apical angle; nr, number of radial ribs.
The systematic classification follows Carter et al. (2011)
and the morphological terminology the glossary of Cox
(1969). The synonymy lists contain only entries considered to
be of major importance for the present study and which have
been carefully checked by the authors. More comprehensive
synonymies can be found in the references cited.
The specimens are kept in the collections of the Museu
Nacional in Rio de Janeiro, Brazil (collection prefix: MN).
For corresponding specimen numbers of Seeling & Bengtson
(2003) see Appendix 1.

SYSTEMATIC PALAEONTOLOGY
Class BIVALVIA Linnaeus, 1758
Subcohort OSTREIONI de Frussac, 1822
Megaorder OSTREATA de Frussac, 1822
Superorder OSTREIFORMII de Frussac, 1822
(= EUPTERIOMORPHIA Boss, 1982)
Order OSTREIDA de Frussac, 1822
Suborder MALLEIDINA J. Gray, 1854
Superfamily PINNOIDEA Leach, 1819
Family PINNIDAE Leach, 1819
Pinna Linnaeus, 1758
Type species. Pinna rudis Linnaeus, 1758, by subsequent
designation of Children (1823).
Remarks. The genus Pinna Linnaeus, 1758, strongly
resembles the genus Atrina Gray, 1842, in outline and
ornamentation. The principal distinguishing characters
of Pinna are (i) the presence of a median groove in each
valve, (ii) the presence of a sulcus that divides the nacreous
layer into dorsal and ventral parts (Yonge, 1953; Seeling
& Bengtson, 2003; Dietl & Alexander, 2005), and (iii) the

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Figure 2. Measurements and orientation of Pinna cretacea. Abbreviations: see Material and Methods.

position of the posterior adductor muscle scar close to the

antero-dorsal margin. According to Seilacher (1984, p. 228,
text-figure 9), the valves of pinnids actively break along a
line that then forms a median ridge. This ridge heals on the
inside by an elastic shell layer and is not covered by the more
rigid nacreous layer. Seilacher argued that the ridge acts as
a pseudo-ligament which, in conjunction with the ligament,
restores the squarish cross-section after it has been deformed
into a rhomb by the adductor. In the genus Atrina, the nacreous
layer lacks a median sulcus, and its posterior adductor muscle
scar is larger and located near the central part of the valve. In
some composite and internal moulds of the Sergipe material,
the median division of the inner nacreous layer of the shell
into ventral and dorsal lobes can be observed.
Subgenus Pinna Linnaeus, 1758
Pinna (Pinna) cretacea (Schlotheim, 1813)
Remarks. The general shape, style of ribbing and size of
some of the Brazilian specimens strongly resemble those
of Pinna cretacea as described and figured by, e.g. Zittel
(1866), Geinitz (1875), Smettan (1997) and Cleevely &
Morris (2002). Other specimens, in contrast, differ in having
wider and strongly ribbed valves (with intercalatory ribs), a
wider posterior, and a large apical angle (Table 1). For two
complete specimens (of nearly the same size) of each group
the transversal sections were traced throughout ontogeny
(H/W; measured at 5 mm intervals). The transversal

sections of both specimens are strongly similar in the early

growth stages but increasingly different in later growth
stages (Figure 3A). The overlap of all individuals in early
growth stages suggests that they belong to a single species.
However, based on differences in other taxonomic characters
as well, such as general outline (H/L), number, regularity
and distribution of radial ribs, and width of the apical angle
(A), two subspecies, P. (P.) cretacea cretacea and P. (P.)
cretacea brevis are recognized. Box-plot analysis indicates
that shell outline (H/L) and apical angle (A) are the most
significant variables to separate P. (P.) cretacea cretacea
and P. (P.) cretacea brevis (Figure 3B). In Table 1, the two
subspecies are compared in detail.
In their description of the material studied here, Seeling
& Bengtson (2003) also included two incomplete, poorly
preserved specimens, which we consider indeterminate and
therefore choose to exclude from the present study. Specimen
no. GPIH-BR C37.720, from the upper Turonian of locality
Cajaba 7, is a cross-section visible on a rock sample
containing four specimens of P. (P.) cretacea. Although it
is highly probable that also this specimen belongs to P. (P.)
cretacea, the state of preservation precludes determination,
at least to species level. The other specimen, no. GPIH-BR
C380.6, collected from middle or upper Cenomanian rocks
of locality Cruzes 16, is the only specimen from outside the
stratigraphic range of the lot, and so would be of particular
interest. However, the state of preservation precludes
specific identification and doubts may even be raised as to
its generic affinity.

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293

B
Figure 3. A, cross-sections (H/W) of the two subspecies of Pinna (P.) cretacea throughout ontogeny (measured at 5 mm intervals in two
representative samples). Note overlap in juvenile stages. B, box-plot of the valve width (H/L) and the apical angle (as distinct taxonomic
features) for the two subspecies.

Pinna (Pinna) cretacea cretacea (Schlotheim, 1813)

(Figures 4A-C; 5A-F; 10B-D,F-G,I-K)
1813 Pinnites cretaceus Fauj.: Schlotheim, p. 113.
1837 Pinna decussata nobis Goldfuss, p. 166, pl. 128, figs.1-2.
1846 Pinna arata Forbes, p. 153, pl. 16, fig. 10.
1866 Pinna cretacea Schlotheim. Zittel, p. 87, pl. 13, fig. 1.
1871 Pinna arata, Forbes. Stoliczka, p. 384, pl. 25, fig. 1.
1875 Pinna cretacea Schlotheim. Geinitz, p. 54, pl. 14, figs. 2-3.
1875 Pinna decussata Goldfuss. Geinitz, p. 53, pl. 15, figs.
2-3, pl. 6, fig. 1.
1906 Pinna decussata Goldfuss. Woods, p. 99, pl. 13, figs.
4-6, pl. 14, fig. 1 (with extensive synonymy).
1909 Pinna decussata Goldfuss. Wanderer, p. 26, pl. 4, fig. 5.
1930 Pinna cf. P. decussata, Goldfuss. Rennie, p. 174, textfigs. 2-3; pl. 19, fig. 9.
?1934 Pinna ovate Blanckenhorn, p. 180, pl. 7, fig. 5.
1955 Pinna laqueata Conrad. Stephenson, p. 110, pl. 16,
figs. 10-12.

1957 Pinna decussata Goldfuss. Dartevelle et al., p. 68, pl.

9, figs. 1a-c.
?1962 Pinna (Pinna) cenomanensis Abbass, p. 44, pl. 5,
figs. 9-10.
?1967 Pinna aff. P. decussata Goldfuss. Brito, p. 7, pl. 1,
figs. 7-9.
1975 Pinna decussata (Goldfuss). Kri & Soukup, p. 47-49,
pl. 1, fig. 6, pl. 2, figs. 1-3, 5 (non pl. 2, fig. 4).
1986 Pinna (Pinna) cretacea (v. Schlotheim). Abdel-Gawad,
p. 140, pl. 30, fig. 1 (with extensive synonymy for northern
and eastern Europe).
1987 Pinna cf. cretacea (Schlotheim). Dhondt, p. 59, pl. 5,
fig. 8.
1997 Pinna cretacea (Schlotheim) s. str. Smettan, p. 110,
pl. 2, fig. 5.
1997 Pinna cf. P. decussata (Goldfuss) s. str. Smettan, p.
110, pl. 2, fig. 6.
2002 Pinna decussata Goldfuss. Cleevely & Morris, p. 101,
pl. 17, fig. 3.

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294

Table 1. Comparison of shell morphology, number, regularity and distribution of radial ribs, and apical angles between Pinna (Pinna) cretacea
cretacea (Schlotheim, 1813) and P. (P.) cretacea brevis subsp. nov. from Turonian and presumed lower Coniacian of the Sergipe Basin, Brazil.
Pinna (P.) cretacea cretacea (Schlotheim, 1813)

Pinna (P.) cretacea brevis subsp. nov.

Shape

Slender, elongate with sharp apex, (H/L: 0.24-0.41).

Wide, less elongate, pyramidal to wedge-shaped, (H/L: 0.45-0.71).

Shell margins

Dorsal and ventral margins nearly straight, posterior margin

truncated and gaping.

Dorsal margin concave, ventral margin convex.

Median ridge
(pseudoligament)

With sharp crest dividing valve into unequal dorsal and ventral
parts.

Well developed, wide, with rounded crest, dividing valve into

equal dorsal and ventral parts.

Radial ribs

7-18.

15-28, with fine intercalations.

Intercalations

None.

Present on ventral and dorsal sides.

Ventral area

Upper third part ornamented with 3 to 7 radial ribs, rest of valve Upper two-thirds ornamented with 5 to 11 radial ribs, rest of
with growth lines only.
valve with growth lines only.

Dorsal area

Ornamented with 4 to 11 radial ribs, separated by regular

interspaces, forming a net-like structure with growth lines.

Umbones

Terminal, sharply pointed anteriorly.

Wide, rounded.

Apical angle

24-38.

51-59.

Cross-section

See Figure 6.

See Figure 6.

2003 Pinna cretacea (Schlotheim). Seeling & Bengtson, p.

477, figs. 4A, C.
2005 Pinna cf. P. cretacea (Schlotheim). Hradeck et al., p.
29, pl. 1, figs. l, m.
2013 Pinna cretacea (Schlotheim). Schneider et al., p. 576,
text-figs. 9C, F.
Material. 27 composite moulds, nos. MN 8750-I to MN 8769I and 8973-I to MN 8979-I, from the lower to upper Turonian
of localities Mucuri 9, Cajaba 7, Mata 10, Laranjeiras 21,
Laranjeiras 22, and Pedra Furada 5, Cotinguiba Formation,
Sergipe Basin, Brazil.
Description. Medium to large (L = 105-200 mm), outline
slender to narrowly triangular, strongly elongate (H/L = 0.35 on
average), strongly inequilateral, equivalved, moderately inflated
and pointed anteriorly (Figures 4A; 5A-B). Cross-section ovate
angular near the anterior end ( of the total valve length from
the anterior end, Figure 6A), rhombic at mid-length of valve,
and elongate-rhombic towards the posterior end (Figure 6B).
Ventral margin straight to slightly convex; dorsal margin
straight to slightly concave. Posterior end gaping and truncated.
Umbones sharply pointed, located at the anterior end, with
small apical angle (24-38). Semi-circular posterior muscle scar
preserved on a few internal moulds, located dorsally (40% of
the total valve length from the anterior end). Anterior muscle
scar not visible. Well-developed median ridge running from
umbo towards posterior end in each valve, dividing the valves
into two unequal parts (Figure 5A). Dorsal part smaller and
more inflated than ventral part, and ornamented with 4 to 11
radial ribs (with rounded crests), separated by wide and shallow
interspaces. Ribs crossed by fine growth lines forming a net-like
structure (Figure 5D). Ventral part of valve ornamented with
three to seven well-developed radial ribs, occupying the upper
part of the median ridge; remaining part of valve crossed only
by commarginal growth lines curving towards the umbones and
forming an acute angle with the radial ribs. Internal structures
such as hinge and ligament not visible.

8 to 20 radial ribs, separated by irregular interspaces.

Measurements. See Table 2.

Discussion. Pinna (Pinna) cretacea cretacea is characterized
by nearly straight dorsal and ventral margins, very elongate,
inflated valves, a slender outline, an acute apical end, a welldeveloped median ridge, and the presence of 7-15 strong radial
ribs separated by broad, smooth interspaces.
Pinna (Pinna) cretacea cretacea is very similar to
P. (P.) decussata Goldfuss, 1837, reported from the Upper
Cretaceous of different parts of the world (see synonymy list).
The two species were synonymised by Abdel-Gawad (1986),
Dhondt (1987) and Seeling & Bengtson (2003). Although P.
(P.) cretacea cretacea strongly resembles P. (P.) decussata in
outline and ribbing style, it differs in being more elongate.
In addition, P. (P.) decussata has a wide apical angle and its
ventral ridges are more curved (Cleevely & Morris, 2002).
According to Abdel-Gawad (1986), the difference between
P. (P.) cretacea cretacea and P. (P.) decussata lies only in
the strength of longitudinal ribs and in the cross-section,
differences assumed to be of ecological significance and
substrate-controlled. Dhondt (1987) also pointed out that the
differences between the two species are minor and have not
been clearly defined. In agreement with the latter author, we
regard the differences between the two species insufficient
for separation at the species level. Similarly, a specimen of P.
laqueata Conrad, 1858, figured and described by Stephenson
(1955) from the Maastrichtian of Missouri, USA, strongly
resembles the present species in general outline, size, and
ribbing style. Abdel-Gawad (1986, p. 141) noted that the
internal characters of Stephensons specimen are very similar
to those of P. (P.) cretacea. Therefore, P. laqueata is also
considered a junior synonym of P. (P.) cretacea cretacea.
Pinna arata Forbes, 1846, from the Upper Cretaceous of
India was originally described on the basis of the presence
of intercalated ribs between the primary ribs on the dorsal
side (Stoliczka, 1871, p. 384, pl. 25, fig. 1b). However,
Forbes species is based on fragmentary internal moulds
and identification must be considered uncertain. In addition,

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295

Table 2. Measurements (mm) of Pinna (Pinna) cretacea cretacea (Schlotheim, 1813) from the lower to upper Turonian of the Sergipe Basin,
Brazil. Abbreviations: see Material and Methods.
Specimen no.

H/L

MN 8750-I

170

50

43

37

0.29

nr
7

MN 8756-I

185

73

42

38

0.39

12

MN 8757-I

195

77

43

33

0.39

11

MN 8758-I

150

58

35

33

0.39

15

MN 8759-I

155

53

24

30

0.34

11

MN 8760-I

125

42

36

27

0.33

MN 8762-I

170

68

37

32

0.40

12

MN 8763-I

200

67

51

31

0.33

MN 8764-I

175

65

38

36

0.37

10

MN 8765-I

175

67

35

38

0.38

11

MN 8766-I

190

78

42

35

0.41

10

MN 8767-I

155

53

22

24

0.34

10

MN 8768-I

170

42

20

25

0.24

13

MN 8977-I

145

50

25

32

0.34

--

MN 8978-I

178

70

35

28

0.39

11

MN 8979-I

105

40

28

38

0.38

--

Range

105-200

40-78

20-51

24-38

0.24-0.41

7-15

Mean

165.19

59.56

34.75

32

0.35

10

intercalatory ribs are also seen in some specimens of P. (P.)

cretacea cretacea from Sergipe. Seeling & Bengtson (2003)
regarded the presence of intercalatory ribs to fall within the
range of variation of P. (P.) cretacea. Therefore, we consider
P. arata a junior synonym of P. (P.) cretacea cretacea.
Pinna aff. P. decussata, described by Brito (1967) from
the Albian of the Camamu Basin, Bahia, Brazil, is here
assigned tentatively to P. (P.) cretacea cretacea, due to the
poor preservation (incomplete internal and external moulds).
P. laticostata Stoliczka, 1871 from the Upper Cretaceous
of India (Stoliczka, 1871, p. 385, pl. 25, figs. 2,3) was also
described on the basis of poorly preserved, incomplete
specimens with damaged umbonal areas. The specimens
differ from P. (P.) cretacea cretacea in having fewer ribs
and lacking a median ridge. We consider identification of P.
laticostata questionable, not only at the species level but also
at the generic level. Similarly, P. (P.) cenomanensis Abbass,
1962, from the Cenomanian of the Western Desert of Egypt,
is based mainly on internal moulds. However, the general
outline, size, and the sharp apical end of Abbass specimens
are similar to those seen in P. (P.) cretacea cretacea, which
makes conspecificity probable.
Pinna robinaldina dOrbigny, 1844, as figured by Woods
(1905, p. 96, pls. 12:11-15, 13:1) from the Lower Greensand
of England, strongly resembles P. (P.) cretacea cretacea but
differs in being more elongate with fewer, irregular radial ribs
on the ventral area, lacking intercalatories, and in having a
small apical angle.
Occurrence. Albian-Maastrichtian of Europe, e.g. Germany
(Geinitz, 1875; Wanderer, 1909; Smettan, 1997; Schneider
et al., 2013), the Czech Republic (Kri & Soukup, 1975),
Poland (Abdel-Gawad, 1986), Austria (Dhondt, 1987;
Hradeck et al., 2005) and England (Woods, 1906; Cleevely

& Morris, 2002); Cenomanian-Turonian of the Middle East,

e.g. Syria and Palestine (Blanckenhorn, 1934); probably the
Cenomanian of Egypt (Abbass, 1962); possibly the Albian of
Bahia, Brazil (Brito, 1967); Turonian of Sergipe, Brazil (this
study); Maastrichtian of North America (Stephenson, 1955);
Upper Cretaceous of India (Forbes, 1846; Stoliczka, 1871).
Pinna (Pinna) cretacea brevis subsp. nov.
(Figures 7; 8A-B; 9A-E; 10A,E,H,L)
?1942 Pinna decussata Goldfuss. Stchepinsky, p. 21, 55,
pl. 4, fig 3.
?1975 Pinna decussata (Goldfuss). Kri & Soukup, p. 47-49,
pl. 2, fig. 4.
2003 Pinna cretacea (Schlotheim). Seeling & Bengtson,
p. 477, fig. 4B.
2011 Pinna cretacea (Schlotheim). Andrade & Santos,
p. 231, figs. 2:1-2.
Etymology. brevis (Latin) for short, referring to the short
valves.
Holotype. Specimen no. MN 8980-I (Figure 7) from the upper
Turonian or lower Coniacian of locality Mucuri 7, Cotinguiba
Formation, Sergipe Basin, Brazil.
Diagnosis. Pinna (Pinna) cretacea with very large, wide,
wedge-shaped valves, numerous radial ribs with fine
intercalatories, well-developed commarginal growth rugae,
relatively wide apical angle, well-developed median ridge
with rounded crest, wide and rounded umbones, slightly
concave dorsal margin, and convex ventral margin.
Material. Eight specimens (most of which with shell
preserved), holotype no. MN 8980-I and paratypes nos.
MN 8981-I to MN 8987-I, from the upper Turonian and/or

REVISTA BRASILEIRA DE PALEONTOLOGIA, 17(3), 2014

296

E
A

Figure 4. Pinna (Pinna) cretacea cretacea (Schlotheim, 1813) from

the upper Turonian of locality Cajaba 7 (MN 8763-I). A, side view of
left valve; B, ventral view; C, dorsal view. Scale bars = 10 mm.

Figure 5. Pinna (Pinna) cretacea cretacea (Schlotheim, 1813). A-B,

articulated composite mould, upper Turonian of locality Cajaba 7 (MN
8765-I): A, side view of left valve showing median groove; B, ventral
view. C, side view of left valve, locality Cajaba 7 (MN 8760-I). D, close-up
showing net-like ornamentation, upper Turonian, locality Mucuri 9
(MN 8750-I). E, left lateral view of internal mould, upper Turonian of
locality Mata 10 (MN 8979-I). F, dorsal view showing concentric rugae,
locality Cajaba 7 (MN 8759-I). Scale bars = 10 mm.

Figure 6. A, sketch showing position of the cross-sections (CS). CS1: 25% of the total shell length from the anterior end; CS2: cross-section
in the middle part of valve; CS3: 75% of the total shell length from the anterior end. B, comparison of transversal sections (in three different
positions from anterior to posterior end of valve CS1, CS 2, and CS3) between Pinna (Pinna) cretacea cretacea (Schlotheim, 1813) and P. (P.)
cretacea brevis subsp. nov.

BENGTSON ET AL. TAXONOMY AND PALAEOECOLOGY OF PINNA (P.) CRETACEA (SCHLOTHEIM, 1813)

297

its posteriorly wide valve and truncated margin but differs

in being smaller (L = 126, H = 68.5 mm, as opposed to L =
220, H = 130 mm on average), in lacking a median ridge and
radial ornamentation, and in its smaller apical angle. Pinna
cottai Geinitz, 1840, from the Upper Cretaceous of Germany
(Geinitz, 1840, p. 55, pl. 11, fig. 1) resembles P. (P.) cretacea
brevis subsp. nov. in having a wide posterior area and a welldeveloped median ridge but differs in having fewer and less
developed radial ribs and an elliptical cross-section at midlength of the valves. Pinna vanhoepeni Rennie, 1930, from the
Upper Cretaceous of South Africa (Rennie, 1930, p. 172, textfig. 1, pl. 18, fig. 9), resembles the present subspecies in having
intercalatory riblets and a concave dorsal margin but differs
in lacking a median ridge, in being taller and narrower, and in
having fewer primary ribs, wide and concave interspaces, and
a straight ventral margin. Although Rennies specimens are
incomplete, the apical angle seems to be smaller than that of P.
(P.) cretacea brevis subsp. nov.. Pinna complanata Stoliczka,
1871, from the Upper Cretaceous of India (Stoliczka, 1871,
p. 384, pl. 14, figs. 3-4), differs in being less elongate and
in having fewer radial ribs, separated by wide interspaces.
Pinna (Plesiopinna) kasimirensis Abdel-Gawad, 1986, from
the Maastrichtian of Poland (Abdel-Gawad, 1986, p. 141, pl.
28, fig. 3, pl. 29, fig. 1) resembles the present subspecies in
having a distinct median ridge, large apical angle (55-65)
and a lenticular cross-section in older growth stages but differs
in being less elongate and in having very wide valves (H/L
= 0.80-0.84) with a thicker prismatic layer. In addition, this
species is ornamented by radial ribs changing into rows of
sparsely distributed scaly nodes on the posterior part.
Occurrence. Upper Turonian and/or lower Coniacian of
Sergipe, Brazil; possibly Cenomanian to Senonian of
Turkey (Stchepinsky, 1942) and Turonian of the Czech
Republic (Kri & Soukup, 1975).

lower Coniacian of locality Mucuri 7, Cotinguiba Formation,

Sergipe Basin, Brazil (Figures 7-9).
Description. Large to very large, pyramidal to broadly wedgeshaped shell, with wide posterior margin (Figure 7), strongly
inequilateral and moderately inflated. Ventral margin convex;
dorsal margin slightly concave. Posterior margin truncated and
slightly gaping. Umbones broad and terminal with an apical
angle ranging from 51 to 59. Median ridge well developed,
with rounded crest, running from umbo towards posterior
margin and dividing valves into two nearly equal parts (Figure
9A). Dorsal part flat to slightly concave, ornamented with 8-20
well-developed radial ribs, separated by irregular, concave,
shallow interspaces. Interspaces widening towards median
ridge, becoming narrower towards dorsal margin. A few
specimens carrying fine secondary ribs intercalated between
the principal radial ribs (Figures 9D-E). Ventral part of shell
flat, ornamented with 5-11 radial ribs, covering two-thirds
of the ventral part from the median ridge towards the ventral
margin (Figures 8A,B). Radial ribs well developed, uniform in
size, and separated by wide, regular, and shallow interspaces.
Commarginal growth rugae well developed towards the
posterior margin in some specimens (Figure 9A).
Measurements. See Table 3.
Discussion. Pinna (Pinna) cretacea brevis subsp. nov. differs
from the nominotypical subspecies in being less elongate (H/L
= 0.51 on average, as opposed to 0.35) and in having more
radial ribs (15-28) with fine intercalatories (Figures 8; 9D,E),
a relatively wide apical angle (51-59, as opposed to 24-38)
and, consequently, a wider valve. For a detailed comparison
of the two subspecies, see Table 1. The subspecies brevis may
reach into the lower Coniacian (Peroniceras spp. ammonite
zone) and is thus slightly younger than the nominotypical
subspecies.
The specimen of Pinna decussata described by Stchepinsky
(1942) from Turkey agrees in general outline with P. (P.)
cretacea brevis subsp. nov., but the poor preservation
precludes a definite assignment. One of the specimens of
Kri & Soukup (1975, pl. 2, fig. 4) from the Czech Republic
also resembles the new subspecies but is incomplete and
assignment must remain tentative. Pinna sp., described by
Smettan (1997, p. 11, pl. 2, fig. 7) from the Cenomanian of
Germany, resembles P. (P.) cretacea brevis subsp. nov. in

PREDATION SCARS
Predation is an important ecological process that often
controls the structure of communities and diversification
and causes long-term morphological and behavioural
trends (Vermeij, 1977, 1987; Kelley & Hansen, 2007).
The repair frequency (f) does not provide quantitative

Table 3. Measurements (mm) of Pinna (Pinna) cretacea brevis subsp. nov. from the upper Turonian and/or lower Coniacian of the Sergipe
Basin, Brazil. Abbreviations: see Material and Methods.
Specimen no.

H/L

nr

MN 8980-I

270

155

45

51

0.57

28

MN 8981-I

--

--

38

51

--

17

MN 8983-I

--

--

41

--

--

16

MN 8984-I

160

100

38

52

0.62

18

MN 8985-I

--

--

43

--

--

15

MN 8986-I

210

95

41

42

0.45

21

MN 8987-I

240

170

38

59

0.71

25

Range

160-270

90-170

38-45

51-59

0.45-0.71

15-28

Mean

220

130

40.57

51

0.59

20

298

REVISTA BRASILEIRA DE PALEONTOLOGIA, 17(3), 2014

B
Figure 7. Holotype of Pinna (Pinna) cretacea brevis subsp. nov. (MN
8980-I) from the upper Turonian or lower Coniacian of locality Mucuri
7, showing left lateral view with shell preservation, the posteriorly wide
valves and the broadly convex ventral margin. Scale bar = 10 mm.

Figure 8. Pinna (Pinna) cretacea brevis subsp. nov. A, right lateral view
in shell preservation showing strongly ribbed and repaired valve (mirrorimage repairs); B, side view of left valve, upper Turonian-lower Coniacian
of locality Mucuri 7, paratype, no. MN 8987-I. Scale bars = 10 mm.

estimates of predation intensity or efficiency but reflects, in

a qualitative way, the relationship between the effectiveness
of the predators crushing ability and the preys resistance to
crushing (Schindel et al., 1982). To calculate repair frequency
(f), we applied the formula of Dietl et al. (2000) and Dietl &
Alexander (2005), defined as the number of repairs in a sample
divided by the total number of specimens in the sample.
Half of the Brazilian specimens of Pinna (P.) cretacea
show patterns of repair scars, cutting the commarginal
growth lines of the valves (Figures 10A-L; 11A). The new
subspecies P. (P.) cretacea brevis subsp. nov. is characterized
by wide valves, a large apical angle, and well-developed
ornamentation (Figures 7-8). The repair frequency of this
subspecies is comparatively high (f = 2.11, Figure 11C), and
most of the repair scars are concentrated in the posterior and
postero-ventral areas. By contrast, the elongate subspecies
P. (P.) cretacea cretacea, with its narrow valves, sharp
anterior end and fewer ribs, exhibits a low repair frequency
(f = 0.86, Figure 11C). The nature of shell breakage was
categorized using the designations of Alexander (1986)
and Dietl et al. (2000), viz. scalloped, embayed, divoted,

cleft. Scalloped injuries include shell breakage that runs

parallel to commarginal growth lines, often cutting the
radial ornamentation, where present. In Recent pinnids
[e.g. Pinna carnea Gmelin, 1791, Atrina rigida (Lightfoot,
1786), Servatrina serrata (G.B. Sowerby, 1825)], the term
scalloped is used to describe the case where the dorsalventral width of the scar exceeds its anterior-posterior length
(Dietl & Alexander, 2005).
In the Sergipe specimens, scalloped scars represent 38.7%
of the total number of scars in Pinna (P.) cretacea cretacea
and 50% in P. (P.) cretacea brevis subsp. nov. (Figure 11B).
Embayed scars are the second largest group of predation
scars, representing 30.7% of the scars in the narrowly elongate
subspecies cretacea and 50% in the wide and short subspecies
brevis (Figure 11B). Embayed valves have lost a large portion
of the shell, showing an irregularly shaped fracture (Figures
10A,B). Divoted valves show triangular to chevron-shaped
depressions, where part of the shell margin was removed,
creating a horseshoe-shaped outline (Figure 10G). This
kind of repair scar represents c. 23% of the scars in P. (P.)
cretacea cretacea. Cleft shell repairs produce slit-like holes

BENGTSON ET AL. TAXONOMY AND PALAEOECOLOGY OF PINNA (P.) CRETACEA (SCHLOTHEIM, 1813)

299

Figure 9. Pinna (Pinna) cretacea brevis subsp. nov. A, right lateral view in shell preservation showing folded growth rugae, upper Turonian-lower
Coniacian of locality Mucuri 7, paratype, no. MN 8986-I; B, side view of left valve of composite mould showing the median groove from the
same locality, paratype, no. MN 8981-I; C, left lateral view of incomplete valve in shell preservation showing the median groove and numerous
radial ribs; D-E, close-up showing intercalatory ribs (Ic) between primary radial ribs (Pr), upper Turonian-lower Coniacian of locality Mucuri 7,
paratype, no. MN 8983-I. Scale bars = 10 mm.

in the valves, more or less parallel to the radial ornamentation

(Figure 10C). Although occasionally curved, the shell rift is
generally confined between any radial ornamentation, and the
growth lines show an inverted V-shaped pattern across the
shear zone. For more details about repair scars, see Alexander
(1986), Dietl et al. (2000), Alexander & Dietl (2001), and
Dietl & Alexander (2005).

DISCUSSION
Pinnid bivalves are suspension-feeders that inhale
water at the posterior end of the valves and, therefore, keep
their truncated posterior margin above the sediment-water
interface. Their pointed anterior end is buried vertically in
soft substrate, attached by numerous byssal threads to hard
particles (e.g. rock or shell fragments) below the sediment

300

REVISTA BRASILEIRA DE PALEONTOLOGIA, 17(3), 2014

B
C

Figure 10. Evidence of shell-crushing predation shown by repair scars in valves of the two subspecies of Pinna (P.) cretacea. A-B, side view of
left valves showing thick irregular folds across the surface and multiple scalloped and embayed repair scars, upper Turonian or lower Coniacian
of localities Mucuri 7 (holotype, no. MN 8980-I) and Cajaba 7 (MN 8757-I), respectively; C, right lateral view showing a slightly curved vertical
cleft repair propagated from the anterior margin in a posterior direction, upper Turonian of locality Cajaba 7 (MN 8765-I); D, side view of left
valve showing a scalloped repair scar, locality Cajaba 7 (MN 8767-I); E, scalloped scar of right valve, locality Mucuri 7 (holotype, no. MN 8980I); F, repair scar of right valve (probably embayed repair), locality Mata 10 (MN 8978-I); G, side view of left valve showing a divoted repair scar
(horseshoe-shaped), locality Cajaba 7 (MN 8759-I); H, large embayed scar of left valve (arrowed), locality Mucuri 7 (MN 8986-I); I, puncture
scar of left valve, locality Cajaba 7 (MN 8768-I); J, side view of right valve showing a puncture scar (divoted? scar), locality Cajaba 7 (MN
8758-I); K, multiple scalloped and embayed repair scars (U-shaped) of right valve, locality Mata 10 (MN 8978-I); L, right valve with mirror-image
repairs of multiple scalloped and embayed scars (U-shaped), locality Mucuri 7 (MN 8987-I). Symbols: + = Pinna (Pinna) cretacea cretacea
(Schlotheim, 1813). * = Pinna (Pinna) cretacea brevis subsp. nov. Scale bars = 10 mm.

BENGTSON ET AL. TAXONOMY AND PALAEOECOLOGY OF PINNA (P.) CRETACEA (SCHLOTHEIM, 1813)

301

C
Figure 11. A, shell repairs (in %) in specimens of Pinna (P.) cretacea from the Sergipe Basin. Note that half of the studied specimens were
subject to predator attacks (repaired valves). B, frequency distribution (in %) of scalloped, embayed, divoted and cleft repairs for each of the
two subspecies. C, repair frequency in the two subspecies.

Figure 12. Suggested scenario of predator-prey interaction and how pinnids repaired their valves after repeated predator attacks during ontogeny.

302

REVISTA BRASILEIRA DE PALEONTOLOGIA, 17(3), 2014

Figure 13. Number of repairs in relation to size (valve length, L) (A); shell outline (B); and number of radial ribs (C) of Pinna (P.) cretacea from
the Sergipe Basin.

surface (Yonge, 1953; Dietl & Alexander 2005). Because

marine shell-crushing predators are active at the surface or
in the uppermost few centimetres of the sediment (Vermeij
& Dudley, 1985), the exposed posterior part of pinnids is
at constant risk of damage by predators, e.g. fishes (Yonge,
1953), currents, storms and/or sediment-loading stress (Checa,
1993). Consequently, epifaunal and semi-infaunal bivalves
generally show higher frequencies of shell repairs than
infaunal taxa. As the Sergipe pinnids occur in fine-grained
lime mudstones and wackestones (Smith & Bengtson, 1991, p.
15) laid down under relatively quiet conditions, predators are
assumed to be the main agent of shell breakage. According to
Vermeij (1983), many bivalves show few shell repairs because
the mantle margins are attached to the shell by pallial muscles,
and a thick periostracum commonly protects against predation
(Yonge, 1953). By contrast, pinnids possess a remarkable
ability of rapid shell repair thanks to their highly retractable
mantle edges, flexible nature of their valves and the vestigial
nature of their periostracum (Yonge, 1953; Seilacher, 1984;
Dietl & Alexander, 2005). As the pinnid shell continuously
grows above the sediment surface, the bivalve strives to
burrow deeper into the sediment in order to reduce the risk
of predatory attacks. After an attack, the mantle extends
towards the posterior end and rapidly repairs the broken
margin through secretion of new shell material. As a result,
irregularly shaped, projecting ridges (repair scars) are formed
across the surface of each valve (Figures 10A,B).
Predatory attacks and subsequent repair of the shell may
occur several times during ontogeny. Figure 12 shows a
suggested scenario of the predator-prey interaction (three
shell-breaking events) during ontogeny and how the pinnids
repaired their valves after repeated predator attacks. According
to Yonge (1953), repeated repair increased the strength of the
shell, because secretion of new shell generally started slightly
anteriorly of the broken edge, forming irregular, thickened
and projecting ridges (Figures 10A,B). Therefore, repeated
predator attacks resulted in increased shell strength and
resistance against subsequent attacks.
According to Taylor & Layman (1972), Carter (1980),
Currey (1988), and Alexander & Dietl (2001) the main
contributing factors of shell breakage in bivalves are: (i) habit

and habitat, (ii) ornamentation and skeletal microstructure,

(iii) soft tissue exposures, and (iv) mechanical or biological
agents. In addition, successful and unsuccessful attacks
depend on shape, size, and rigidity of the valve (Vermeij,
1982). With respect to ornamentation, Checa (1993) and
Dietl et al. (2000) noted a correlation between shell-repair
frequency and number of ribs in Cretaceous exogyrine
oysters. Ornamented forms exhibit higher frequency of
repair than less ornamented and thinner forms. We consider
three possibilities to explain the variation of repair frequency
between the two subspecies of Pinna (P.) cretacea from
Sergipe (f = 2.11 vs. 0.86). Firstly, as discussed above, features
such as size, outline, and ornamentation strongly influence
shell breakage. For instance, repair frequency in the large,
wide, and strongly ribbed valves of the subspecies brevis is
higher than that of the narrow, elongate and less ornamented
subspecies cretacea. Figures 13A-C show a significant
relationship between the number of repairs and shell length,
outline and number of radial ribs. As both subspecies are
thin-shelled, valve thickness is not significantly correlated
with repair frequency. Secondly, the high repair frequency in
P. (P.) cretacea brevis subsp. nov. indicates that shell-crushing
predators were present. By contrast, the low repair frequency
in the nominotypical subspecies was probably related to
complete crushing of the shells by predators. Differences
in burrowing depth between the two subspecies could also
account for the different frequencies of predatory attacks. If
so, then P. (P.) cretacea brevis subsp. nov. was probably less
deeply burrowing than the narrowly elongate P. (P.) cretacea
cretacea and therefore more often subject to predator attacks.

CONCLUSIONS
Two subspecies of Pinna (Pinna) cretacea, P. (P.) cretacea
cretacea (Schlotheim, 1813) and P. (P.) cretacea brevis subsp.
nov., are described from the Turonian to lower Coniacian
of the Cotinguiba Formation, Sergipe Basin, north-eastern
Brazil. The subspecies cretacea is recorded mainly from the
upper Turonian, with a few occurrences from the lower or
middle Turonian, whereas subspecies brevis is only recorded
from the highest levels of the Turonian, possibly reaching into

BENGTSON ET AL. TAXONOMY AND PALAEOECOLOGY OF PINNA (P.) CRETACEA (SCHLOTHEIM, 1813)
the lower Coniacian. Box-plot analysis indicates that shell
outline (H/L) and apical angle (A) are the most significant
features to separate the two subspecies.
Half of the specimens studied show different patterns
of repair scars, formed mainly after predator attacks. Other
ecological factors, such as strong currents and storms,
apparently played no role as the specimens occur in finegrained lime mudstones and wackestones deposited in
relatively quiet waters.
The frequency of shell repair (f) for the two subspecies
differs: f = 2.11 in P. (P.) cretacea brevis subsp. nov. and 0.86
in P. (P.) cretacea cretacea. Variations in repair frequency
may reflect differences in size, shape, ribbing or life habits
(burrowing depths) between the subspecies. The high repair
intensity in P. (P.) cretacea brevis subsp. nov. indicates that
predator attacks were common, though often not lethal. The
low repair frequency in P. (P.) cretacea cretacea may be
related to complete crushing of the shells by predators or to
a more deeply-infaunal life habit, exposing less of the shell
above the sediment surface.

ACKNOWLEDGMENTS
We greatly appreciate the review and valuable suggestions
of S.E. Damborenea (Museo de La Plata, Argentina) and an
anonymous reviewer. Great thanks are also due to B. Leipner
(Institut fr Palontologie, Universitt Erlangen-Nrnberg,
Germany) for the preparation of some of the specimens. P.B.
thanks S.I. Bengtson (Heidelberg), D.A. Campos (DNPM, Rio
de Janeiro) and Petrobras S.A. (Aracaju and Rio de Janeiro)
for field and logistic assistance. The fieldwork was financed
mainly by the Swedish Natural Science Research Council
(NFR, now Vetenskapsrdet, VR).

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Received in April, 2014; accepted in November, 2014.

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306

Appendix 1. List of specimens studied with corresponding specimen numbers of Seeling & Bengtson (2003), localities and chronostratigraphic
assignments.
GPIH-BR numbers and figures
of Seeling & Bengtson (2003)

MN (Museu Nacional, Rio de Janeiro)

numbers and figures herein

Remarks

Locality

Substage

Pinna (Pinna) cretacea cretacea (Schlotheim, 1813)

C32.1

MN 8750-I*; Fig. 5D

C37.7

MN 8751-I

C37.8

MN 8752-I

C37.17

MN 8753-I

C37.18

MN 8754-I

C37.55

MN 8755-I

C37.129

MN 8756-I*

C37.130

MN 8757-I*; Fig. 10B

C37.131

MN 8758-I*; Fig. 10J

C37.132

MN 8759-I*; Figs. 5F, 10G

C37.133

MN 8760-I*; Fig. 5C

C37.189

MN 8761-I

C37.190

MN 8762-I*

C37.191

MN 8763-I*; Fig. 4

C37.192; Fig. 4A

MN 8764-I*

C37.588

MN 8765-I*; Figs. 5A-B, 10C

C37.598

MN 8766-I*

C37.716; Fig. 4C

MN 8767-I*; Fig. 10D

Mucuri 9

Cajaba 7

upper Turonian

Misprint for C37.589

C37.717; Fig. 4C

MN 8768-I*; Fig. 10I

C37.718

MN 8769-I

C37.719

MN 8973-I

C37.720

-------------

C61.38

MN 8974-I

Laranjeiras 21

C62.38

MN 8975-I

Laranjeiras 22

C96.4

MN 8976-I

Pedra Furada 5

C367.3

MN 8977-I*

C367.4

MN 8978-I*; Fig. 10F, K

C367.39

MN 8979-I*; Fig. 5E

C380.6

-------------

Accumulation of specimens,
of which no. C37.720 an
indeterminate cross-section

Indeterminate

lower or middle
Turonian

Mata 10

upper Turonian

Cruzes 16

middle or upper
Cenomanian

Mucuri 7

upper Turonian or
lower Coniacian

Pinna (Pinna) cretacea brevis subsp. nov.

C26.14; Fig. 4B

MN 8980-I*; Figs. 7, 10A, E

Holotype

C26.4

MN 8981-I*; Fig. 9B

Paratype

C26.5

MN 8982-I

Paratype

C26.15

MN 8983-I*; Figs. 9C-E

Paratype

C26.51

MN 8984-I*

Paratype

C26.53

MN 8985-I*

Paratype

C26.77

MN 8986-I*; Figs. 9A, 10H

Paratype

C26.78

MN 8987-I*; Figs. 8, 10L

Paratype

* = measured specimen (Tables 23).