See discussions, stats, and author profiles for this publication at: https://www.researchgate.

net/publication/241075573

Formation processes of shell concentrations in the Lower Cretaceous

estuarine sediments of the Okurodani Formation, Tetori Group, Gifu
Prefecture, central Japan

Article  in  Palaeogeography Palaeoclimatology Palaeoecology · August 2011

DOI: 10.1016/j.palaeo.2011.05.053

CITATIONS READS

5 290

3 authors:

Shota Yamashita Hiroshige Matsuoka

JX Nippon Oil & Gas Exploration Corpolation Kyoto University
12 PUBLICATIONS   236 CITATIONS    62 PUBLICATIONS   737 CITATIONS   

SEE PROFILE SEE PROFILE

Hajime Naruse
Kyoto University
92 PUBLICATIONS   1,314 CITATIONS   

SEE PROFILE

Some of the authors of this publication are also working on these related projects:

Inverse modeling of tsunami deposits using deep learning neural network View project

Inverse analysis of turbidites View project

All content following this page was uploaded by Shota Yamashita on 28 January 2019.

The user has requested enhancement of the downloaded file.

 Palaeogeography, Palaeoclimatology, Palaeoecology 308 (2011) 476–491

Contents lists available at ScienceDirect

Palaeogeography, Palaeoclimatology, Palaeoecology

j o u r n a l h o m e p a g e : w w w. e l s ev i e r. c o m / l o c a t e / p a l a e o

Formation processes of shell concentrations in the Lower Cretaceous estuarine

sediments of the Okurodani Formation, Tetori Group, Gifu Prefecture, central Japan
Shota Yamashita a,⁎, Hiroshige Matsuoka b, Hajime Naruse a
a
Department of Earth Sciences, Graduate School of Science, Chiba University, Japan
b
Department of Earth and Planetary Science, Graduate School of Science, Kyoto University, Japan

a r t i c l e i n f o a b s t r a c t

Article history: The formation processes of shell concentrations in estuarine environments are complicated by the interplay
Received 1 December 2009 between the original biogenic and overprinting sedimentary processes. In order to better understand this
Received in revised form 18 May 2011 interplay, the formation processes of shell concentrations composed exclusively of monospecific Myrene
Accepted 30 May 2011
tetoriensis (“corbiculoid” bivalve) occurring in the Lower Cretaceous estuarine deposits of the Okurodani
Available online 6 June 2011
Formation of the Tetori Group in central Japan were considered within in the context of their mode of life. In
Keywords:
addition, the size–frequency distribution of shells was discussed in relation to the formation processes of shell
Cretaceous concentrations. As a result, the autochthonous occurrence of M. tetoriensis clarified a gregarious mode of life in
Tetori Group the muddy sediments of a central estuary. In contrast, the post-mortem exhumation, accumulation, and
Shell beds degree of transport of dead shells by hydrodynamic processes were reflected in various occurrence modes of
Taphonomy allochthonous shell concentrations. The size distribution patterns of these shell concentrations suggested that
Estuary the primary size composition of autochthonous living and dead assemblages could have been retained in the
absence of sufficient overprinting, even through reworking and transport. This was probably possible because
of the high production of dead shells as compared to the relative effect of sedimentary processes.
© 2011 Elsevier B.V. All rights reserved.

1. Introduction environments, these processes are relatively weak in embayed coastal

The present study attempted to reveal the formation processes of
molluscan shell concentrations in an estuarine environment by
analyzing their preservation and shell size distribution. Molluscan
shell concentrations have been the focus of several previous studies,
demonstrating their role as important indicators in paleoenviron-
ments (e.g., Kidwell et al., 1986; Kidwell, 1991; Fürsich and
Oschmann, 1993; Fürsich and Pandey, 1999, 2003). Molluscan shell
concentrations are a ubiquitous feature of many modern and ancient
coastal environments, including estuaries (Trewin and Welsh, 1976;
Davies et al., 1989; Meldahl and Cutler, 1992; Damborenea and Lanés,
2007) and open sea environments (Fürsich and Oschmann, 1993;
Fürsich and Pandey, 2003). Numerous studies have investigated the
taphonomic signatures of shell concentrations, e.g., species diversity,
disarticulation rates, and the degree of fragmentation and abrasion,
and have demonstrated that the taphonomic signatures of shell
concentrations are good indicators of sedimentary processes such as
waves and tidal and fluvial currents (Müller, 1951; Schäfer, 1972;
Fürsich and Oschmann, 1986; Kidwell, 1986; Fürsich et al., 1991;
Kidwell, 1991; Meldahl and Cutler, 1992; Radley and Barker, 2000;
Matsukawa and Nakada, 2003). However, compared to open-sea
⁎ Corresponding author.
E-mail address: yama-c@graduate.chiba-u.jp (S. Yamashita).
environments (Kidwell, 1986; Dalrymple et al., 1992). Alternatively,
the high productivity of biogenic hardparts originating from cohorts
of opportunistic species is expected to play a major role in the
formation of shell concentrations (Aigner et al., 1978; Kidwell, 1986).
However, an accurate reconstruction of the formation processes of
shell concentrations in coastal environments is relatively difficult due
to the interplay of biogenic and sedimentological factors (Kidwell,
1986). Since the original biogenic information, particularly of non-
sessile epifauna and shallow infauna, can easily be erased by
sedimentological reworking (Kidwell, 1986, 1991), further examples
of shell concentrations that are closely related to the ecology of
specimens are required from a variety of sedimentary environments.
To this end, the present study focuses on various occurrence
modes of shell concentrations composed exclusively of the brackish
neomiodontid (“corbiculoid”) bivalve, Myrene tetoriensis (Kobayashi
and Suzuki, 1937; Hayami, 1958; Matsukawa and Ido, 1993), in
estuarine sediments of the Lower Cretaceous Okurodani Formation of
the Tetori Group in central Japan (Fig. 1). First, autochthonous
assemblages of M. tetoriensis were identified and their mode of life
was reconstructed. Second, the taphonomic signatures of M. tetor-
iensis shells occurring in the same horizon were observed. Since the
physical transport of shells plays a relatively minor role in the calm
depositional environments of embayed coasts, unlike in other
depositional environments, fragmentation and abrasion rates may
not be useful as indicators of shell transport (Fürsich and Oschmann,

0031-0182/$ – see front matter © 2011 Elsevier B.V. All rights reserved.
doi:10.1016/j.palaeo.2011.05.053
 S. Yamashita et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 308 (2011) 476–491 477

Fig. 1. A. Distribution of the Tetori Group in central Japan (modified after Maeda, 1961). B. Locality of the study area. Lithologies of transects 1 through 5 are shown in Fig. 3.

1986; Fürsich et al., 1991). We therefore focused on the size–
frequency distribution of shells, because this distribution is expected
to contain information about the paleoecology of M .tetoriensis and
transport processes of their shells (Hallam, 1967; Surlyk, 1972, 1974;
Alexander, 1977; Cadée, 1983; Shimoyama, 1985; Cummins et al.,
1986).
As a result, the ecology of M. tetoriensis, i.e., the gregarious
behavior of this species in the muddy sediments of a central estuary, is
clarified herein. In addition, it is demonstrated that the primary size
composition of living and death assemblages can be retained despite
reworking and transport because of a very high production of bio-
genic hardparts. The present study represents a remarkable case study
for clarifying the relationship between the original biogenic and
sedimentological processes leading to the formation of shell
concentrations.
1996; Manabe and Evans, 1996; Yabumoto, 1997; Hirayama et al.,
2000) have been found in the Tetori Group in the study area. The
Tetori Group is composed of three subgroups, namely, the Kuzuryu,
Itoshiro, and Akaiwa subgroups. The Itoshiro Subgroup of the study
area has been subdivided into three formations, namely, the
Otaniyama, Okurodani, and Amagodani Formations (Maeda, 1961)
(Fig. 2). The Otaniyama Formation is composed of an arkosic
sandstone bed and is conformably overlain by the Okurodani
Formation. The Okurodani Formation comprises alternating beds of
mudstone and sandstone. The bottom part of this formation is
composed of a black mudstone bed. This formation is conformably
overlain by the Amagodani Formation, which is predominantly
composed of coarse-grained arkosic sandstone and conglomerate
beds. The Itoshiro Subgroup of the Shokawa District has been

2. Geological setting

The shell concentrations described in the present paper are from

the Lower Cretaceous Tetori Group in the Shokawa District of central
Japan (Fig. 1). The Tetori Group is composed of Middle Jurassic to
Lower Cretaceous sedimentary rocks and is distributed in the
Hokuriku and Hida districts of central Japan (Maeda, 1961) (Fig. 1).
The Tetori Group is composed of marine and non-marine deposits.
The Tetori Basin is interpreted as having initially developed in the
Middle Jurassic by subsidence and incipient sedimentation (Isozaki
and Maruyama, 1991; Maruyama et al., 1997; Matsukawa et al., 1997,
1998, 2006). In the Early Cretaceous, the Tetori Basin expanded and
shifted northward due to the duplication of Jurassic to Early
Cretaceous accretionary complexes and an accompanying strike-slip
movement (Matsukawa et al., 1997, 1998, 2006).
The Tetori Group occurs in three separate regions, namely, the
Kuzuryugawa, Mt. Hakusan, and Jinzu regions (Fig. 1). The Tetori
Group in the Shokawa District represents the easternmost distribu-
tion of the Mt. Hakusan region (Fig. 1) (Maeda, 1961; Matsukawa
et al., 2003). The study area comprises the Kobudani and Okurodani
areas, which are located in the southwestern part of the Shokawa
District (Fig. 1). Various macrofossils, including plants and molluscs
(Yokoyama, 1889; Kobayashi and Suzuki, 1937; Maeda, 1952, 1954,
1961), as well as vertebrate fossils, such as fishes, reptiles, turtles, and Fig. 2. Stratigraphic table of the Tetori Group in the Shokawa District, based on
dinosaurs (Gifu-ken Dinosaur Research Committee, 1993; Hirayama, Kusuhashi et al. (2006).
478 S. Yamashita et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 308 (2011) 476–491

placed in the Lower Cretaceous based on U–Pb radiometric dating
(Kusuhashi et al., 2006) (Fig. 2).
In the present study, three transects in the Kobudani area
(Transects 1 through 3) and two transects in the Okurodani area
(Transects 4 and 5) were investigated (Figs. 1 and 3). According to
the stratigraphic classifications of The Gifu-ken Dinosaur Fossil
Excavation Party (1999, 2000), the uppermost part of the Otaniyama
Formation and the Okurodani and Amagodani formations are
distributed in these areas (Figs. 2 and 3). The strata dip 20 to 30°
toward the northwest and compose homoclinal structures.
3. Sedimentary environments
Based on lithology, sedimentary structures, and fossils, four sedi-
mentary facies (Facies A through D) were identified in the investigated
succession (Figs. 3 and 4), and the depositional environment of the
middle part of the Tetori Group in the study area was reconstructed.
3.1. Facies A: shallow-marine deposits
Description: Facies A comprises alternating beds of fine- to coarse-
grained sandstone with common intercalation of fine to coarse
conglomerates (Fig. 4). Muddy sediments, organic matter, and trace
fossils are absent. The sandstone beds are 0.2 to 20 m thick. The
sandstone beds are very well sorted and do not exhibit grading
(Fig. 4). Parallel and small-scale trough cross-laminations are
observed in the sandstone beds (Fig. 4). The conglomerate beds are
0.05 to 1 m thick. In addition, the conglomerate beds are well sorted
and exhibit faint grading (Fig. 4). The conglomerate beds persist
laterally and exhibit a tabular or sheet-like geometry. The conglom-
erate and sandstone beds have a flat, non-erosional contact (Fig. 4).
The bivalves Myrene tetoriensis, Megasphaerioides okurodaniensis
(Komatsu et al., 2003), and Ostrea sp. occur. Belemnites have also
been reported to exist within this facies (Kumon et al., 1994).
Facies A occurs in the lowermost part of the investigated section,
corresponding to the uppermost part of the Amagodani Formation
(Fig. 3). The total thickness of this succession is more than 100 m, and
is laterally continuous. This facies is directly overlain by laminated
siltstones (Facies B) and occasionally by sandy mudstone beds (Facies
C) (Fig. 3).
Interpretation: Facies A was probably deposited in a nearshore
marine environment, such as an upper shoreface. The horizontal
persistence of tabular conglomerate beds indicates deposition by
processes that acted uniformly over a broad area, such as waves or
coastal currents (Bourgeois and Leithold, 1984). These beds can be
interpreted as representing fluvially supplied gravels that were
reworked and widely distributed by long-period swells (Kumar and
Sanders, 1976; Clifton, 1973, 1981; Bourgeois and Leithold, 1984).
Trough cross-laminated beds were probably formed by wave or
current ripple migration (Reineck and Singh, 1980; Bourgeois, 1980).
The occurrence of Ostrea sp. and belemnites indicates marine or
brackish-water conditions (Stenzel, 1971).

Fig. 3. Lithology of the study area (right: Kobudani area, left: Okurodani area). Sections of transects 2 and 3 are correlated by a tuff bed, whereas other sections are correlated
geometrically. A through D refer to sedimentary facies discussed in the text. The horizons of detailed sections (Sections 1 though 6, see Fig. 7) are indicated by asterisks.
 S. Yamashita et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 308 (2011) 476–491
Fig. 4. Sedimentary facies table in the study area.
3.2. Facies B: central estuarine deposits
Description: Facies B consists primarily of well-sorted, laminated
siltstones of 5 to 20 m in thickness (Facies B-1), accompanied by thin
(5 to 50 cm thick) sandstone beds (Facies B-2) (Fig. 4). Bioturbation is
very weak in Facies B-1. Lenticular sandstone laminae (0.1 to 0.2 cm
thick, 5 to 10 cm wide) commonly occur in Facies B-1 (Fig. 4). Facies
B-2 is characterized by very fine- to fine-grained 5- to 50-cm-thick
sandstone beds. Erosional surfaces and normal grading is common,
and flaser or wavy bedding (Reineck and Wunderlich, 1968) with
combined-flow ripple lamination is occasionally observed (Fig. 4). The
upper boundary is in gradual contact with Facies B-1.
Myrene tetoriensis shell concentrations are very commonly inter-
layered in Facies B (at intervals of 30 to 100 cm). The brackish to
freshwater bivalves Megasphaerioides okurodaniensis and Ostrea sp.,
and the small Melanoides sp. gastropods also co-occur.
Facies B occurs in the lowermost part of the Okurodani Formation.
Lateral thickness variations of this facies are typical of the Kobudani
area where this facies thins distinctly toward the southeast from 10 to
30 m to less than 10 m (Fig. 3). The facies directly overlies Facies A and
is overlain by sandy mudstone beds (Facies C), although the boundary
is not exposed (Fig. 3).
Interpretation: Facies B is interpreted as central estuarine deposits.
The flaser or wavy beddings with combined-flow ripples in Facies B-2
were likely formed in tidal-influenced environments (Reineck, 1967;
Reineck and Wunderlich, 1968; Terwindt, 1971; Terwindt and
Breusers, 1972). The thick, laminated mudstone beds (Facies B-1)
probably indicate calm conditions along an embayed coast (Reineck
and Wunderlich, 1968; Reineck and Singh, 1980; Dalrymple et al.,
1992; Reading and Collinson, 1996). The weakness of bioturbation
may suggest a high-stress environment owing to fluctuations in
salinity, temperature, or oxygen levels, along with turbidity, erosion/
479
deposition, and mobility of bottom surface sediments (Droser and
Bottjer, 1986, 1989; Gingras et al., 2002; Dashtgard and Gingras, 2005;
Pearson and Gingras, 2006; Dalrymple and Choi, 2007). The thin
sandstone beds showing normal grading and erosional surfaces
(Facies B-2) could have been intermittently deposited by hydrody-
namic processes, such as waves, tidal and fluvial currents, or storm
flows (Reineck and Singh, 1980; Dalrymple et al., 1992; Einsele et al.,
1996; Myrow and Southard, 1996).
3.3. Facies C: fluvial overbank deposits
Description: Facies C consists primarily of intensely bioturbated,
poorly sorted, sandy mudstone beds. Facies C ranges in thickness from
2 to 20 m (Fig. 4) and contains a large amount of organic matter, such
as plant debris, wood fragments, and rootlets are also observed. Fine-
to medium-grained sandstone beds that are 0.5 to 2 m thick are
commonly interlayered (Fig. 4). Some of the sandstone beds exhibit a
weak fining-upward trend (Fig. 4). The freshwater bivalve Unio
ogamigoensis and the gastropod Viviparus sp. occur in Facies C.
Facies C occurs primarily in the middle to upper part of the
Okurodani Formation and is often overlain by medium- to very
coarse-grained sandstone beds (Facies D) with sharp erosional
contact (Fig. 4). This facies occurs above Facies B in the middle to
upper part of the Okurodani Formation. Facies B and C exhibit a
nesting pattern at Transect 1 in the Kobudani area (Fig. 3). It is
common for the lateral thickness of Facies C to vary (Fig. 3).
Interpretation: Facies C is interpreted as a fluvial overbank deposit.
The occurrence of in situ rootlets along with the abundance of plant
debris suggests subaerial vegetation (Bridge, 2003). The fine to
medium sandstone beds intercalated into mudstone beds may have
been deposited by intermittent flooding events (Hughes and Lewin,
1982; Bridge, 2003).
480 S. Yamashita et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 308 (2011) 476–491

Fig. 5. Detailed lithology of Sections 1 through 6. The positions of the sections are shown in Fig. 3. The horizons indicated by arrows denote the shell concentrations.
 S. Yamashita et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 308 (2011) 476–491 481

Table 1
Details of shell concentrations of Myrene tetoriensis.

locality name Lithofacies Type Approximate size composition Co-occurrence

Loc. 1 KB1-01 A 2 Uni-modal

KB1-02 A 4 (fragment)
KB1-03 A 4 (fragment)

Loc.2 KB2-01 B-2 2 Uni-modal

KB2-02 B-1 3-1 Bi-modal
KB2-03 B-1 3-1 Uni-modal
KB2-04 B-1 3-1 Bi-modal
KB2-05 B-1 3-1 Uni-modal (juveniles)
KB2-06 B-1 3-1 Uni-modal
KB2-07 B-1 3-1 Uni-modal
KB2-08 B-1 3-1 Bi-modal
KB2-09 B-1 3-1 Bi-modal
KB2-10 B-1 3-1 Uni-modal (juveniles)
KB2-11 B-2 2 Uni-modal
KB2-12 B-2 2 Uni-modal
KB2-13 B-2 3-2 Bi-modal

Loc.3 KB3-01 B-2 3-2 Bi-modal Melanoides sp.

KB3-02 B-1 3-1 Uni-modal (juveniles) Melanoides sp.
KB3-03 B-1 3-1 Uni-modal (juveniles) Melanoides sp.
KB3-04 B-1 3-1 Bi-modal Melanoides sp.
KB3-05 B-1 3-1 Bi-modal Melanoides sp.
KB3-06 B-1 3-1 Uni-modal (juveniles) Melanoides sp.
KB3-07 B-1 3-1 Bi-modal Melanoides sp.
KB3-08 B-1 3-1 Bi-modal Melanoides sp.

Loc.4 KB4-104 A 4 (fragment)

KB4-103 A 4 (fragment)
KB4-102 A 4 (fragment)
KB4-101 A 4 (fragment)
KB4-01 B-1 3-1 Uni-modal
KB4-02 B-1 3-1 Uni-modal
KB4-03 B-1 3-1 Uni-modal
KB4-04 B-2 2 Uni-modal
KB4-05 B-2 2 Uni-modal
KB4-06 B-2 2 Uni-modal
KB4-07 B-2 2 Uni-modal
KB4-08 B-2 3-2 Uni-modal
KB4-09 B-1 1 Uni-modal
KB4-10 B-1 1 Uni-modal
KB4-11 B-2 2 Uni-modal Megasphaerioides
okurodaniensis

Loc.5 KB5-01 A 4 (fragment)

KB5-02 A 2 Uni-modal
KB5-03 A 4 (fragment)

Loc.6 OK1-01 A 4 (fragment)

OK1-02 A 4 (fragment)
OK1-03 A 4 (fragment)
OK1-04 A 4 (fragment)

3.4. Facies D: fluvial channel deposits
Description: Facies D comprises poorly sorted, medium to very
coarse sandstone beds and granule to pebble conglomerate beds that
range from 1 to 20 m in thickness. The sandstone and conglomerate
beds exhibit a distinct fining-upward trend (Fig. 4). Most of the beds
have sharp, convex erosional bottom surfaces (Fig. 4). Mud clasts are
commonly found near the erosional surfaces. Muddy sediments and
organic matter are rare. The gastropod Viviparus sp. occurs only
occasionally.
Facies D occurs in the middle to upper part of the investigated
succession, which is correlated to the lower part of the Amagodani
Formation. In most cases, Facies D is accompanied by Facies C
(overbank deposits) and covers Facies C with erosional contact
(Fig. 4). The thickness of Facies D increases from 4 to 5 m in the
lowermost part to approximately 20 m in the middle part of the
Otaniyama Formation (Fig. 3).
Interpretation: Sharp, convex erosional surfaces and a fining-
upward trend are typical of channel-fills (Fielding, 1984, 1986). The
mud clasts and pebbles associated with the erosion surfaces are
probably related to channel migration (Lucchi, 1995). The coarse
sediments and low mud contents suggest a braided-river depositional
environment (McPherson et al., 1987).
3.5. Depositional environment
The Otaniyama, Okurodani, and Amagodani formations examined
in the present study area are interpreted as an incised-valley fill
system. The thick package of central estuarine deposits (Facies B)
overlying the nearshore marine deposits (Facies A) suggests marine
transgression. The braided channel (Facies D) and overbank deposits
(Facies C) overlaying the central estuarine deposits can be interpreted
as being of fluvial and bay-head delta origin (McPherson et al., 1987).
482 S. Yamashita et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 308 (2011) 476–491

Fig. 6. Shell concentration KB4-10 (Type 1). A. Photograph of the lower bedding plane. Articulated specimens are preserved with their commissural planes almost perpendicular to
the bedding plane with their umbos downward (arrows). (B) Photograph and (C) sketch of the cross-section. The horizon of KB4-11 is also shown in the uppermost part of the cross-
section. A large Megasphaerioides okurodaniensis specimen is observed near the top.

Depositional sequences are defined based on facies stacking. The
Otaniyama, Okurodani, and Amagodani formations examined in the
present study area are considered to reflect a regressive transition
from marine to fluvial environments. Two smaller-order transgres-
sive–regressive cycles occur in the succession. The thick, shallow
marine deposits (Facies A) suggest highstand deposition (Zaitlin et al.,
1994). The first sequence boundary is estimated to be between Facies
A and Facies B or C in the uppermost part of the Otaniyama Formation
(Fig. 3). The succession, which is composed primarily of central
estuarine deposits (Facies B) that directly overlie the boundary,
corresponds to Lowstand Systems Tract (LST) and Transgressive
Systems Tract (TST) (Zaitlin et al., 1994). The second sequence
boundary is estimated in the lower to upper part of the Okurodani
Formation, dividing the central estuarine succession and fluvial
successions above it (Fig. 3). The upward thickening succession
composed of fluvial overbank (Facies C) and channel-fill deposits
(Facies D) above the second sequence boundary suggests bayhead
delta top and fluvial deposition, corresponding to LST and TST (Zaitlin
et al., 1994). Based on these interpretations, the Otaniyama,
Okurodani, and Amagodani formations in the study area are regarded
as a regressive transition from a nearshore marine to a fluvial
environment via an estuarine setting through two transgressive–
regressive cycles. This is conformable to the interpretations of
Umezawa (1997) and The Gifu-ken Dinosaur Fossil Excavation Party
(1999, 2000).
4. Taphonomic analysis
4.1. General remarks on Myrene tetoriensis
The focus of the present study, namely, Myrene tetoriensis, is a
“corbiculoid” bivalve. The “Corbiculoids” are a polyphyletic group
composed of three families: Neomiodontidae, Corbiculidae, and
Pisidiidae (Casy, 1955). In other words, it is a term used for bivalves
that advanced into brackish-water environments during the mid-
Mesozoic. Various neomiodontid and corbiculid bivalve species occur
in non-marine Lower Cretaceous deposits in Japan (Matsukawa and
Nakada, 2003). Whereas the Neomiodontidae became extinct during
the Late Cretaceous, contemporary species of the Corbiculidae were
distributed in freshwater and brackish water environments, such as
estuaries (Matsukawa and Nakada, 2003).
 S. Yamashita et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 308 (2011) 476–491
Fig. 7. Shell concentration KB4-11 (Type 2). (A) Photograph and (B) sketch of the cross-section.
Myrene tetoriensis was first described by Kobayashi and Suzuki
(1937), who designated this species Corbicula tetoriensis in the family
Corbiculidae, whereas Hayami (1958) redesignated this species as M.
tetoriensis in the Neomiodontidae. Numerous specimens exhibiting
various growth stages occur in the Tetori Group. During the growth
stage, the height and convexity of Myrene tetoriensis increase and the
specimens become subcircular- or subelliptical-shaped, and truncated
obliquely behind the beak (Kobayashi and Suzuki, 1937). Adult
specimens exhibit a triangular form (Kobayashi and Suzuki, 1937).
Their habitat is interpreted as being shallow infaunal, because they
exhibit a very shallow pallial sinus (Kobayashi and Suzuki, 1937;
Matsukawa and Ido, 1993).
Matsukawa and Ido (1993) defined several fossil assemblages,
including a monospecific M. tetoriensis assemblage from the Tetori
Group of this area. Matsukawa and Nakada (1999) inferred the
salinity of the habitats of these assemblages by comparisons with
related taxa from modern environments. They reported that the
habitat preference of M. tetoriensis ranged from freshwater to brackish
water, and that the monospecific occurrence of M. tetoriensis may
have corresponded to a salinity of approximately 5‰.
4.2. Occurrence of shell concentrations of Myrene tetoriensis
A total of 30 beds of M. tetoriensis shell concentrations were
investigated along six transects (Fig. 5, Table 1). Based on the degree
of articulation and fragmentation, the occurrence modes of the
Myrene tetoriensis shell concentrations were classified into four types.
Then, their taphonomic features, i.e., bed geometry and the distribu-
tion of shell orientation, were described (Figs. 5 through 10). The
density of occurrence was defined as half the total number of shells
(regarding two shells as one individual) per unit area (1 m 2). In order
to avoid over-counting, fragmented specimens lacking umbones were
not counted.
4.2.1. Type 1: Concentration of articulated shells
Almost all of the shells are articulated (percentage of articulation:
N90%) (Figs. 6 and 11). The fine details of the surface ornaments of the
shells are well preserved, and more than half of the specimens occur
483
with their valves closed (Fig. 6). Encrustation and boring are not
observed. The commissure planes are perpendicular to the bedding
plane or recline at an angle of approximately 30° to the bedding plane
with their umbones directed downward (Fig. 6). Small clusters
composed of five to six articulated specimens are frequently observed
(Fig. 6). The density of shells is less than 1000 specimens/1 m 2 (Fig. 6).
This type of shell concentration occurs at only two locations within
Facies B-1 in the upper part of Section 4 (KB4-09, 10) (Fig. 5). Shelly
horizons extend for 0.5 to 1 m laterally, and are directly overlain by a
thin sandstone bed (Facies B-2) (Fig. 6). At KB4-10, the contact
between the shelly horizon and the sandstone bed exhibits a sharp
erosional surface, whereas it is disturbed by downward directed
burrows 2–3 mm in diameter, at KB4-09 (Fig. 6).
4.2.2. Type 2: Concentration of partly articulated shells
This type of shell concentration includes both articulated and
disarticulated shells (Figs. 7 and 11). The shell-surface sculptures are
nearly perfect. Encrustation and boring are not observed. The
percentage of articulation ranges from 5 to 50%. Most of the
articulated specimens are gaping at 120°, and some of the specimens
are nested (Fig. 7). Muddy sediment fills the drafts of shells (Fig. 7).
Shells are randomly oriented both in the plan and cross-sectional
views (Fig. 7). This type of concentration exhibits a matrix-supported
fabric in the mudstone beds of Facies B-2 and occasionally in
sandstone and conglomerate beds of Facies A (Fig. 5). The shelly
horizons extend for 0.5–2 m laterally and are 3–5 cm thick. Articu-
lated shells are relatively rare in Facies A (5–10% on average).
4.2.3. Type 3: Concentration of disarticulated shells
Most of the shells are disarticulated but complete (Figs. 8, 9, and
11). The molds of these shells exhibit fine hinge details, and
encrustation and boring are not observed (Fig. 8). Shell concentra-
tions of this type are subdivided into two subtypes (Types 3–1 and 3–
2) corresponding to their geometry (Figs. 8 and 9). Type 3–1 forms
pavements extending laterally for 1 to 5 m in the mudstone beds of
Facies B-1 (Fig. 8). Shells are very densely packed along the bedding
plane and are arranged in a convex-up position (Fig. 8). In plan
orientation, the long axes of the shells tend to point in a NS–SE
484 S. Yamashita et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 308 (2011) 476–491

Fig. 8. (A–C) Shell concentrations KB2-07, (D, E) KB3-06, and (F, G) KB3-07 (Type 3–1). A, Photograph of the bedding plan of KB2-07 with the inner molds of the shells. B, C. Close-up
photographs of shells. Note that the fine details of the hinges are preserved. D. Photograph of the bedding plan of KB3-06 with the inner molds of the shells. E. Cross-section of a large shell
surrounded by a dense concentration of small shells. F. Photograph of the bedding plan of KB3-07. G. Cross-section. Small shells of 1 to 3 mm in length are horizontally aligned (arrow).

direction (Fig. 12). The density of the occurrences is quite high (KB4-09, KB4-10), one Type 2 bed (KB4-11), and four Type 3 beds
(approximately 2000–50,000/1 m 2). Type 3–2 is characterized by a (KB2-05, KB2-07, KB3-06, and KB3-07) (Fig. 13). Type 4 shells were
matrix-supported fabric in the sandstone beds of Facies B-2 (Fig. 9). not measured due to their fragmentation. Descriptive statistic values
The shelly beds extend laterally for 1 to 2 m (Fig. 9). The cross- were calculated as follows:
sectional orientation of shells is almost random. Shell size commonly
decreases upward (Fig. 9). ∑pd
Mean shell size ðmÞ = ðmmÞ
100
4.2.4. Type 4: Disarticulated-shell concentration including shell sffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffi
2
fragments ∑pðd−mÞ
Standard deviation ðσ Þ = ðmmÞ
Most of the shells are disarticulated and are often fragmented 100
(Fig. 10). The edges of the shell fragments are sharp. Encrustation and
boring are not observed. This concentration type exhibits a matrix- ∑pðd−mÞ3
supported fabric in the sandstone and conglomerate beds of Facies A Skewness ðSk Þ = ðdimensionlessÞ;
100σ 3
(Fig. 10). Shelly horizons extend 1 to 2 m laterally. Shells are
randomly arranged in cross-sectional view (Fig. 10). where d is a representative value of each shell-size class (in
increments of 2.0 mm) and p is a fraction (%).
4.2.5. Size–frequency distributions of Myrene tetoriensis shell The size–frequency distribution of Type 1 shell concentrations has
concentrations a symmetric, bell-shaped form with a single distinct peak (Fig. 13).
The sizes of the Myrene tetoriensis shells occurring in the shell The shell size included in this type is 10 to 20 mm on average (Fig. 13).
concentrations were measured at seven horizons: two Type 1 beds The standard deviation (σ) is approximately 1.9 (Fig. 13), and thus the
 S. Yamashita et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 308 (2011) 476–491 485

Fig. 9. Cross-section through the shell concentration KB3-01 (Type 3–2). A. Photograph and B. sketch. Note the sharp erosional surface of the sandstone bed. Shells are graded, and
small shells occur in the uppermost part of the bed.

degree of size dispersion is quite low. The skewness value is highly
variable (from approximately −0.4 to 1.0) (Fig. 13).
Although the size–frequency distribution of Type 2 also has a
distinct, sharp peak at approximately 15 mm, and the shape of the
distribution curve is negatively skewed (approximately −0.5), the
standard deviation is low (approximately 2.1) (Fig. 13).
Type 3–1 shell concentrations exhibit more complicated features.
Based on the approximate sizes of the component shells, three types
of size-distribution patterns are recognized: A) bimodal distribution,
B) unimodal distribution composed of larger shells (N5 mm), and C)
unimodal distribution composed of smaller shells (b4 mm). Among
the 22 beds with Type 3 shell concentration, 10 beds exhibit Pattern A,

Fig. 10. Cross-section of shell concentration OK1-01 (Type 4). A. Photograph and B. sketch. Shells and shell fragments (arrow) show matrix-support fabric in the sandstone bed.
486 S. Yamashita et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 308 (2011) 476–491

Fig. 11. Degree of articulation of shell concentrations of Types 1 through 3.

seven beds exhibit Pattern B, and five beds exhibit Pattern C (Table 1).
The size–frequency distribution of Pattern A has a strong positive-
skewness (approximately 0.7) because smaller specimens are
dominant in the distribution (Fig. 13). The largest shell is approx-
imately 30 mm and the degree of dispersion is quite high (8.2)
(Fig. 13). The size–frequency distribution of Pattern B is moderately
positively skewed (Sk N 0.9) continuous histogram lacking small shells
(b4 mm) (Fig. 13). One well-defined peak appears at approximately
10 mm (Fig. 13) and the degree of dispersion is also high (σ ~ 4)
(Fig. 13). The same size patterns of component shells (described
above) are found for Type 3–2. One horizon of Pattern A and two
horizons of Pattern B are discriminated from three horizons of Type 3–
2 occurrences.

5. Discussion

5.1. Autochthonous assemblage of Myrene tetoriensis and implication for

their ecology

Type 1 occurrence revealed that Myrene tetoriensis inhabited the

Fig. 12. Plane-view orientation patterns of Type 3–1 shell concentrations (KB2-05, KB2-07).
muddy sediments of a central estuary. The rather high degree of
articulation (N90%) suggests the preservation of autochthonous
specimens (Fig. 14). The preferred arrangement of their commissural
plane perpendicular to the bedding may be explained as more or less
reflecting their life position, because this orientation is physically
unstable as a result of hydrodynamic transport (Allen, 1990; Kondo,
1997). This type of concentration can thus be interpreted as an
autochthonous assemblage representing the mode of life of this
 S. Yamashita et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 308 (2011) 476–491 487

Fig. 13. Size–frequency distribution of Myrene tetoriensis shells in concentrations of Types 1 through 3. σ: standard deviation, Sk: skewness.
488 S. Yamashita et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 308 (2011) 476–491
Fig. 14. Paths leading to Myrene tetoriensis shell concentrations with the corresponding size–frequency distribution of shells.
species. A cross-sectional view of this type of concentration reveals
that the preservation of autochthonous specimens of M. tetoriensis
resulted from the deposition of sand, which may have protected these
specimens from post-mortem reworking and transport (Fig. 14).
The size-distribution pattern of the shells suggests that this type
of concentration is a census assemblage rather than an autochtho-
nous death assemblage that originated from the in-situ accumulation
of dead shells from several generations. The size–frequency
distributions of living and death assemblages exhibit different
patterns. Life assemblages are generally characterized by several
well-defined peaks representing each age class and exhibit a bi- or
polymodal distribution pattern arising from seasonal recruitment
(e.g., Shimoyama, 1985; Noe-Nygaard et al., 1987). In contrast, death
assemblages exhibit strongly positively-skewed and continuous
size–frequency distributions, with the abundance of juveniles
being due to the higher rate of juvenile mortality (Surlyk, 1972,
1974; Alexander, 1977). The Type 1 size-distribution, which exhibits
a unimodal histogram composed of similar-sized shells, clearly
differs from the death assemblage. Therefore, the Type 1 size-
distribution can be interpreted as a single generation having
composed a living assemblage, with the absence of other generations
being due to several factors. One possible factor is the short duration
of the specimens. Living assemblages composed exclusively of a
single generation can arise when most of the specimens of the last
generation die soon after subsequent recruitments (Shimoyama,
1985). Another explanation is related to the dispersal of juveniles.
This behavior is commonly observed in contemporary bivalves,
including corbiculoids, as a response to environmental deterioration
(Bayne, 1964; Sigurdsson et al., 1976; Beukema and Vlas, 1989;
Shibuya and Kato, 1989; Takada et al., 2000). Estuarine environ-
ments, in which salinity, temperature, or oxygen levels fluctuate
frequently, support the proposition that M. tetoriensis exhibited this
behavior.
In summary, M. tetoriensis exhibited gregarious behavior in a
central estuary, and their living assemblages were commonly
composed of single generations. The high input rate of biogenic
hardparts, due primarily to their gregarious mode of life, played a
major role in the formation of shell concentrations of this type. Rapid
deposition of a thick layer of sand may have been responsible for the
in-situ burial of living assemblages of the shallow-infaunal bivalves
and would explain the absence of escape behavior (Kranz, 1974;
Kondo, 1997). However, the possibility that in-situ mass mortality was
caused by some other factor, such as environmental fluctuation or
toxic dinoflagellate blooms, before the sand sealed the assemblage
cannot be overlooked (e.g., Noe-Nygaard et al., 1987).
5.2. Formation processes of allochthonous shell concentrations of
Myrene tetoriensis
While Type 1 concentrations are interpreted as autochthonous
assemblages, Type 2 through 4 concentrations show evidence of
reworking and transport.
The Type 2 concentration of partially articulated shells shows
obvious signs of reworking and transport (Fig. 14). The shelly
sandstone beds with an erosional base and a faint fining-upward
pattern (Facies B-2) indicate the transport of shells by hydrodynamic
processes (Fig. 14). Articulated shells filled with muddy sediment
signify the reworking of autochthonous specimens in muddy
sediments and their short-term transport (Kidwell, 1991; Matsukawa
et al., 1993). The occurrence of Type 2 shell concentrations in marine
 S. Yamashita et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 308 (2011) 476–491
sediments (Facies A) suggests that shells were commonly transported
from their habitats.
Both Type 3–1 and 3–2 shell concentrations show evidence of
shell transport and accumulation via post-mortem exhumation
(Fig. 14). The predominance of disarticulated shells of Type 3–1
concentration indicates the transport of shells (Matsukawa et al.,
1993). Compared to autochthonous assemblages (Type 1), the higher
shell densities of the Type 3–1 concentrations corroborate their post-
mortem accumulation. The predominance of the convex-up orien-
tation and the preferred plan-view orientation of shells suggest their
rearrangement by hydrodynamic processes (Fig. 14) (Brenchley and
Newall, 1969; Allen, 1990). Shelly sandstone beds with an erosional
lower surface (Facies B-2) of Type 3–2 suggest hydrodynamic
reworking and transport of shells (Fig. 14). The dense packing
consisting almost completely of disarticulated shells, which differ-
entiate this concentration type from the Type 2 shell concentration,
may be explained as being the result of reworking of the Type 3–1
concentration (Fig. 14).
The disarticulation and fragmentation of shells in Type 4 shell
concentrations implies transport and physical breakage of shells by
hydrodynamic processes (Fürsich and Oschmann, 1993) (Fig. 18). The
occurrence of this type in only the marine sediments (Facies A)
indicates transport from their original habitat (Fürsich and Oschmann,
1993; Fürsich and Pandey, 1999).
In summary, transport by hydrodynamic processes is interpreted
as the final formation process of allochthonous M. tetoriensis shell
concentrations (Types 2 through 4), and post-mortem exhumation,
accumulation, and the degree of transport of dead shells are reflected
in the various occurrence modes. In addition to these processes, the
high production of dead shells due to the gregarious behavior of M.
tetoriensis probably contributed to the origin of these types of shell
concentrations.
5.3. Taphonomic processes and size–frequency distribution of Myrene
tetoriensis shell concentrations
The size–frequency distribution of living and death assemblages of
shelled animals (molluscs and brachiopods) is related to both their
ecology and taphonomic biases, i.e., size-selective removal by currents
or predation by birds or crabs (Cadée, 1983; Shimoyama, 1985). In the
present study, the size distribution of typical residual assemblages, i.e. a
moderately positive skewness and a continuous histogram without
juveniles (Cadée, 1983, Matsukawa et al., 1996), is reflected in Pattern B
of the Type 3 concentrations (Fig. 13). The taphonomic evidence
regarding the Type 3 concentrations, which indicates the hydrodynamic
transport of shells, suggests that Pattern B of the Type 3 was formed by
the selective removal of small shells from autochthonous death
assemblages by currents.
However, the size–frequency distributions of numerous other shell
concentrations of the present study do not agree with the taphonomic
evidence described in previous studies. First, as described above, the
unimodal size–frequency distribution of autochthonous assemblages
(Type 1) is typically considered to be the result of size-sorting during
hydrodynamic transport (e.g., Zuschin et al., 2005). Thus, the well
defined unimodal size–frequency distribution of Type 2 shell
concentrations may be regarded as representing populations that
were originally unimodal living assemblages, rather than being the
result of size-sorting during transport. Conversely, the size–frequency
distribution of Pattern A in the Type 3 shell concentrations exhibits
characteristics typical of autochthonous death assemblages, which are
characterized by a continuous histogram with abundant juveniles,
despite the apparent evidence of transport (Fig. 14). This mismatch is
probably because of the imperfect overprinting of the original
biogenic information. The gregarious behavior of M. tetoriensis is
considered to have resulted in a very high production of dead shells,
which is believed to have played a major role in the formation of
489
various types of shell concentrations, particularly since the hydrody-
namic processes are relatively weak in a central estuarine environ-
ment (Kidwell, 1986; Dalrymple et al., 1992). These findings imply
that the primary size composition of autochthonous living and death
assemblages may have been retained without significant overprint-
ing, even throughout reworking and transport. Additional studies
should therefore be conducted in a variety of depositional environ-
ments in order to clarify the detailed characteristics of the over-
printing processes on the original size–frequency distribution through
sedimentary processes.
6. Conclusions
1) Autochthonous assemblages of the fossil “corbiculoid” bivalve,
Myrene tetoriensis occur in the Okurodani Formation of the Tetori
Group in central Japan. This occurrence indicates that M. tetoriensis
lived gregariously in the muddy sediments of a central estuary and
that their living assemblages were commonly composed of a single
generation.
2) The formation processes of M. tetoriensis shell concentrations were
clarified based on taphonomic analysis. Transport by currents is
considered to have been the final process of shell concentration
formation, and post-mortem exhumation, accumulation, and
degree of transport of dead shells were reflected in the various
modes of their occurrence. In any case, the high production of dead
shells due to the gregarious behavior of M. tetoriensis is likely to
have played an important role in the formation of shell
concentrations.
3) The size–frequency distribution of shells was discussed in relation
to the formation processes of shell concentrations. Compared to
the relative effect of sedimentary processes, the primary size
composition of autochthonous living and death assemblages can
be retained in the absence of sufficient overprinting, even
throughout reworking and transport, because of the high
production of dead shells.
Acknowledgements
The authors would like to thank Prof. Harutaka Sakai, Dr.
Haruyoshi Maeda, and Prof. Atsushi Yamaji of Kyoto University for
their advice and scientific discussions. Several members of the
biosphere group and members of the geology and mineralogy group
of Kyoto University provided help and motivation. Special thanks are
due to Mr. Shizuo Shimojima of the Shokawa branch office of
Takayama City and his family for providing everything we needed
during our fieldwork.
References
Aigner, T., Hagdorn, H., Mundlos, R., 1978. Biohermal, biostromal, and storm-generated
coquinas in the Upper Muschelkalk. Neues Jahrbuch für Geologie und Paläonto-
logie, Abhandlungen 157, 42–52.
Alexander, R.R., 1977. Growth, morphology and ecology of Paleozoic and Mesozoic
opportunistic species of brachiopods from Idaho–Utah. Journal of Paleontology 51,
1139–1149.
Allen, J.R.L., 1990. Transport -hydrodynamics: shells. In: Briggs, D.E.G., Crowther, P.R.
(Eds.), Palaeobiology: A Synthesis. Blackwell, Oxford, pp. 227–230.
Bayne, B.L., 1964. Primary and secondary settlement in Mytilus edulis L. (Mollusca).
Journal of Animal Ecology 33, 513–523.
Beukema, J.J., Vlas, J., 1989. Tidal-current transport of thread-drifting postlarval
juveniles of the bivalve Macoma balthica from the Wadden Sea to the North Sea.
Marine Ecology Progress Series 52, 193–200.
Bourgeois, J., 1980. A transgressive shelf sequence exhibiting hummocky stratification:
the Cape Sebastian Sandstone (Upper Cretaceous), southeastern Oregon. Journal of
Sedimentary Petrology 50, 681–702.
Bourgeois, J., Leithold, E.L., 1984. Wave-worked conglomerates — depositional
processes and criteria for recognition. In: Koster, E.H., Steel, R.J. (Eds.),
Sedimentology of Gravels and Conglomerates: Memoir of the Canadian Society of
Petroleum Geologists, 10, pp. 331–344.
490 S. Yamashita et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 308 (2011) 476–491
Brenchley, P.J., Newall, G., 1969. Flume experiments on the orientation and transport of
models and shell valves. Palaeogeography, Palaeoclimatology, Palaeoecology 7,
185–220.
Bridge, J.S., 2003. Rivers and Floodplains. Blackwell, London.
Cadée, G., 1983. Low juvenile mortality of fossil brachiopods, some comments. Internal
reports Nederlands Instituut voor Onderzoek der Zee, Texel, The Netherlands, p. 29.
Casy, R., 1955. The pelecypod family Corbiculidae in the Mesozoic of Europe and the
Near East. Journal of the Washington Academy of Science 45 (12), 366–372.
Clifton, H.E., 1973. Pebble segregation and bed lenticularity in wave-worked versus
alluvial gravel. Sedimentary Geology 20, 173–187.
Clifton, H.E., 1981. Progradational sequences in Miocene shoreline deposits, south-
eastern Caliente Range, California. Journal of Sedimentary Petrology 51, 165–184.
Cummins, H., Powell, E.N., Stanton, R.J., Staff, G., 1986. The size–frequency distribution
in palaeoecology: effects of taphonomic processes during formation of molluscan
death assemblages in Texas bays. Palaeontology 29, 495–518.
Dalrymple, R.W., Choi, K., 2007. Morphologic and facies trends through the fluvial-
marine transition in tide-dominated depositional systems: a schematic framework
for environmental and sequence-stratigraphic interpretation. Earth-Science Re-
views 81, 135–174.
Dalrymple, R.W., Boyd, R., Zaitline, B.A., 1992. Estuarine facies models: conceptual basis
and stratigraphic implications. Journal of Sedimentary Petrology 62, 1130–1146.
Damborenea, S.E., Lanés, S., 2007. Early Jurassic shell beds from marginal marine
environments in southern Mendoza, Argentina. Palaeogeography, Palaeoclimatol-
ogy, Palaeoecology 250, 68–88.
Dashtgard, S.E., Gingras, M.K., 2005. Facies architecture and ichnology of recent salt-
marsh deposits: Waterside Marsh, New Brunswick, Canada. Journal of Sedimentary
Research 75, 596–607.
Davies, D.J., Powell, E.N., Stanton Jr., R.J., 1989. Taphonomic signature as a function of
environmental processes: shells and shell beds in a hurricane-influenced inlet on
the Texas coast. Palaeogeography, Palaeoclimatology, Palaeoecology 72, 317–356.
Droser, M.L., Bottjer, D.J., 1986. A semiquantitative field classification of ichnofabric.
Journal of Sedimentary Petrology 56, 558–559.
Droser, M.L., Bottjer, D.J., 1989. Ichnofabric of sandstone deposited in high-energy
nearshore environments: measurement and utilization. Palaios 4, 598–604.
Einsele, G., Chough, S.K., Shiki, T., 1996. Depositional events and their records—an
introduction. Sedimentary Geology 104, 1–9.
Fielding, C.R., 1984. Upper delta plain lacustrine and fluvio-lacustrine facies from the
Westphalian of the Durham coalfield, NE England. Sedimentology 31, 119–140.
Fielding, C.R., 1986. Fluvial channel and overbank deposits from the Westphalian of the
Durham coalfield, NE England. Sedimentology 33, 119–140.
Fürsich, F.T., Oschmann, W., 1986. Storm shell beds of Nanogyra virgula in the Upper
Jurassic of France. Neues Jahrbuch für Geologie und Paläontologie, Abhandlungen
172, 141–161.
Fürsich, F.T., Oschmann, W., 1993. Shell beds as tools in basin analysis: the Jurassic of
Kachchh, western India. Journal of Geological Society, London 150, 169–185.
Fürsich, F.T., Pandey, D.K., 1999. Genesis and environmental significance of Upper
Cretaceous shell concentrations from the Cauvery Basin, southern India. Palaeo-
geography, Palaeoclimatology, Palaeoecology 145, 119–139.
Fürsich, F.T., Pandey, D.K., 2003. Sequence stratigraphic significance of sedimentary
cycles and shell concentrations in the Upper Jurassic –Lower Cretaceous of
Kachchh, western India. Palaeogeography, Palaeoclimatology, Palaeoecology 193,
285–309.
Fürsich, F.T., Oschmann, W., Jaitly, A., Singh, I.B., 1991. Faunal response to
transgressive–regressive cycles: example from the Jurassic from Western India.
Palaeogeography, Palaeoclimatology Palaeoecology 85, 149–159.
Gifu-ken Dinosaur Research Committee, 1993. Report on the Dinosaur Fossil Excavation
in Gifu Prefecture, Japan. Gifu Prefectural Museum, Gifu. p. 46.
Gingras, M.K., Rasamen, M., Ranzi, A., 2002. The significance of bioturbated inclined
heterolithic stratification in the southern part of the Miocene Solimoes Formation,
Rio Acre, Amazonia, Brazil. Palaios 17, 591–601.
Hallam, A., 1967. The interpretation of size–frequency distributions in molluscan death
assemblages. Palaeontology 1, 25–42.
Hayami, I., 1958. A review of so-called “Cyrenoides” in Japan. Japanese Journal of
Geology and Geography 29, 11–27.
Hirayama, R., 1996. Fossil land turtles from the Tetori Group (early Cretaceous) of
Shokawa Village, Gifu Prefecture, central Japan. Abstracts of the 1996 Annual
Meeting of the Geological Society of Japan, p. 162.
Hirayama, R., Brinkman, D.B., Danilov, I.G., 2000. Distribution and biogeography of non-
marine Cretaceous turtles. Russian Journal of Herpetology 7, 181–198.
Hughes, D.A., Lewin, J., 1982. A small-scale flood plain. Sedimentology 29, 891–895.
Isozaki, Y., Maruyama, S., 1991. Studies on orogeny based on plate tectonics in Japan
and new geotectonic subdivisions of the Japanese Island. Journal of Geography 100,
697–761.
Kidwell, S.M., 1986. Models for fossil concentrations: palaeobiologic implications.
Palaeobiology 12, 6–24.
Kidwell, S.M., 1991. The stratigraphy of shell concentrations. In: Allison, P.A., Briggs, D.E.
G. (Eds.), Taphonomy: Releasing the Data Locked in the Fossil Record. Plenum
Press, New York, pp. 115–209.
Kidwell, S.M., Fürsich, F.T., Aigner, T., 1986. Conceptual framework for the analysis and
classification of fossil concentrations. Palaios 1, 228–238.
Kobayashi, T., Suzuki, K., 1937. Non-marine shells of the Jurassic Tetori series in Japan.
Japanese Journal of Geology and Geography 14, 33–51.
Komatsu, T., Chen, J.H., Wang, Q.F., 2003. Bivalve and charophyte fossils from the Tetori
Group: a clue to stratigraphic correlation of late Mesozoic non-marine deposits
between Japan and China — a preliminary work. Memoir of the Fukui Prefectural
Dinosaur Museum 2, 43–49.
Kondo, Y., 1997. Inferred bivalve response to rapid burial in a Pleistocene shallow-
marine deposit from New Zealand. Palaeogeography, Palaeoclimatology, Palaeoe-
cology 128, 87–100.
Kranz, P.M., 1974. The anastrophic burial of bivalves and its paleoecological
significance. Journal of Geology 82, 237–265.
Kumar, N., Sanders, J.E., 1976. Characteristics of shoreface storm deposits: modern and
ancient examples. Journal of Sedimentary Petrology 46, 145–162.
Kumon, F., Umezawa, T., Nishikawa, T., Matsuzawa, T., 1994. New occurrence of
belemnite from the Itoshiro Subgroup of the Tetori Group in Shokawa-mura, Gifu
Prefecture. Abstracts of the 1994 Annual Meeting of the Geological Society of Japan,
p. 95.
Kusuhashi, N., Matsumoto, A., Murakami, M., Tagami, T., Hirata, T., Iizuka, T., Handa, T.,
Matsuoka, H., 2006. Zircon U–Pb ages from tuff beds of the upper Mesozoic Tetori
Group in the Shokawa district, Gifu Prefecture, central Japan. Island Arc 15,
378–390.
Lucchi, F.R., 1995. Sedimentgraphica: Photographic Atlas of Sedimentary Structures.
Columbia University Press, New York.
Maeda, S., 1952. A stratigraphical study on the Tetori series in the upper Shiokawa
district in Gifu Prefecture. Journal of the Geological Society of Japan 58, 145–153.
Maeda, S., 1954. On the Tetori Forest Fossils from the upper reaches of the Sho River
district, Gifu Prefecture. Studies of the Geological and Mineralogical Institute of the
Tokyo University of Education 3, 43–47.
Maeda, S., 1961. On the geological history of the Mesozoic Tetori Group in Japan. Journal
of the College of Arts and Science, Chiba University 3, 396–426.
Manabe, M., Evans, S.E., 1996. The earliest record of a choristodere from East Asia (the
middle early Cretaceous Okurodani Formation, Tetori Group, Shokawa, Gifu, Japan).
Abstracts of the 1996 Annual Meeting of the Palaeontological Society of Japan, p. 55.
Maruyama, S., Isozaki, Y., Kimura, G., Terabayashi, M., 1997. Paleogeographic maps of
the Japanese Island: plate tectonics synthesis from 750 Ma to Present. Island Arc 6,
121–142.
Matsukawa, M., Ido, K., 1993. Nonmarine molluscan communities and palaeoecology in
the Jurassic–Cretaceous Tetori Group, Japan. Cretaceous Research 14, 365–381.
Matsukawa, M., Nakada, K., 1999. Stratigraphy and sedimentary environment of the
Tetori Group in its central distribution based on nonmarine molluscan assem-
blages. Journal of the Geological Society of Japan 105, 817–835.
Matsukawa, M., Nakada, K., 2003. Adaptive strategy and evolution of corbiculoids based
on the Japanese Mesozoic fossils. Bulletin of Tokyo Gakugei University Section 4
(55), 161–189.
Matsukawa, M., Higuchi, M., Yamamoto, T., Ido, K., 1993. Primary information on death
assemblages in fluvial environments — comparative analysis between living and
death fresh-water molluscan assemblages in the Kunichigawa River, southern
Matsuyama, Japan—. Journal of the Geological Society of Japan 99, 643–657.
Matsukawa, M., Kawashima, K., Koarai, K., 1996. Nonmarine molluscan assemblages in
the middle part of the Tetori Group (Upper Jurassic to Lower Cretaceous), Gifu
Prefecture, central Japan. Bulletin of Tokyo Gakugei University Section 4 (48),
101–113.
Matsukawa, M., Takahashi, O., Hayashi, K., Ito, M., Konovalov, V.P., 1997. Early
Cretaceous paleogeography of Japan, based on tectonic and faunal data. Memoires
of the Geological Society of Japan 48, 29–42.
Matsukawa, M., Ito, M., Hayashi, K., Takahashi, O., Yang, S.Y., Lim, S.K., 1998. Evaluation
of nonmarine bivalves as chronological indices, based on examples from the Lower
Cretaceous of East Asia. New Mexico Museum Natural History and Science Bulletin
14, 125–133.
Matsukawa, M., Koarai, K., Shionoya, S., Shinkai, T., Nakada, K., Matsui, T., Aono, H.,
Kobayashi, N., Okubo, A., Hayashi, K., Ito, M., 2003. Stratigraphy and sedimentary
basin developments of the Tetori Group in its main area, central Japan. Journal of
the Geological Society of Japan 109, 383–398.
Matsukawa, M., Ito, M., Nishida, N., Koarai, K., Lockley, M.G., Nichols, D.J., 2006. The
Cretaceous Tetori biota in Japan and its evolutionary significance for terrestrial
ecosystem in Asia. Cretaceous Research 27, 199–225.
McPherson, J.G., Shanmugam, G., Moiola, R.J., 1987. Fan-deltas and braid deltas:
varieties of coarse-grained deltas. Geological Society of America Bulletin 99,
331–340.
Meldahl, K.H., Cutler, A.H., 1992. Neotectonics and taphonomy: Pleistocene molluscan
shell accumulations in the northern Gulf of California. Palaios 7, 187–197.
Müller, A.H., 1951. Grundlagen der Biostratinomie. Abhandlungen der Deutschen
Akademie der Wissenschaften Berlin, Klasse Mathematik und allgemeine Nat-
urwissenschaften, Jahrgang 1950 (3), 1–147.
Myrow, P.M., Southard, J.B., 1996. Tempestite deposition. Journal of Sedimentary
Research 66, 875–887.
Noe-Nygaard, N., Surlyk, F., Piasecki, S., 1987. Bivalve mass mortality caused by toxic
dinoflagellate blooms in a Berriasian–Valanginian lagoon, Bornholm, Denmark.
Palaios 2, 263–273.
Pearson, N.J., Gingras, M.K., 2006. An ichnological and sedimentological facies model for
muddy point-bar deposits. Journal of Sedimentary Research 76, 771–782.
Radley, J., Barker, M.J., 2000. Palaeoenvironmental significance of storm coquinas in a
Lower Cretaceous coastal lagoonal succession (Vectis Formation, Isle of Wight,
southern England). Geological Magazine 137, 193–205.
Reading, H.G., Collinson, J.D., 1996. Clastic coast, In: Reading, H.G. (Ed.), Sedimentary
Environments: Processes, Facies and Stratigraphy, 3rd ed. Blackwell Science Ltd.,
Oxford, pp. 154–231.
Reineck, H.E., 1967. Layered sediments of tidal flats, beaches, and shelf bottoms of the
North Sea. In: Lauff, G.H. (Ed.), Estuarine: Sediments and Sedimentation: American
Association for the Advancement of Science Publication, 83, pp. 191–206.
Reineck, E.H., Singh, I.B., 1980. Depositional Sedimentary Environments. Springer-
Verlag, Berlin.
 S. Yamashita et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 308 (2011) 476–491
Reineck, E.H., Wunderlich, F., 1968. Classification and origin of flaser and lenticular
bedding. Sedimentology 11, 99–104.
Schäfer, W., 1972. Ecology and Palaeoecology of Marine Environments. Oliver & Boyd,
Edinburgh.
Shibuya, K., Kato, J., 1989. The mass generation of bivalve, Corbicula japonica in
Hachirou-gata lagoon. The Business Reports of Aquatic Resources in Akita
Prefecture, 15, pp. 60–105.
Shimoyama, S., 1985. Size–frequency distribution of living populations and dead shell
assemblages in a marine intertidal sand snail, Umbonium (Suchium) moniferum
(Lamarck), and their palaeoecological significance. Palaeogeography, Palaeoclima-
tology, Palaeoecology 49, 327–358.
Sigurdsson, J.B., Titman, C.W., Davies, P.A., 1976. The dispersal of young post-larval
bivalves molluscs by byssus threads. Nature 262, 386–387.
Stenzel, H.B., 1971. Oysters. In: Moore, R.C. (Ed.), Treatise on Invertebrate Paleontology.
: Part N. Mollusca 6. Bivalvia, Vol. 3. Geological Society of America & University of
Kansas, Boulder, Colorado & Lawrence, Kansas, pp. 953–1197.
Surlyk, F., 1972. Morphological adaptations and population structures of the Danish
chalk brachiopods (Maastrichtian, Upper Cretaceous). Det Kongelige Danske
Videnskabernes Selskab Biologiske Skrifter, 19, p. 57.
Surlyk, F., 1974. Life habitat, feeding mechanism and population structure of the
Cretaceous brachiopod genus Aemula. Palaeogeography, Palaeoclimatology,
Palaeoecology 15, 185–203.
Takada, Y., Sonoda, T., Nakamura, M., Nakao, S., 2000. Growth and settlement of the
bivalve Corbicula japonica population in Lake Shinji. Nippon Suisan Gakkaishi 64,
678–686.
Terwindt, J.H.J., 1971. Lithofacies of inshore estuarine and tidal inlet deposit. Geologie
en Mijnbouw 50, 515–526.
View publication stats
491
Terwindt, J.H.J., Breusers, H.N.C., 1972. Experiments on the origin of flaser, lenticular
and sand-clay alternating bedding. Sedimentology 19, 85–98.
The Gifu-ken Dinosaur Fossil Excavation Party, 1999. Sedimentary environments of the
Tetori Group in the Okurodani area, Shokawa-mura, Gifu Prefecture, central Japan.
Report on Gifu Prefectural Museum, 20, pp. 1–8.
The Gifu-ken Dinosaur Fossil Excavation Party, 2000. Sedimentary environments of the
Tetori Group in the Kobudani area, Shokawa-mura, Gifu Prefecture, central Japan.
Report on Gifu Prefectural Museum, 21, pp. 1–10.
Trewin, N.H., Welsh, W., 1976. Formation and composition of graded estuarine shell
beds. Palaeogeography, Palaeoclimatology, Palaeoecology 19, 219–230.
Umezawa, T., 1997. Sedimentary facies and sedimentary environments of the Tetori
Group in the Shokawa Village, Gifu Prefecture, Japan. Report on Gifu Prefectural
Museum, 18, pp. 73–83.
Yabumoto, Y., 1997. Fresh water fish fossils from the Tetori Group in Shokawa Village,
Gifu Prefecture. Abstracts of the 1997 Annual Meeting of the Palaeontological
Society of Japan, p. 26.
Yokoyama, M., 1889. Jurassic plants from Kaga, Hida, and Tosa. Journal of the Faculty of
Science, Imperial University of Tokyo 3, 1–66, pls. 1–14.
Zaitlin, B.A., Dalrymple, R.W., Boyd, R., 1994. The stratigraphic organization of incised-
valley systems associated with relative sea-level change. In: Dalrymple, R.W., Boyd,
R., Zaitlin, B.A. (Eds.), Incised-Valley Systems: Origin and Sedimentary Sequences:
SEPM Special Publication, 51, pp. 45–60.
Zuschin, M., Harzhauser, M., Mandic, O., 2005. Influence of size-sorting on diversity
estimates from tempestitic shell beds in the Middle Miocene of Austria. Palaios 20
(2), 142–158.