Cretaceous Research 33 (2012) 21e41

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Cretaceous Research
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Adaptive morphologies and guild structure in a high-diversity bivalve fauna

from an early Campanian rocky shore, Ivö Klack (Sweden)
Anne Mehlin Sørensen a, *, Finn Surlyk a, John W.M. Jagt b
a
Department of Geography and Geology, University of Copenhagen, Øster Voldgade 10, DK-1350 Copenhagen K, Denmark
b
Natuurhistorisch Museum Maastricht, de Bosquetplein 6-7, NL-6211 KJ Maastricht, The Netherlands

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

Article history: The bivalve fauna from a late early Campanian rocky shore at Ivö Klack (southern Sweden), comprises
Received 10 March 2011 just over sixty species, a very high diversity in comparison to other Late Cretaceous and modern rocky
Accepted in revised form 31 July 2011 shore bivalve assemblages. This high diversity is here considered to represent a reliable census of the
Available online 27 August 2011
fauna; only in part can it be explained by the cumulative effect of generations of bivalves inhabiting this
coastal environment. The high density and diversity and the wide range of shell morphologies allow
Keywords:
interpretation of different modes of life in this variable environment with many contrasting habitats.
Bivalves
Study of the functional morphology of bivalve shells and comparison with extant relatives has resulted in
Palaeoecology
Rocky shore
a subdivision of the fauna into seven guilds and five habitats. The bivalve fauna represents a within-
Late Cretaceous habitat, time-averaged assemblage to which none of the species was introduced from adjacent envi-
Sweden ronments. It includes some of the most northerly known, very small rudistid bivalves, in addition to the
oldest known occurrences of Mytilus and Barbatia in association with rocky shores. Bivalves constituted
the most important invertebrate group inhabiting the late early Campanian rocky shore at Ivö Klack, in
terms of diversity, density and biomass.
Ó 2011 Elsevier Ltd. All rights reserved.

1. Introduction 1885) are predominant. The lowermost, photonegative zone

Rocky shores provide many different habitats for the colonising
fauna and are therefore often characterised by a high faunal density
and diversity. Studies of ancient rocky shore faunas are few and far
between, primarily because rocky shores are sites of erosion rather
than of deposition, and the shelly fauna easily undergoes severe
fragmentation in such high-energy settings. However, good
examples of ancient rocky shores do exist, with the encrusting
fauna well preserved (Surlyk and Christensen, 1974; Lescinsky et al.,
1991; Johnson and McKerrow, 1995; Desrochers, 2006). A well-
exposed late early Campanian rocky shore is preserved at Ivö
Klack in southern Sweden; it yields an invertebrate fauna of both
high density and diversity (Surlyk and Sørensen, 2010). A biological
interpretation, comprising three zones, of the cementing epifauna
has been mapped out on large gneiss boulders and hummocks
(Surlyk and Christensen, 1974). The highest zone is dominated by
the spondylid bivalve Spondylus labiatus (Wahlenberg, 1821), while
in the next, lower, zone on the near-vertical sides of the boulders
the oyster Amphidonte haliotoideum (J. Sowerby, 1813) and the
inarticulate craniid brachiopod Ancistrocrania stobaei (Lundgren,
* Corresponding author. Tel.: þ45 35322401.
E-mail address: anne@geo.ku.dk (A. M. Sørensen).
shows a predominance of serpulid species which occupy the over-
hanging lower portions of the boulders and hummocks (Sørensen
and Surlyk, 2010). The commonly excellent preservation of this
ancient rocky shore fauna offers an opportunity to increase the
knowledge of faunal composition, life habits and nature of the
ecosystem. Thus, the aim of the present paper is to interpret the
mode of life of the bivalve assemblages, to compare these with
other Late Cretaceous and modern rocky shores faunas and to
subdivide the bivalves into guilds and preferred habitats in order to
interpret their ecological role on the early Campanian rocky shore.
2. Geological setting
The Ivö Klack site is a disused kaolin quarry, on the island of Ivö
in lake Ivösjön, in the northern part of the Kristianstad Basin in
north-east Skåne, southern Sweden (Fig. 1). This basin was situated
at a palaeolatitude of approximately 50 N in a warm-temperate to
subtropical climate (Surlyk, 1997; Surlyk and Sørensen, 2010).
Global sea level was in the order of 100 m higher than today
(Kominz et al., 2008), and the basin was transgressed repeatedly
from the south, resulting in the formation of an archipelago
with low islands and peninsulas (Surlyk and Christensen, 1974;

0195-6671/$ e see front matter Ó 2011 Elsevier Ltd. All rights reserved.
doi:10.1016/j.cretres.2011.07.004
22 A.M. Sørensen et al. / Cretaceous Research 33 (2012) 21e41
Fig. 1. Simplified geological map of north-east Skåne (southern Sweden), showing the Kristianstad Basin and the locality of Ivö Klack (modified after Norling and Bergström, 1987).
Christensen, 1975). As the Late Cretaceous transgression pro-
gressed, the deeply weathered basement at Ivö Klack was flooded
and washed clean by waves, leaving a steep, irregular, boulder-
strewn and hummocky rocky coast. The shore occupied the inter-
tidal and subtidal zones with a maximum water depth probably not
more than c. 5 m (Surlyk and Christensen, 1974). The succession at
Ivö Klack consists of Precambrian gneiss overlain by its weathering
product, kaolin. On top of the kaolin follow a few metres of Cam-
panian quartz sand washed out from the kaolin and, in turn,
overlain by coarse-grained marine carbonates composed of skeletal
fragments of bivalves (with a predominance of oysters), bryozoans,
brachiopods and echinoids (Surlyk and Sørensen, 2010). Belemnites
are abundant and show that the onlapping carbonates belong to the
upper lower Campanian Belemnellocamax mammillatus zone, c.
81e80 Ma (Christensen, 1975).
3. Material and methods
A large portion of the material studied was collected in
1928e1929 by Alfred Rosenkrantz and in the early 1970s by Finn
Surlyk and Walter Kegel Christensen; this is now housed at the
Statens Naturhistorisk Museum, Copenhagen, with the prefix GMI
(Geological Museum Ivö). The abundant, large-sized shells are well
preserved, whereas juvenile specimens are almost absent, probably
due to shell breakage and fragmentation in this turbulent environ-
ment. Bivalve shells are normally represented by isolated, dissoci-
ated valves and many of the thick-shelled species reveal numerous
sponge borings. A total of 19 species had aragonitic shells; these are
represented by internal moulds which make proper taxonomic
assignment rather difficult. One species, a trigoniid, is represented
by external moulds as well. The abundance of moulds of formerly
aragonitic species is indicative of the slow dissolution of aragonite in
the turbulent, near-shore environment (compare Cherns and
Wright, 2000). Epifaunal encrustation of oyster muscle scars,
subsequent to aragonite dissolution, indicates that dissolution
occurred on the seafloor or under very shallow burial and subse-
quent reworking of the sediment during storms, exposing the oyster
shells on the seafloor. The bivalve guilds and habitats are interpreted
on the basis of studies of functional morphology of the shells and
a comparison with modern relatives.
4. Results
The bivalve fauna from Ivö Klack comprises just over sixty
species, in around 40 genera, and inclusive of four indeterminate
forms which are briefly described below and listed in Table 1.
Commonest are oysters; these make up c. 90 per cent of all bivalve
shells recorded in the present study. The assemblages are sub-
divided into seven guilds, on the basis of tiering and postulated
mode of life (Fig. 2; Table 1). It has not been possible to assign
a single indeterminate form, GMI 2602 (Fig. 5KeL) to any guild. The
epifaunal, free-lying guild comprises three species, the epifaunal
cemented guild 14 (primarily oysters and spondylids), the epi-
byssate guild 23, the semi-infaunal byssally attached guild one, the
shallow infaunal guild 11, and the deep infaunal guild eight. Finally,
the boring guild is documented by borings assignable to the ich-
nogenus Gastrochaenolites Leymerie, 1842, found in oyster shells
(Fig. 3). No shells of boring bivalves have been found, but at least
one species must have belonged to this guild.
Five habitats that represent different hydrodynamic conditions
and levels of light penetration are distinguished (Table 2). The
interpretation of habitat preferences of the bivalve species listed is
based primarily on attachment strategy. A total of 20 species
occurred in the within-sediment habitat 1; they are all infaunal and
 A.M. Sørensen et al. / Cretaceous Research 33 (2012) 21e41 23

Table 1 open spaces within bundles of marine plants. Thirty-five species

Abundance, shell composition and guild structure of the bivalve fauna from Ivö represent the relatively protected habitat 3, between and beneath
Klack. GMI numbers indicate unidentified species. GMI 2602 is not included in this
table, since the guild could not be inferred. *Assignment to guild is tentative;
boulders, and these include byssally attached, cemented as well as
a ¼ abundant (>100 specimens), c ¼ common (5 < specimens<100), r ¼ rare free-lying species. The semi-exposed habitat 4 hosted twenty-two
(specimens<5), A ¼ aragonite, C ¼ calcite. species, inclusive of byssally attached, cemented as well as free-
Species Abundance Composition Guild
lying species. The most exposed habitat, habitat 5, was occupied
by twelve cemented and byssally attached species (Table 2).
Neithea quinquecostata c CeA Epifaunal free-lying
*Vultogryphaea lacinata c CeA guild
*GMI 3501 r CeA 4.1. The bivalve fauna
Amphidonte haliotoideum a CeA Epifaunal cemented
Acutostrea incurva c CeA guild Size indications below include shell height (between umbo and
Ceratostreon sp. r CeA ventral margin) and length (measured from anterior to posterior).
*Exogyra? sp. r CeA Identification of material has relied on various sources, inclusive of
Gryphaeostrea canaliculata c CeA
Nilsson (1827), Lundgren (1885, 1894), Holzapfel (1887e1889),
Hyotissa semiplana r CeA
Pycnodonte vesicularis c CeA Hennig (1897), Cox et al. (1969), Stanley (1970), Stenzel (1971),
Rastellum diluvianum a CeA Dhondt (1972, 1973), Marquet (1982), Abdel-Gawad (1986), Mal-
Biradiolites suecicus c CeA chus (1990), Waller and Marincovich (1992), Dhondt and Dieni
Spondylus labiatus a CeA (1993), Malchus et al. (1994, 1996) and Elder (1996).
Spondylus latus r CeA
Spondylus truncatus c CeA
*Spondylus lamellatus c CeA Arcidae
Spondylus sp. A c CeA 1. Barbatia sp. (Fig. 9B)
Anomia? sp. r CeA Epibyssate guild
Mytilus sp. r CeA Internal moulds only; of medium size (length up to 55 mm);
Barbatia sp. c A outline elongate, oval; long, near-straight taxodont hinge line;
Barbatia? sp. r A pallial line visible along entire ventral margin; outer surface with
Modiolus sp. A r CeA
radial ribs.
Lima ovata c CeA
Ctenoides sp. r CeA
Limatula semisulcata c CeA 2. Barbatia? sp. (Fig. 9H)
Limea sp. r CeA
Pseudolimea sp. r CeA
Internal mould only; one specimen known, of medium size
*Camptonectes virgatus r CeA
Dhondtichlamys pulchella c CeA
(length 42 mm); outline triangular, suboval; only part of hinge line
Dhondtichlamys subarata c CeA preserved; pallial line visible along entire ventral margin. In
Mimachlamys undulata r CeA comparison to Barbatia sp. (see above), umbo less acuminate, wider
Lyriochlamys faujasi r CeA and straighter; outer surface with radial ribs.
Chlamys septemplicata c CeA
Lyriochlamys dentata c CeA
Lyriochlamys ternata r CeA 3. Arca? sp. (Fig. 9I)
Lycettia sp. r CeA
*Plagiostoma hoperi r CeA Internal mould only; one specimen known, of large size (length
Isognomon sp. r CeA
81 mm); outline elongate; valve inflated; beak prominent and
Pectinidae indet. r CeA
GMI 3701 r ?
inflated; umbo positioned anteriorly; ventral margin concave
centrally; no dentition or ornament seen.
Modiolus sp. B c CeA Semi-infaunal
byssus-attached guild
Cucullaeidae
Cardiinae sp. c A Shallow infaunal guild 4. Cucullaea exaltata (Nilsson, 1827) (Fig. 6R and S)
Arctica sp. c A
Granocardium? sp. r A
Cucullaea exaltata c A Internal moulds only, of medium size (up to 53 mm in height);
Cucullaea sp. A c A outline subtrigonal to subtrapezoidal; posterior margin obliquely
Cucullaea sp. B c A truncated; rounded posterior umbonal carina in both valves; pallial
Trigonia sp. c A
line prominent, visible along entire ventral margin; no gapes; no
Glycymeris lens c A
Glycymeris sp. A c A ornament or dentition preserved.
*Icanotia grosseplicata c A
Liopistha sp. r A 5. Cucullaea sp. A (Fig. 6M and N)
Goniomya sp. c A Deep infaunal guild
Arcomya sp. c A Internal moulds only, of medium size (up to 40 mm in height);
Arca? sp. r A outline subtrigonal to subtrapezoidal; posterior margin obliquely
Panopaea regularis c A
truncated; rounded posterior umbonal carina in both valves; pallial
Tellinidae? indet. c ?
Laternulidae sp. r A
line prominent, traceable along entire ventral margin; no gapes.
Tellina? sp. r A Compared to C. exaltata (see above), this has a deeper scar posterior
GMI 1715 r ? of the carina, a rounded postero-ventral margin and a more
no fossils ? Boring guild acuminate beak; no ornament or dentition preserved.

6. Cucullaea sp. B (Fig. 6G and H)

semi-infaunal. Seventeen species of byssate nestling and fissure-
dwelling bivalves inhabited the protected photonegative habitat Internal moulds only, of medium size (up to 49 mm in height);
2, between and beneath hummocks and boulders and probably in outline subtrigonal to suboval; posterior margin obliquely
24 A.M. Sørensen et al. / Cretaceous Research 33 (2012) 21e41
Fig. 2. The seven bivalve guilds from Ivö Klack. Representing the free-lying epifaunal guild is Vultogryphaea lacinata (A); typical of the cemented epifaunal guild are Rastellum
diluvianum (B) and spondylids (C); representing the epibyssate guild are Modiolus sp. A (D) and Mimachlamys undulata (E); a typical member of the semi-infaunal guild is Modiolus
sp. B (F); representing the shallow infaunal guild is Cucullaea sp. A (G) and Arcomya sp. (H) is typical of the deep infaunal guild. The boring guild is represented by a boring in an
oyster valve (see also Fig. 3).
truncated; rounded posterior umbonal carina in both valves. Nar-
rower than C. exaltata and lacking a prominent pallial line. No
ornament or dentition preserved.
Glycymerididae
7. Glycymeris lens (Nilsson, 1827) (Fig. 6F)
Internal moulds only, of small size (up to 25 mm in height);
outline subcircular; umbo orthogyrate and relatively straight; both
valves evenly inflated; pallial line visible along entire ventral
margin, internally fluted ventrally; taxodont hinge well preserved;
no ornament seen.
8. Glycymeris sp. A (Fig. 6P)
Internal mould only; one specimen known, of small size (32 mm
in height); outline subcircular; umbo orthogyrate and relatively
straight; ventral margin internally fluted; outer surface with radial
ribs. No dentition preserved.
Mytilidae
9. Mytilus sp. (Fig. 8N)
Internal mould only; one specimen known, of medium size
(length 60 mm); outline mytiliform; beak acuminate; anterior
adductor scars small, yet distinct; valve inflated, ventral shell
margin straight and flattened; outer surface with concentric
growth lines; no dentition preserved.
Fig. 3. Bivalve boring, assignable to the ichnogenus Gastrochaenolites, in a right valve
of the oyster Rastellum diluvianum.
10. Modiolus sp. A (Fig. 8O)
Internal moulds only; of medium size (length up to 62 mm);
outline modioliform; umbo obtuse and ventral margin curved;
ligament area fairly long; both valves equally inflated; no dentition
or ornament seen.
11. Modiolus sp. B (Fig. 8H)
Internal moulds only; of medium size (length up to 52 mm); outline
modioliform, wider than Modiolus sp. A (see above); umbo obtuse,
ventral margin more clearly curved than in previous form; ligament
area fairly long; valve inflated; no dentition or ornament seen.
12. Lycettia sp. (Fig. 8I)
Internal mould only, one specimen known; of small size (length
34 mm); outline mytiliform; beak more acuminate than in Mytilus
sp. (see above); sharp carina extending from beak to postero-ventral
margin; valve inflated; ventral margin straight and flattened; outer
surface with concentric growth lines; no dentition seen.
Isognomonidae
13. Isognomon sp. (Fig. 9D and J)
Internal moulds only, of large size (at least 90 mm in width);
beak area acuminate; only umbonal area preserved, but with
characteristic numerous and regularly arranged ligament grooves,
each about 3 mm wide and 9 mm long; no ornament preserved.
Pectinidae
14. Camptonectes virgatus (Nilsson, 1827) (Fig. 8P)
Small sized (up to 11 mm in height); outline drop shaped; valves
relatively convex, with unequal auricles; umbonal angle 70 ; right
anterior auricle elongate, wing like, with byssal sinus; left anterior
auricle near rectangular; posterior auricles smaller and more
obtusely angled; outer surface with radial striae, diverging from
umbo towards pallial and lateral margins; striae crossed by
concentric growth lines.
15. Lyriochlamys faujasi (Defrance, 1825) (Fig. 8J)
Large sized (up to 85 mm in height); outline drop shaped; valves
prosocline and flattened; umbonal angle 75e83 ; anterior auricle
rather wide, elongate and wing like, with deep byssal sinus and
covered with concentric elevated lines which bend along byssal
sinus; posterior auricle smaller, triangular and acutely angled;
outer surface with 25e35 radial, divided ribs.
 A.M. Sørensen et al. / Cretaceous Research 33 (2012) 21e41 25

Fig. 4. Gryphaeid and ostreid oysters from Ivö Klack; all specimens natural size. AeB. Vultogryphaea lacinata (GMI 4101), left valve in outer and anterior views; CeD. Vultogryphaea
lacinata (GMI 4102), right valve in internal and external views; EeF. Gryphaeostrea canaliculata (GMI 4503), left valve in external and posterior views (attachment area facing
upwards); G. Ceratostreon sp. (GMI 4201), left valve (attachment area facing upwards); H. Hyotissa semiplana (GMI 4107), right valve in internal view; IeJ. Pycnodonte vesicularis (GMI
4505), right valve in internal and external views; KeL. Pycnodonte vesicularis (GMI 4504), right valve in internal and external views; MeN. Ceratostreon sp. (GMI 4203), left valve in
external and internal views; O. Amphidonte haliotoideum (GMI 4105), left valve in internal view; PeQ. Amphidonte haliotoideum (GMI 4104), right valve in internal and external
views; R. Pycnodonte vesicularis forma hippopodium (GMI 4502), right valve in internal view; SeT. Pycnodonte vesicularis forma hippopodium (GMI 4501), left valve in internal and
ventral views; UeV. Gryphaeostrea canaliculata (GMI 4401), right valve in external and internal views.
26 A.M. Sørensen et al. / Cretaceous Research 33 (2012) 21e41

Fig. 5. Bivalves from Ivö Klack; all specimens natural size, except AeC, which are x 0.8. AeC. Rastellum diluvianum (GMI 6501), right valve in internal view (A), right valve in external
view (B) and ventral view (C), left valve at base; D. Acutostrea incurva (GMI 6401), left valve in external view; EeF. Gryphaeostrea canaliculata (GMI 4202), left valve in anterior and
internal views; G. Rastellum diluvianum (GMI 4901), left valve in external view; H. Tellina? sp. (GMI 1704), internal mould of right valve; IeJ. Exogyra? sp. (GMI 4506), right valve in
internal and external views; KeL. indeterminate (GMI 2602), internal mould of right valve in posterior and external views; MeN. Acutostrea incurva (GMI 6402), left valve in internal
and external views.
 A.M. Sørensen et al. / Cretaceous Research 33 (2012) 21e41 27

Table 2
The bivalve fauna and its link with five habitats at Ivö Klack. The habitats represent different hydrodynamic conditions and light penetration.

Increasing energy and illumination

Life habit Species Habitat 1 Habitat 2 Habitat 3 Habitat 4 Habitat 5

Within-sediment Protected photonegative Relatively protected Semi-exposed parts Most exposed parts
with limited environments between and environments of the boulders and of the boulders with
turbulence beneath boulders and in between and on the seafloor high water
spaces within bundles of beneath boulders turbulence
marine plants
Byssate current Mytilus sp. X X
loving Modiolus sp. A
Lycettia sp.
GMI 3701

Cemented by the Amphidonte haliotoideum X X X

entire left valve Ceratostreon sp.
Exogyra? sp.
Pycnodonte vesicularis
Hyotissa semiplana
Spondylus sp. A
Spondylus latus
Spondylus truncatus

Cemented by a small Acutostrea incurva X X

area near the umbo Gryphaeostrea canaliculata
Rastellum diluvianum
Biradiolites suecicus
Spondylus labiatus
Spondylus lamellatus

Epifaunal, free-lying Neithea quinquecostata X X

Vultogryphaea lacinata
GMI 3501
GMI 3803

Byssate nestling and Barbatia sp. X X

fissure dwellers Barbatia? sp.
Lima ovata
Ctenoides sp.
Limea sp.
Pseudolimea sp.
Limatula semisulcata
Camptonectes virgatus
Dhondtichlamys pulchella
Dhondtichlamys subarata
Mimachlamys undulata
Lyriochlamys faujasi
Chlamys septemplicata
Lyriochlamys dentata
Plagiostoma hoperi
Isognomon sp.
Pectinidae indet.

Semi-infaunal Modiolus sp. B X

Shallow infaunal Cardiinae sp. X
Arctica sp.
Granocardium? sp.
Cucullaea exaltata
Cucullaea sp. A
Cucullaea sp. B
Trigonia sp.
Glycymeris lens
Glycymeris sp. A
Icanotia grosseplicata
Liopistha sp.

Deep infaunal Goniomya sp. X

Arcomya sp.
Arcomya? sp.
Panopaea regularis
Laternulidae sp.
Tellinidae? indet.
Tellina? sp.
GMI 1715
28 A.M. Sørensen et al. / Cretaceous Research 33 (2012) 21e41
16. Lyriochlamys dentata (Nilsson, 1827) (Fig. 8L)
Large sized (up to 125 mm in height); outline drop shaped;
umbonal angle 72e76 ; right anterior auricle elongate and wing like,
with deep byssal sinus covered with radial ribs; left anterior
auricle large, with slight byssal sinus; posterior auricles much
smaller, obtusely angled and with few radial ribs; outer surface
with numerous, commonly tripartite ribs with scabrous
spinelets.
17. Lyriochlamys ternata (Münster in Goldfuss, 1833) (Fig. 8B)
Medium sized (32 mm in height), flattened; only a single valve
known; outline drop shaped; outer surface with rounded radial ribs
and riblets, broad intercostal intervals with radial lines and well-
developed, undulating concentric growth lines.
18. Dhondtichlamys pulchella (Nilsson, 1827) (Fig. 8E and F)
Small sized (up to 14 mm in height); outline rounded; umbonal
angle 101 105 ; right valve more convex than left; anterior auricle
elongate and wing like, with deep byssal sinus in right valve;
posterior auricle smaller and obtusely angled; outer surface with
a large number of radially divided ribs.
19. Dhondtichlamys subarata (Nilsson, 1827) (Fig. 8Q and R)
Small sized (up to 13 mm in height); outline rounded; umbonal
angle 90e97 ; shell relatively convex, left valve more so; anterior
auricle elongate and wing like, with deep byssal sinus; posterior
auricle smaller and triangular; outer surface with a large number of
divided ribs.
20. Mimachlamys undulata (Nilsson, 1827) (Fig. 8G)
Medium sized (38 mm in height); outline rounded; umbonal
angle at first 90 , but widening ventrally to 95 ; right valve fairly
convex, auricles relatively small; both auricles triangular but
anterior one with deep byssal sinus on right valve and covered with
radial ribs and pronounced concentric growth lines; outer surface
with thin, close-set radial ribs crossed by concentric, irregularly
distributed growth lines. Only a single right valve available.
21. Chlamys septemplicata (Nilsson, 1827) (Fig. 8K)
Medium sized (up to 50 mm in height); outline drop shaped;
umbonal angle 72e80 ; valves prosocline and flattened; right
anterior auricle broad, with acute angle and byssal sinus; posterior
auricle smaller with an obtuse angle; outer surface with rounded
radial ribs, broader intercostal intervals with finer radial ribs and
few, well-developed concentric growth lines.
22. Neithea quinquecostata (J. Sowerby, 1814) (Fig. 8C and D)
Medium sized (up to 50 mm in height); outline triangular; left
valve cup shaped, right valve flattened; auricles small, triangular
and of equal size; hinge with two divergent cardinal teeth, one on
each side of ligament pit; outer surface with six primary radial ribs
between with are four intercostal ribs.
23. Pectinidae indet. (Fig. 8V)
Small sized (16 mm in height); outline rounded; umbonal angle
90 ; left valve convex, with anterior auricle relatively small and
triangular with deep byssal sinus; outer surface with pronounced
concentric ornament, with distinct sinuous imbrication pattern,
continuing onto auricle. Only a single left valve available.
Anomiidae
24. Anomia? sp. (Fig. 7O)
Medium sized (45 mm in height); outline rounded; single valve
known weakly inflated; outer surface with concentric growth lines.
Spondylidae
25. Spondylus sp. A (Fig. 7B, K and Q)
Large sized (up to 100 mm in height); outline subrounded; right
valve bulbous and outer surface with strong radial ribs, rough
spines emerging from ribs; left valve deep and with large attach-
ment area; free portion of left valve with long, prominent spines
following ill-developed ribs and growth lines.
26. Spondylus labiatus (Wahlenberg, 1821) (Fig. 7A)
Medium sized (up to 50 mm in height); outline elongate to oval; left
valve inflated, cylindrical and cemented to substrate along small
portion of shell (boulders and hummocks); right valve lid like; outer
surface with strong radial ribs and few prominent growth lines. Species
commonly found in clusters, with radial, petal-like arrangement
pattern; easily recognised by strong radial ribs and elongate outline.
27. Spondylus latus (J. Sowerby, 1814) (Fig. 7C)
Medium sized (up to 50 mm in height); outline highly variable,
but mostly elongate to oval; right valve inflated, cylindrical; outer
surface of both valves with strong radial ribs and few, prominent
growth lines, may merge lamellae fused to either substrate or other
epifauna; left valve probably cemented to substrate by large
attachment area and by lamellae.
28. Spondylus truncatus (Lamarck, 1819) (Fig. 7F, G and L)
Medium sized (up to 40 mm in height); outline variably sub-
rounded; right valve gibbous with straight hinge line and small
auricles; outer surface with strong radial ribs, every fourth to sixth
rib being more prominent with rough spines; left valve cemented
to substrate by entire surface; shape of substrate recognisable on
right valve (xenomorphic expression).
29. Spondylus lamellatus (Nilsson, 1827) (Fig. 7E)
Medium sized (up to 48 mm in height); outline highly variable,
but mostly elongate to oval; valves inflated, cylindrical; outer
surface with strong radial ribs and prominent growth lines, all
merging into lamellae.
Limidae
30. Plagiostoma hoperi (Mantell, 1822) (Fig. 7P)
Medium sized (up to 35 mm in height); outline suborbicular to
suboval; opisthocline and length commonly slightly exceeding
height; beak anterior, with moderately long cardinal area, broad
ligament pit and obtuse auricles; outer surface radially striated,
with weak concentric growth lines.
31. Lima ovata (Nilsson, 1827) (Fig. 7H)
Small sized (up to 33 mm in height); outline suborbicular to
oval, height exceeding width; hinge margin relatively short;
 A.M. Sørensen et al. / Cretaceous Research 33 (2012) 21e41 29

Fig. 6. Bivalves from Ivö Klack; all specimens natural size. AeB. Cardiinae sp. (GMI 1601), internal mould of left valve in posterior and external views; CeD. Cardiinae sp. (GMI 1602),
internal mould of right valve in external and posterior views; E. ‘Trigonia’ sp. (GMI 1402), rubber peel of external surface of right valve, showing ornament; F. Glycymeris lens (GMI
1708), internal mould; G H. Cucullaea sp. B (GMI 1504), internal mould of right valve in external and anterior views; IeJ. ‘Trigonia’ sp. (GMI 1401), internal mould of right valve in
external and dorsal views; KeL. Granocardium? sp. (GMI 1705), internal mould of right valve in external and posterior views; MeN. Cucullaea sp. A (GMI 1502), internal mould of
right valve in external and anterior views; O. Liopistha sp. (GMI 1703), internal mould of right valve; P. Glycymeris sp. A (GMI 1716), internal mould; Q. Arctica sp. (GMI 1713), internal
mould of left valve; ReS. Cucullaea exaltata (GMI 1501), internal mould of right valve in external and anterior views; TeU. Arctica sp. (GMI 1503), internal mould of left valve in
external and posterior views.
30 A.M. Sørensen et al. / Cretaceous Research 33 (2012) 21e41

Fig. 7. Bivalves from Ivö Klack; all specimens natural size. A. Spondylus labiatus (GMI 304), right valve in external view; B. Spondylus sp. A (GMI 401), right valve in external view; C.
Spondylus latus (GMI 306), two left valves cemented together; D. Limatula semisulcata (GMI 302), right valve in external view; E. Spondylus lamellatus (GMI 308), right valve in
external view; F. Spondylus truncatus (GMI 307), right valve in external view; G. Interior of a left valve of an indeterminate spondylid, possibly S. truncatus (GMI 101); H. Lima ovata
(GMI 301), left valve in external view; I. Ctenoides sp. (GMI 1701), internal mould of left valve; J. indeterminate (GMI 1715), internal mould of right valve; K. Spondylus sp. A (GMI
309), right valve in external view; L. Spondylus truncatus (GMI 305), right valve in external view; M. Pseudolimea sp. (GMI 310), right valve in external view; N. Limea sp. (GMI 303),
right valve in external view; O. Anomia? sp. (GMI 3801), external view; P. Plagiostoma hoperi (GMI 311), right valve in external view; Q. Spondylus sp. A (GMI 21), right valve in
external view.
 A.M. Sørensen et al. / Cretaceous Research 33 (2012) 21e41 31

Fig. 8. Bivalves from Ivö Klack; all specimens natural size, except P. A. indeterminate pectinid (GMI 3501), right valve in external view; B. Lyriochlamys ternata (GMI 3803), right
valve in external view; C. Neithea quinquecostata (GMI 3503), left valve in external view; D. Neithea quinquecostata (GMI 3802), right valve in internal view; E. Dhondtichlamys
pulchella (GMI 3605), external view; F. Dhondtichlamys pulchella (GMI 3602), right valve in external view; G. Mimachlamys undulata (GMI 3502), left valve in external view; H.
Modiolus sp. B (GMI 1707), internal mould of right valve; I. Lycettia sp. (GMI 402), internal mould of left valve; J. Lyriochlamys faujasi (GMI 3601), external view; K. Chlamys sep-
templicata (GMI 3401), right valve in external view; L. Lyriochlamys dentata (GMI 3402), in external view; M. indeterminate (GMI 3701), internal mould of left valve; N. Mytilus sp.
(GMI 403), internal mould of left valve; O. Modiolus sp. A (GMI 1607), internal mould of left valve; P. Camptonectes virgatus (GMI 3504), left valve in external view; Q. Dhondtichlamys
subarata (GMI 3603), left valve in external view; R. Dhondtichlamys subarata (GMI 3604), right valve in external view; S T. Biradiolites suecicus (GMI 4001), attached valves in top
and lateral views; U. Icanotia grosseplicata (GMI 1605), internal mould of right valve; V. Pectinidae indet. (GMI 3702), left valve in external view.
32 A.M. Sørensen et al. / Cretaceous Research 33 (2012) 21e41

Fig. 9. Bivalves from Ivö Klack; all specimens natural size. A. Panopaea regularis (GMI 1706), internal mould of right valve; B. Barbatia sp. (GMI 1710), internal mould of right valve; C.
Goniomya sp. (GMI 1604), internal mould of right valve; D. Isognomon sp. (GMI 1709), internal mould of left valve; E. Tellinidae? indet. (GMI 1712), internal mould of left valve; F G.
Laternulidae sp. (GMI 1609), internal mould of articulated shell (left valve) in external and dorsal views; H. Barbatia? sp. (GMI 1714), internal mould of right valve; I. Arca? sp. (GMI
1610), internal mould of right valve; J. Isognomon sp. (GMI 1606), internal mould of left valve; K. Arcomya sp. (GMI 1608), internal mould of left valve.
 A.M. Sørensen et al. / Cretaceous Research 33 (2012) 21e41
auricles well differentiated, with anterior one being slightly
smaller; no gapes; outer surface with radial ribs and concentric
growth lines.
32. Ctenoides sp. (Fig. 7I)
Medium sized (49 mm in height); outline oval, height exceeding
width; beak acuminate and anterior margin straight; outer surface
with concentric growth lines. Only a single internal mould avail-
able; no dentition seen.
33. Limatula semisulcata (Nilsson, 1827) (Fig. 7D)
Small sized (up to 26 mm in height); outline oval, height exceeding
width; strongly inflated, hinge edentulous, auricles small and sub-
equal; no gapes; outer surface with fine radial ribs, most conspicuous
towards valve centre and absent from posterior and anterior ends.
34. Limea sp. (Fig. 7N)
Small sized (up to 30 mm in height); outline suborbicular or
ovate, height exceeding width; hinge margin relatively short;
hinge with series of short denticles on either side; auricles well
differentiated and appearing larger than in L. ovata (see above);
no gapes; outer surface with radial ribs; crenulated shell margin.
35. Pseudolimea sp. (Fig. 7M)
Small sized (13 mm in height); outline obliquely ovate to
orbicular; gibbous; beak near centre of moderately long cardinal
area, ligament pit broad; outer surface with numerous concentric
growth lines and radial ribs. Only a single valve available.
Gryphaeidae
36. Vultogryphaea laciniata (Nilsson, 1827) (Fig. 4AeD)
Medium sized (up to 45 mm in height); outline rounded to oval;
left valve cup shaped, outer surface with concentric growth lines
and few strong radial ribs with tendency to form hollow spines;
attachment area small; right valve as thin lid, slightly concave, with
concentric growth lines; shell typically exogyrine and coiled,
resulting in narrow ligamental and subligamental areas; commis-
sural shelf well-developed, with chomata dorsally.
Species here interpreted to have been cemented during juvenile
stage and later becoming free-lying, as based on very small
attachment area, morphological differences between valves and
relatively thin-shelled and concave left valve.
37. Gryphaeostrea canaliculata (J. Sowerby, 1813) (Figs. 4EeF, UeV
and Fig. 5EeF)
Medium sized (up to 42 mm in height); outline oval, height
exceeding width; left valve cup shaped, with small opisthogyrate
beak; right valve forming thin lid, slightly convex or concave and
much smaller than left valve, leaving wide free margin on left valve;
ligament area triangular; first opisthogyrate and later straight;
resilifer broader than bourrelets; commissural shelf well-
developed, in some cases with subligamental gutter without cho-
mata; outer surface of both valves with concentric growth lines,
becoming squamate at flanks; attachment area large.
38. Pycnodonte vesicularis (Lamarck, 1806) (Fig. 4IeL, ReT)
Medium sized (up to 75 mm in height); outline round to
obliquely oval; right valve forming weakly convex or concave lid;
33
ligament area normally short, straight and triangular, with bour-
relets and resilifer of near-equal size; hinge line usually straight and
short; commissural shelf well-developed with chomata restricted
to small, commonly fairly deep, gutters on both sides of hinge;
outer surface with growth lines near commissure; shell with
vesicular structure.
39. Amphidonte haliotoideum (J. Sowerby, 1813) (Fig. 4OeQ)
Medium sized (up to 70 mm in height); outline oval to comma
shaped; commissural shelf well-developed in both valves, with
chomata along entire margin; left valve deep, with a spirally curved
umbonal region; right valve flattened convex or concave with
a curved umbonal region; outer surface with smooth growth lines;
attachment area relatively large, in some cases, whole left valve
cemented to substrate.
40. Ceratostreon sp. (Fig. 4G, M and N)
Medium sized (up to 41 mm in height); outline elongate to
oval; left valve cup shaped with umbonal region forming larger
end; commissural shelf well-developed with chomata dorsally;
outer surface with smooth growth lines, plus keel rising into
prominent spines; attachment area relatively large; only left
valves available.
41. Exogyra? sp. (Fig. 5I and J)
Medium sized (38 mm in height); outline elongate, oval; valve
cup shaped with umbonal region forming larger end; twisted beak;
ligament area triangular, resilifer broader than bourrelets;
commissural shelf poorly developed; no chomata visible; outer
surface with smooth growth lines plus a keel rising from beak to
halfway down shell. Only a single right valve available.
42. Hyotissa semiplana (J. de C. Sowerby, 1825) (Fig. 4H)
Large sized (up to 90 mm in height); outline variably lobate;
right valve flat to slightly convex, with well-developed commis-
sural shelf, chomata present along entire shelf; adductor muscle
scar reniform, with both ends well rounded. Only right valves
available.
Ostreidae
43. Acutostrea incurva (Nilsson, 1827) (Fig. 5D, M and N)
Medium sized (up to 80 mm in height); outline variable, but
mostly elongate, straight or, more commonly, curved; ligament
area in left valve acuminate, high triangular, resilifer broader than
bourrelets; commissural shelf with rounded subligamental gutter
with chomata, and both gutters and chomata well-developed
from hinge to adductor muscle area, but effacing ventrally; outer
surface of both valves with concentric undulating growth lines,
becoming squamate in last stages of growth; attachment area
normally small.
44. Rastellum diluvianum (Linnaeus, 1767) (Fig. 5AeC and G)
Large sized (up to 160 mm in height); outline elongate, occa-
sionally crescentic to triangular-crescentic; shells with crescentic
outline have convex anterior and concave posterior flanks, tending
to be vertical to commissural plane; chomata present along entire
commissure; adductor muscle scars large and subtriangular to
comma shaped; ligament area broad and straight to opisthogyrate;
outer surface of both valves with numerous branching costae,
34 A.M. Sørensen et al. / Cretaceous Research 33 (2012) 21e41
turning into strongly plicate shell margin, forming an interlocking,
acutely angled, zigzag commissure; posterior shell margin not
always plicate; attachment area relatively large, commonly occu-
pying entire posterior flank.
Trigoniidae
45. ‘Trigonia’ sp. (Fig. 6E, I and J)
Internal moulds and external casts only; of small size (up to
27 mm in height); outline triangular to suboval; marginal carina
prominent; two symmetrical, striated teeth visible in mould;
outer surface with concentric ribs on anterior part, crossed by
radial ribs in median portion of shell; posterior part with radial
ribs.
Cardiidae
46. Cardiinae indet. (Fig. 6AeD)
Internal moulds only; of small size (up to 27 mm in height);
outline subtrigonal to suboval; both valves evenly inflated; umbo
prosogyrate, turned forwards; posterior margin obliquely trun-
cated; pallial line visible along entire margin; no ornament or
dentition seen.
47. Granocardium? sp. (Fig. 6K and L)
Internal mould only; one specimen available, of medium size
(40 mm in height); outline subtrigonal to suboval; both valves
evenly inflated, posterior margin obliquely truncated; umbo pro-
sogyrate, beak relatively straight; outer surface with radial ribs and
concentric growth lines; no dentition seen.
Tellinidae
48. Tellina? sp. (Fig. 5H)
Internal moulds only; small sized (up to 14 mm in height);
outline elongate, suboval; umbo positioned almost centrally, beaks
orthogyrous; siphonal gape posteriorly; outer surface with
concentric growth lines; no dentition seen.
49. Tellinidae? indet. (Fig. 9E)
Internal moulds only; of medium size (up to 30 mm in height);
outline elongate, suboval; umbo positioned almost centrally,
siphonal gape posteriorly; outer surface with concentric growth
lines; no dentition seen.
Arcticidae
50. Arctica sp. (Fig. 6Q, T and U)
Internal moulds only; of medium size (up to 48 mm in height);
outline subtrigonal to suboval; umbo prosogyrate, both valves
evenly inflated; posterior margin obliquely truncated; pallial line
visible along entire margin; hinge of heterodont type; no ornament
preserved.
Icanotiidae
51. Icanotia grosseplicata (Lundgren, 1894) (Fig. 8U)
Internal moulds only; of medium size (length up to 70 mm);
outline elongate; both valves equally weakly inflated, umbo posi-
tioned anteriorly; siphonal gape posteriorly; anterior end much
narrower than posterior; outer surface with concentric growth
lines and five prominent radial ribs posteriorly; no dentition seen.
Hiatellidae
52. Panopaea regularis (d’Orbigny, 1849) (Fig. 9A)
Internal moulds only; of large size (up to 68 mm in height); outline
elongate, suboval; umbo positioned almost centrally; siphonal gape
posteriorly; pallial line visible along entire margin, with wide pallial
sinus posteriorly; no dentition or ornament preserved.
Radiolitidae
53. Biradiolites suecicus (Lundgren, 1870) (Fig. 8S and T)
Medium sized (up to 40 mm in height); attached valve conical,
free valve forming lid; attached valve straight or arched, with no
teeth and posterior ligament; outer surface with longitudinal
ornament and growth lines.
Pholadomyidae
54. Goniomya sp. (Fig. 9C)
Internal moulds only; of medium size (length up to 56 mm);
outline elongate to suboval; both valves equally weakly inflated,
umbo positioned anteriorly; siphonal gape posteriorly; beaks
almost orthogyrous; outer surface with concentric growth lines; no
dentition seen.
55. Arcomya sp. (Fig. 9K)
Internal moulds only; of medium size (length up to 75 mm);
outline elongate; both valves equally weakly inflated, umbo posi-
tioned anteriorly; siphonal gape posteriorly; pallial line visible
along entire margin; no dentition or ornament preserved.
Laternulidae
56. Laternulidae sp. (Fig. 9F and G)
Internal moulds only; of large size (up to 44 mm in height);
outline elongate, oval; umbo positioned almost centrally; siphonal
gape posteriorly; deep holes, positioned dorso-posteriorly probably
reflect internal umbonal plate; pallial line visible along entire
margin, wide pallial sinus posteriorly; no dentition or ornament
seen.
Poromyidae
57. Liopistha sp. (Fig. 6O)
Internal moulds only; of small size (up to 28 mm in height);
outline triangular to suboval; anterior end shorter than posterior;
outer surface with coarse radial ribs, except on dorsal part of
posterior slope; no dentition seen.
Indeterminate
58. GMI 1715 (Fig. 7J)
Internal mould only; only a single specimen known, of medium
size (length 42 mm); outline elongate; valve inflated, umbo posi-
tioned anteriorly; siphonal gape posteriorly; pallial line visible
along entire valve margin; no dentition or ornament seen.
59. GMI 2602 (Fig. 5K and L)
Internal mould only; only a single specimen known, of small size
(14 mm in height); outline subcircular; valve weakly inflated, umbo
positioned anteriorly; pallial line visible along entire valve margin;
no dentition or ornament seen.
 A.M. Sørensen et al. / Cretaceous Research 33 (2012) 21e41 35

60. GMI 3501 (Fig. 8A)
Internal mould only; only a single valve known, of medium size
(46 mm in height); outline drop shaped; valve cup shaped, with
portions of auricles preserved; outer surface with rounded radial
ribs and riblets, broad intercostal intervals with radial lines and
well-developed, undulating concentric growth lines. Clearly a pec-
tinid, but generically and specifically indeterminate.
61. GMI 3701 (Fig. 8M)
Internal mould only; only a single specimen known, of small
size (length 29 mm); outline mytiliform; beak more strongly
acuminate than in Mytilus (see above); valve inflated; ventral shell
margin straight and flattened; no dentition or ornament seen.
5. Discussion
The bivalves constitute the most diverse and abundant faunal
group which inhabited the rocky shore at Ivö Klack, being repre-
sented by just over 60 species. This diversity is high in comparison
with other Late Cretaceous rocky shore bivalve assemblages from
Germany, the Czech Republic, New Zealand and Mexico (Table 3).
The late Cenomanian e early Turonian fauna from the Czech
Republic was studied with respect to cementing species found on
boulders; fifteen taxa have been recorded (Nekvasilová and Zítt, 

Table 3
The bivalve fauna from four Late Cretaceous rocky shore localities, and their modes of life as interpreted here. All species present are suspension feeders. # indicate presence at
 1988; Lescinsky et al.,
Ivö Klack, C ¼ cemented, By ¼ byssally attached, Bo ¼ boring, I ¼ infaunal, F ¼ free-lying (based on Pietzsch, 1962; Crampton, 1988; Nekvasilová and Zítt,

1991; Zítt,  et al., 1997).
1992; Johnson and Hayes, 1993 and Zítt

Species Germany Cenomanian Mexico CampanianeMaastrichtian Czech Republic Late New Zealand
CenomanianeEarly Turonian Santonian
Opis bicornis I
Gastrochaena ostreae Bo
Lima reichenbachi By
Pecten laminosus By
Pecten acuminatus By
Pecten elongatus By
Neithea quinquecostata # F
Neithea digitalis ? ?
Spondylus striatus C
Spondylus hystrix C
Lopha carinata C C
Rastellum diluvianum # C C
Pycnodonte vesicularis # C
Ostrea? cf. hippopodium # C
Exogyra lateralis C
Exogyra sigmoidalis C C
Amphidonte haliotoideum # C C
Exogyra reticulata C
Hyotissa semiplana C
Ostrea operculata C
Gryphaeostrea cf. canaliculata C
Spondylus omalii C
Ostrea sp. # C
?Amphidonte sp. C
Lyriochlamys cf. traskii By
Spondylus sp. # C
?Atreta n. sp. C
Coralliochama orcutti C
Oyster sp. A C
Atreta? sp. 1 C
Atreta? sp. 2 C
Spondylus sp. # C
Arca sp. By
Cucullaea sp. By
Modiolus sp. # By
Pteriidae sp. By
Isognomon sp. # By
Inoceramus sp. By
Oxytomidae sp. By
Chlamys sp. # By
Camptonectes sp. # By
Regalilima cf. marlburiensis By
Ostreidae sp. C
Trigonia aff. marwicki I
Pterotrigonia waitangiensis I
Psammobiidae sp. I
Bivalvia sp. ?
Total number of species 16 6 15 15
Total number of species in guilds I¼1 By ¼ 1 C ¼ 14 I¼3
Bo ¼ 1 C¼5 ?¼1 By ¼ 10
By ¼ 4 C¼1
F¼1 ?¼1
C¼8
?¼1
36 A.M. Sørensen et al. / Cretaceous Research 33 (2012) 21e41


1988; Zítt, 
1992; Zítt and Nekvasilová, 1996; Zítt  et al., 1997)
(Table 3). This number slightly exceeds that for Ivö Klack (14
cementing species), suggesting that the entire rocky shore bivalve
fauna in the Czech Republic may have been as diverse as at Ivö
Klack. The Cenomanian rocky shore bivalves from Germany
(Pietzsch, 1962) constitute the most diverse Cretaceous assemblage
described to date outside Ivö Klack; it comprises 16 species, here
assigned to five guilds, namely the cemented epifaunal, the epi-
byssate, the infaunal, the free-lying and the boring guild (Table 3).
The CampanianeMaastrichtian rocky shore fauna from Mexico is
represented by one byssate and five cementing species, indicating
that only species directly associated with the rocky shore were
described, similar to the Czech fauna (Lescinsky et al., 1991;
Johnson and Hayes, 1993). A Santonian fauna from New Zealand
(Crampton, 1988) includes 15 species of bivalve which are here
assigned to the cemented epifaunal, the epibyssate and the infaunal
guild. This guild structure indicates that both the subtidal, intertidal
and the adjacent sedimentary seafloor environments were
included in that study. However, that particular fauna is from the
Southern Hemisphere and has no species in common with any of
the other Late Cretaceous rocky shore faunas. Comparisons may
thus be difficult, although the diversity would be expected to be
similar at similar latitudes and in comparable climates. All the
Fig. 11. Pie-diagram showing the distribution of 60 bivalve species from Ivö Klack
Cretaceous faunas are less diverse in comparison to Ivö Klack as far
amongst the various guilds.
as number of species and number of guilds are concerned. The
German and Czech faunas, roughly 10 myr older, lived at similar
latitudes and in comparable climatic regimes within the same on the diversity at Ivö Klack. The fauna is interpreted as comprising
epeiric sea as that described from Ivö Klack. Furthermore, the a within-habitat, time-averaged assemblages (sensu Walker and
diversity of cementing species from the Czech Republic is similar to Bambach, 1971). According to Kidwell (1998, 2002), the relative
that at Ivö Klack, which indicates that at least these two European abundance of within-habitat, time-averaged assemblages is slightly
rocky shores may have hosted a comparable species diversity as higher than their modern counterparts. The much higher bivalve
recorded from Ivö Klack. Thus, it is most likely that the lower diversity at Ivö Klack is, however, interpreted as representing
diversity of other Late Cretaceous faunas can be explained by a genuine picture.
taphonomic processes, such as mechanical destruction, or by The cementing and epibyssate guilds at Ivö Klack are particu-
incomplete taxonomic assessment of taxa not directly cemented to larly important and comprise 61 per cent of all species recognised
the rocky shore. (Fig. 11). The guild structure provides a source for interpretation of
The species diversity among modern rocky shore bivalve faunas the former environment along the rocky shore and is a way to
from different latitudes is low in comparison to the Ivö Klack tackle palaeoecological aspects, independent of the number of
assemblages, even when infaunal species inhabiting associated specimens of each species preserved and, thereby, independent of
sediment are included. The highest bivalve diversity is found at preservational and collection biases. The high number of species in
Otranto in the southern Adriatic Sea (Italy), where 33 subtidal the attached guilds can probably be ascribed to the turbulent
species have been counted (Fig. 10) (Terlizzi et al., 2003). The high conditions along the steep coast, where it was advantageous to
diversity at Ivö Klack may represent the cumulative effect of be attached firmly to the substrate. Cementation provides the
generations of bivalves accumulated in the associated 22e24 m strongest attachment and all of the commoner species found at
thick carbonate succession along the rocky shore. The accumulation Ivö Klack belong to this guild, indicating relatively strong turbu-
of many generations of bivalves will lead to a higher total diversity lence along the shore (Table 1). The high number of species in the
compared to any snapshots of faunas from similar modern settings, shallow infaunal guild demonstrates that life just below the sed-
but, on the other hand, large taphonomic losses of aragonitic imentewater interface was attractive, with good water circulation
species and fragile, thin-shelled species have had a negative impact and oxygenation, but lower energy levels, precluding erosion of

Fig. 10. The bivalve fauna from four modern rocky shore localities and their positional relation to the substrate.
 A.M. Sørensen et al. / Cretaceous Research 33 (2012) 21e41
Fig. 12. Reconstruction of the bivalve fauna from Ivö Klack in life positions on and along the rocky shore.
shells or uprooting and destruction of the commonly thin-shelled
valves. The water depth must thus have been at least a few
metres, so as not to be affected by the relatively strong turbulence
along the shore. The distribution of the bivalves amongst the guilds
is similar to other Late Cretaceous bivalve faunas, with the majority
of species belonging to the cementing and bysally attached guilds
(Table 3). The Late Cretaceous guild structure is similar to that
known for modern rocky shores where the intertidal zones are
included in the various studies (Fig. 10). Only the Italian bivalve
community differs with a predominance of infaunal species,
probably because only bivalves from the subtidal zone were studied
and the most turbulent environment excluded. The similarity
between the Late Cretaceous and modern guild structures places
the bivalve fauna from Ivö Klack as a within-habitat, time-averaged
assemblage and thereby documents it as part of a true rocky shore
community.
The high species diversity and the wide variety of bivalve shell
morphologies document an environment along the rocky shore
with a variety of habitats occupied by species with different
attachment strategies. The preference of bivalves for different
habitats is based on their resistance to turbulence and on compar-
ison with modern relatives (Fig. 12; Table 2). This palaeoecological
subdivision is, like the guild structure, independent of preservation
and collection biases. It is assumed that species which preferred
37
high-energy conditions were also able to live in lower-energy
settings, whereas the opposite can be ruled out. Habitat 3, the
relatively protected environment, is thus occupied by the most
diverse fauna, because species which lived in habitats 4 and 5 were
also able to cope with habitat 3, following the above assumption.
Some species interpreted to have been present in a certain habitat
due to their capability to resist wave energy may actually not have
been present, due to biological factors such as competition for space
and species interactions. The preferred habitats of bivalves have
been interpreted in order to obtain an overview of their distribution
along the rocky shore. This shows that bivalves were all able to
inhabit the rocky shore environment and thus were not subse-
quently swept in from adjacent environments, which is supported
by the well-preserved fossils and the mainly angular nature of shell
fragments, indicative of limited or no transport (Surlyk and
Sørensen, 2010).
A well-developed epifaunal zonation, comprising three zones, is
preserved on gneiss boulders and hummocks (Surlyk and
Christensen, 1974). The highest zone is dominated by S. labiatus,
commonly the sole fossil found in this zone, which occupies the
uppermost part of the vertical sides of boulders and hummocks. The
next lower zone is dominated by A. haliotoideum and A. stobaei; this
represents the more or less vertical sides of boulders. Oysters are able
to live and feed in suspension-rich waters due to their self-cleaning
38 A.M. Sørensen et al. / Cretaceous Research 33 (2012) 21e41

mechanism. The closer the vertical wall was to the seafloor, the more
sediment was brought into suspension by waves. Many spondylids
are also found in this zone, indicating that sediment in suspension was
not the causal zonation factor between the two upper zones. The
difference in bivalve assemblages of the two highest zones probably
reflects variable tolerance to turbulence and competition for space.
The lowermost zone is dominated by serpulids and occupies the
overhanging lower part of the boulders, where energy levels were low
and light poor (Surlyk and Christensen, 1974; Sørensen and Surlyk,
2010). The absence of photonegative bivalves in the lowest zone
may be due to attachment strategies. Most photonegative bivalves are
nestlers and fissure dwellers which normally are attached by a byssus;
this means that they will not be preserved in situ.
Oysters are the commonest invertebrates at Ivö Klack; they
played a significant ecological role in this environment. Many of the
oysters have a high density of cementing and boring fauna which
made them important for the overall biodiversity by producing
small-scale habitats for encrusting and boring fauna (Fig. 13). Their Fig. 14. Right valve of Amphidonte haliotoideum encrusted by a large number of
stationary shells formed a stable substrate for small epifaunal conspecific juveniles.

species and many of these probably benefitted from the strong

feeding currents of the oysters (Sørensen and Surlyk, 2010). The
oysters probably reduced competition for space by occupying
different substrates, and the two dominant oysters, R. diluvianum
and A. haliotoideum had different life positions. The medium-sized
A. haliotoideum is found encrusting hummocks and boulders by the
whole of the left valve, in close proximity with conspecific indi-
viduals and even onto them (Fig. 14). The large-sized, thick-shelled
R. diluvianum is commonly found grown together and not
encrusting boulders (Fig. 15). Their heavy shells and relatively small
attachment scars would probably make it impossible for them to be
attached to boulders where they would have been sitting at an
almost right angle to the substrate in the turbulent environment.
They are thus interpreted as having constructed oyster banks on
the seafloor immediately adjacent to the rocky shoreline. The
preference for different substrates would have minimised compe-
tition for space, as high densities suggest.
Rudistid bivalves were widely distributed during the Late
Cretaceous and Biradiolites suecicus from Ivö Klack represents their
most northerly occurrence known, together with Canadian (Sas-
katchewan) records (Caldwell and Evans, 1963). These radiolitids
are small sized, less than 30 mm in diameter, compared to taxa
from the Maastrichtian of Peru which reached diameters up to
600 mm, probably as a result of symbiosis with zooxanthellae
(Philip and Jaillard, 2004). The small size and paucity of rudistids at
Ivö Klack is probably due to the northern location where symbiosis
with zooxanthellae may have been reduced or altogether absent,
due to lower water temperatures and illumination. Zooxanthellate,
non-reef forming corals are known from Ivö Klack, indicating that
the average water temperature must have been at least 18  C
(Sørensen and Surlyk, 2011). It is impossible to state whether or not
the rudistids lived in symbiosis with algae, but both zooxanthellate
corals and rudistids are rare in comparison to tropical settings,
indicating that the cooler waters were not favourable for these taxa
or their potential symbionts.
The byssate bivalve genera Mytilus and Barbatia are both
successful on rocky shores today; their record extends back to the
Late Jurassic and Triassic, respectively. The earliest known occur-
rence of Mytilus in association with ancient rocky shores is from the
late Oligocene of Oregon and that of Barbatia from the Maas-
trichtian of Alabama (Miller and Orr, 1988; Bryan, 1992; Johnson
and Baarli, 1999). Their presence on the early Campanian rocky

Fig. 13. Right valve of Rastellum diluvianum, showing a high density of encrusting fauna.
 A.M. Sørensen et al. / Cretaceous Research 33 (2012) 21e41
Fig. 15. Nine valves of Rastellum diluvianum cemented together, illustrated in two views. The oysters grew upon each other and constructed banks along the rocky coast. Their
common growth perpendicular to the substrate and their relatively small attachment areas can be seen on the specimens in the upper part of the figure. Living orientation of the
specimens cannot be inferred. LV ¼ left valve; RV ¼ right valve; ? ¼ unidentified valve.
shore at Ivö Klack thus documents the earliest example of both
Mytilus and Barbatia in association with such an environment.
The extent to which the bivalves described here are represen-
tative of the original fauna is difficult to estimate, since the
aragonite-shelled taxa are preserved as internal/external moulds
only and are mainly represented by few specimens of each species.
This indicates that the taphonomic loss of this group is high. The
ratio between aragonitic and calcitic bivalve species (i.e, 19 vs 37),
indicates a reasonable preservation potential for aragonitic species,
since it has been shown that the majority of bivalve species from
Ivö Klack were attached and epifaunal in the turbulent environ-
ment and most epifaunal species are calcitic (Heinberg, 1999;
Hautmann, 2006). Thus, a higher number of calcitic species is to
be expected in all cases. The preserved bivalve fauna is accordingly
considered to be representative of the bivalve community living on
the rocky shore where the intertidal, the subtidal and the associ-
ated skeletal sands are included in this environment.
6. Conclusions
 The rocky shore bivalve fauna from Ivö Klack is mainly well
preserved and, with just over 60 species on record, it is more
diverse than any other Cretaceous and modern rocky shore
faunas described to date. The high diversity most likely is
a genuine reflection of the fauna living along the shore at any
time and can only in part be explained by the cumulative effect
of generations of bivalves; taphonomic loss would probably
have equalised this.
 Seven guilds are recognised; 59 species are referred to these.
The cementing and epibyssate guilds comprise the largest
number of species due to the advantage of being firmly
attached to the substrate in a turbulent, high-energy environ-
ment. The high number of species in the shallow infaunal guild
39
indicates good water circulation but rather limited energy
levels, precluding erosion, uprooting and destruction of the
commonly thin-shelled valves. This distribution of bivalve
species between the guilds is similar to what has been
described for other Late Cretaceous rocky shore communities
and what is known from modern rocky shore, with a predom-
inance of attached species.
 Five habitats are recognised, representing different hydrody-
namic conditions and light penetration (illumination levels).
The distribution of bivalves between the habitats shows that all
species were able to live in the rocky shore environment which
included the intertidal, the subtidal and the associated loose
sediments, and thus were not subsequently washed in from
nearby settings. This is supported by the well-preserved fossils
and the mainly angular shell fragments which are indicative of
limited or no transport.
 Three zones of cementing fauna are recognised on large gneiss
boulders and hummocks. The distribution of encrusting
bivalves is interpreted to be the result of competition for space
and tolerance to water turbulence.
 Oysters were the commonest invertebrates along the shore and
are important for biodiversity as they offered abundant, small-
scale habitats for encrusting and boring fauna and thus
increase both the number of niches and faunal diversity. Some
of the species appear to have formed oyster banks along the
rocky shore, whereas others encrusted the large boulders,
thereby minimising competition for space.
 The small rudistid B. suecicus is, together with forms from
Saskatchewan (Canada), the most northerly known Late
Cretaceous occurrence of these bivalves. The small size and
paucity of rudistids indicate that the cooler water temperature,
compared to tropical settings, was not favourable for them or
their potential symbionts.
40 A.M. Sørensen et al. / Cretaceous Research 33 (2012) 21e41
 The successful rocky shore bivalve genera Mytilus and Barbatia
had their earliest known occurrence on rocky shores at Ivö
Klack.
 The bivalves thus formed the most important invertebrate
group living along the late early Campanian rocky shore at Ivö
Klack in terms of diversity, density and biomass. This contrasts
with modern rocky shores which are dominated by gastropods,
but this may be an artefact since the gastropod fauna at Ivö
Klack has suffered significant taphonomic losses directly linked
to their aragonitic shell mineralogy.
Acknowledgements
This study was funded by the Danish National Research Council
and the Carlsberg Foundation. Claus Heinberg is thanked for
constructive comments on an early manuscript version.
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