Radiolitid rudist colonisation strategies and biostrome development

in moderate-energy inner-platform environments

(Campanian, Bra Island, Croatia)
Dominik K. Hennhfer
a,
, Enric Pascual-Cebrian
a
, Tvrtko Korbar
b
, Wolfgang Stinnesbeck
a
, Stefan Gtz
a,1
a
Institute of Earth Sciences, Heidelberg University, Im Neuenheimer Feld 234, 69120 Heidelberg, Germany
b
Croatian Geological Survey, Sachsova 2, 10001 Zagreb, Croatia
a b s t r a c t a r t i c l e i n f o
Article history:
Received 21 June 2013
Received in revised form 17 March 2014
Accepted 19 March 2014
Available online 27 March 2014
Keywords:
Radiolitid rudists
3D reconstructions
Settlement
Palaeoenvironment
Population dynamics
During the Late Cretaceous radiolitid rudist bivalves were abundant calciers on Tethyan shallow-water carbon-
ate platforms. Previous studies have demonstratedvarious lowto high-energetic subtidal environments inwhich
vertically growing rudist bivalves lived. However, the precise mode of colonisation in moderate to higher ener-
getic environments remains unclear. The purpose of this study is to elucidate the attachment of rudist juveniles
and colonisation strategies on loose grain sediment using three-dimensional reconstructions and other quantita-
tive approaches. The present data demonstrate that small and thinly walled rudists colonised energetic environ-
ments over several generations. The main contributors to the biostrome, Distefanella, Bournonia, and
Pseudopolyconites, followed different strategies within the same ecosystem. For instance, cylindrical elevator
morphotype Distefanella built up a generally loose but anchored framework to resist sedimentation and wave
energy. In contrast, Bournonia and Pseudopolyconites followed a solitary strategy of settlement on any hard sub-
strate. Preferred growth direction affecting all genera was possibly linked to a predominant direction of water
currents.
2014 Elsevier B.V. All rights reserved.
1. Introduction
Radiolitid and hippuritid rudists were widespread benthic calciers
in Late Cretaceous tropical to sub-tropical carbonate platforms of the
Tethys Ocean and colonised various environments (Ross and Skelton,
1993; Gili et al., 1995; Sanders and Pons, 1999). Fewelevator rudist gen-
era were able to colonise shallowsettings with relatively high hydrody-
namic regimes (Sanders, 1998; Simone et al., 2003; Cestari, 2005). For
instance, the radiolitid genus Distefanella has been interpreted to have
tolerated high sedimentation rates and moderate to high-energy hydro-
dynamics (Cestari and Pons, 2007). A crucial factor for colonisation by
rudist elevator morphotypes was secure attachment on a hard sub-
strate. This attachment protected the rudist against wave energy and
avoids toppling and subsequent burying in the soft sediment (Skelton
et al., 1995). It is yet uncertain whether recruitment in rudist bivalves
happened either through primary or secondary attachment. Primary
attachment as veliger larvae can be observed in recent oysters (e.g.
Waller, 1981), whereas juvenile mussels settle as small dissoconchs
(Bayne, 1964). Korbar (2007) documented the attachment of rudist
juveniles to larger shells on the sea oor. Hennhfer et al. (2012)
described settlement of Biradiolites on a larger recumbent morphotype
of the genus Titanosarcolites in a relatively calm environment. In the
absence of an epibenthic hard substrate, smaller bioclasts (e.g. shell
fragments) represent an alternative to avoid settlement in the soft sed-
iment. Skelton et al. (1995) presented this option for hippuritid clusters
initiating on blankets of coarser bioclastic sediment. Little is known,
however, how rudist bivalves managed to secure and stabilise their
right valves during early stages of growth when inhabiting higher ener-
getic environments. The objective of the present analysis is to visualise
and describe these early juvenile growth strategies and to specify envi-
ronmental effects such as water energy and currents on the rudist com-
munity. Serial grinding and scanning of a decimetre scale sample
allowed three-dimensional reconstructions and quantitative analysis
of an in situ rudist biostrome. In the biostrome we examined the
radiolitid bivalve genera Distefanella, Bournonia and Pseudopolyconites
in a paucispecic rudist community. The material originates from
Middle Campanian subtidal carbonate platform deposits located on
the island of Bra off the coast of Croatia.
Palaeogeography, Palaeoclimatology, Palaeoecology 403 (2014) 8087
Corresponding author at: Institut fr Geowissenschaften, Universitt Heidelberg, Im
Neuenheimer Feld 234, 69120 Heidelberg, Germany. Tel.: +49 6221 544891.
E-mail addresses: dominik.hennhoefer@geow.uni-heidelberg.de (D.K. Hennhfer),
enric.pascual@geow.uni-heidelberg.de (E. Pascual-Cebrian), tvrtko.korbar@hgi.cgs.hr
(T. Korbar), wolfgang.stinnesbeck@geow.uni-heidelberg.de (W. Stinnesbeck).
1
Deceased.
http://dx.doi.org/10.1016/j.palaeo.2014.03.027
0031-0182/ 2014 Elsevier B.V. All rights reserved.
Contents lists available at ScienceDirect
Palaeogeography, Palaeoclimatology, Palaeoecology
j our nal homepage: www. el sevi er . com/ l ocat e/ pal aeo
2. Geological setting
During Late Triassic to Early Jurassic times, the Adriatic (Dinaridic)
Carbonate Platformformed the largest of several isolated carbonate plat-
forms in the central-southern Tethys (Jelaska, 2002; Vlahovi et al.,
2005). Lasting up to the Palaeocene, this platform allowed for the accu-
mulation of a several kilometre thick succession of mostly shallow ma-
rine carbonate (Jelaska, 2002). The island of Bra, located in Dalmatia
(Croatia), structurally represents an antiform as a part of the Cenozoic
fold-and-thrust belt of the External Dinarides (Korbar, 2009). The shal-
low water carbonate deposits cropping out on Bra are Late Cretaceous
to Eocene in age and were deposited on the Adriatic Carbonate Platform
andthe overlying forelandbasin(Gui andJelaska, 1990; Cvetko Teovi
et al., 2001; Moro et al., 2002; Steuber et al., 2005; Korbar et al., 2012).
Samples were collected near Povlja village on the northern coast of the
island of Bra (Fig. 1). East of the shing harbour of Tija Luka, Lower
to Middle Campanian limestone of the Puia Formation include the
Bra Marbles (Braki Marmori), Rasotica and Lovreina Members,
which crop out towards the headland of Tiji Rat (Fig. 1). The Puia
Formation is predominantly represented by carbonate packstone to
grainstone forming massive layers towards the base of the formation in
the Bra Marbles Member. This unit is commercially used as a famous
building stone. The mud and grain-supported limestone was deposited
in close geographic vicinity to rudist buildups below, probably above
the fair weather wave base (Cvetko Teovi et al., 2001). It alternates
with laterally extensive (N20 m) and decimetre to metre-scale thick
biostromes built up almost exclusively by rudist bivalve communities.
Radiolitids (Family Radiolitidae D'Orbigny 1847) are dominant and
assigned to the genera Distefanella, Biradiolites, Gorjanovicia, Bournonia,
Pseudopolyconites, Sauvagesia, Kuehnia and Durania.
The biostrome analysed here (Puia Formation, Lovreina
Member) is a paucispecic radiolitid community formed by Distefanella,
Bournonia, Pseudopolyconites, and Kuehnia. It was sampled from the
uppermost metre-thick rudist-bearing bed of the Tiji Rat section
(Fig. 2ac) and it overlies peloidal wackestone to packstone with
skeletal components (Fig. 2e, f). Centimetre-scale grainstone beds
were also observed (Fig. 2e). Distefanella radoicicae (=D. salmojraghii
sensu Cestari, 2008), Pseudopolyconites sp. and Bournonia excavata are
abundant and occur either as small bouquets or grewsolitarily, whereas
Kuehnia sp. (Kuehnia braciana?) is only locally present. Oriented
samples were extracted from the biostrome (Fig. 2a) and one sample
block was analysed in detail. Furthermore, reference samples were col-
lected from the underlying beds and laterally within the rudist-bearing
layer (Fig. 2di).
3. Visualisation method and application
The biostrome sample was processed with grinding tomography
allowing quantitative analysis of palaeontological and sedimentological
data as well as three-dimensional reconstructions. Pascual-Cebrianet al.
(2012a,b) provide a detailed description of the tomographic method
used here.
The oriented sample TR1 was embedded in an epoxy resin cast of
15 15 20 cm
3
and processed with semi-automatic serial grinding
andscanning. We usedanoptimisedG&NMPS 2-R300 precisiongrinding
machine for serial grinding and a custombuilt Epson Perfection V750 Pro
for data capture. We obtained 400 high-resolution digital image scans
(tomograms) with inter-slice spacing of 0.3 mm (vertical resolution).
The digital images were saved as TIFF les at a resolution of 1200 dpi.
Vertical resolution (space between tomograms) and horizontal resolu-
tion (pixel size on the tomogram) determine the dimension of a volu-
metric pixel. The data set covers a height of over 120 mm vertically,
including over one hundred individual rudists.
Tomograms were processed with OsiriX64 V.3.9 biomedical image
software (Ratib and Rosset, 2006), an open-source PACS workstation
(picture archiving and communication system). 3D reconstructions
were created to illustrate the distribution of rudists in the sediment
andto create virtual cuts through the sample volume. Involume render-
ing (VR) the reconstruction can be illustrated in false colour, enhancing
a certain area of the spectrum (e.g. colour of the shell), and reducing
Povlja
L
.

P
o
v
lja
6
4
0
7
.
0
0
0
4800.000
6
4
0
5
.
0
0
0
4
5 km
BOL
SUMARTIN
SELCA
POVLJA
DOL
SUPETAR
SUTIVAN
MILNA
N
S
W
1
6

3
0

4320
E
Foraminiferal Limestone
Sumartin Fm.
Milna Fm.
Sveti Duh Fm. Dol Fm.
Gornji Humac Fm.
Quaternary
0.5 km
CROATIA
Adriatic Sea
SPLIT
Fig. 1. Geological map of Bra Island and overviewof the major geological formations (modied after Steuber et al., 2005). Puia Formation crops out along the northern shore of Bra
(grey hatched area). Regional overview (top left) and detail of the section studied North of the village of Povlja (top right). (For interpretation of the references to colour in this gure
legend, the reader is referred to the web version of this article.)
81 D.K. Hennhfer et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 403 (2014) 8087
other areas resulting in transparency (e.g. sediment) (Figs. 3b, 4). Other
visualisation approaches, such as the illustration of shell attachment and
growth direction, allowed the creation of isosurface reconstructions
fromthe data set (Fig. 5). The smoothness of these isosurface reconstruc-
tions was enhanced by using MeshLab (V.1.3.2 64bit) mesh processing
software with a Laplacian lter algorithm and shells were coloured in
two different grey shades for better distinctness.
For quantitative evaluation, the basal part of the rudist buildup was
digitally measured for overall area coverage (percentage of horizontal
area covered by rudist right valves) and population density (number
of rudist specimens per area and genus) identied per tomogram over
a vertical extent of 120 mm(covering 400 layers at 0.3 mminterspace).
In total, over one thousand rudists have been recorded. Gtz and
Stinnesbeck (2003), Gtz (2007), and Hennhfer et al. (2012) have pre-
viously worked on quantitative evaluation of population dynamics in
dense rudist associations and provide further explanation and discus-
sion on the terminology used in this work.
4. Results
4.1. Facies description
The rudist biostrome analysed here as well as the underlying ne-
grained mud to grain-supported peloidalskeletal limestone beds
correlate with the top of the Lovreina Member (Puia Formation)
and, based on foraminiferal biostratigraphy, are Campanian in age.
Foraminifera detected in thin sections are assigned to Scandonea
mediterranea (Fig. 2g), Dicyclina schlumbergeri (Fig. 2h), and Accordiella
conica (Fig. 2i). Limestone underlying the rudist buildup contains ne-
grained peloids, orbitoids and miliolids, undened small gastropods,
and rare larger skeletal fragments (?Kuehnia sp.) as the major bioclasts
(Fig. 2e). Two types of matrix dominate in these lithologies: (1) peloid
grains in sparite matrix (peloidal grainstone), and (2) a carbonate
mud (wackestone to packstone) (Fig. 2df).
Within the biostrome the radiolitid rudist bivalves Distefanella,
Bournonia and Pseudopolyconites built up an in situ framework, which
partly trapped sediment and angular bioclasts such as fragmented
rudist shells in its interspaces (Fig. 3a). The different genera of radiolitid
rudists form a generally loose rudist biostrome. Closer observation
shows that monospecic Distefanella bouquets of fewer than 10 speci-
mens built up the framework. However, no articulated rudists (in situ
LV and RV) were found. The free valves (LV) are generally absent and
the in situ attached valves (RV) are bioeroded and/or mechanically
brokentowards the top. Body cavities are inlled with muddy sediment,
which is ner than the sediment surrounding the rudists (Fig. 3b, white
arrows). A few disarticulated free valves were found in the interstitial
sediment and one in a body cavity. Yet, the aragonitic cardinal
apparatus of these free valves is mostly missing and has been dissolved
early after deposition. In a single unidentied free valve preserved in
the body cavity of Bournonia the teeth are recrystallised to calcite
(Fig. 3c, black arrow). In macroscopic scale toppled right valves are pre-
dominantly North South orientated. These horizontally lying shells
were used as a hard substrate for newsettlers of the same species, lead-
ing to a lined-up growth of a new generation of Distefanella. Towards
Fig. 2. Schematic section with outcrop and microfacies photographs of the analysed biostrome; a schematic section at Tiji Rat including occurrences of rudist sub-families within the
succession and species in the top biostrome; b weathered surface towards top of the biostrome with transverse cuts of Distefanella; c South to North toppled right valve of Distefanella
(top centre) and N-S aligned growing Distefanella (top right) on weathered surface; d transverse section photomicrograph of Distefanella right valve near commissure, note the outer
shell layer (osl), the former aragonitic inner shell layer (isl), the body cavity lled with ne sediment (top right), and the coarser interstitial sediment; e scan image from the bed
underlying the rudist biostrome showing interbedding of wackestone/packstone (w-p) and grainstone (g); f photomicrograph of typical peloidal miliolid wackestone to packstone
underlying the radiolitid biostrome; g photomicrograph of abundant Scandonea mediterranea; h photomicrograph of Dicyclina schlumbergeri (right), a juvenile rudist right valve in
peloid matrix; i photomicrograph of Accordiella conica (top left).
82 D.K. Hennhfer et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 403 (2014) 8087
the top of the examined biostrome layer, the rudist diversity generally
decreases to a denser and apparently monospecic Distefanella associa-
tion (Fig. 2b).
4.2. Three-dimensional reconstructions
The volume rendering reconstructions (Fig. 4a) display the lower
90 mmof the examined limestone block (layers 001 to 300). This recon-
struction can be inspected by using virtual cuts of any direction. Fig. 4b
shows the biostrome under conditions of transparent sediment and the
right valves of Distefanella displayed under enhanced contrast. Most
right valves show a growth inclination towards the ENE (59). For
example, the isolated D. radoicicae shows an average inclination of 28
towards the ENE.
Fig. 5 illustrates the isosurface reconstructions of a single specimen
(D0, grey) of D. radoicicae during complete growth, fromthe early juve-
nile attachment on an adult (D1) of the same species, to its own adult
stage and death. Other rudist shells attached or in contact with D0
were also reconstructed. Distefanella specimen D0 (Fig. 5ae, in grey)
reaches a total height of 74 mm and a maximum thickness of
10.5 mm (dorso-ventral). The D0 juvenile settled and attached to the
shell in between two prominent costae of the adult. First growthfollow-
ing attachment shows that the juvenile inclined towards another adult
specimen (D2). When reaching D2, a change in growth direction
towards sub-parallel growth occurred with respect to D2. Fig. 5e
shows an abrupt correction of growth direction after outgrowing the
top of specimen D2. The rst growth stage without a direct inuence
of other shells is characterised by 14 mmof sub-vertical growth follow-
ed by a change to more inclined growth at anangle of about 45 towards
E, followed by up to 19 towards the NNE. At this approximately hori-
zontal stage the shell experienced heavy bioerosion and fracturing
after its growth (Fig. 5b, c). In the topmost growth stages of D0 growth
direction changed back to sub-vertical (Fig. 5b). During this stage juve-
niles of a later generationattached inthe interspace between two costae
of the strongly inclined adult shell of D0 (Fig. 5d). Fig. 5b and c indicates
that two of three specimens (D3, D5) outgrew their lateral position by
Fig. 3. Oriented scan images of the biostrome; a overview image (tomogram 187) with
predominant Distefanella shells (right valves, dark), body cavities are either inlledby sed-
iment or by secondary blocky calcite; b detail of image (a, top right square) and volume
rendering reconstruction; white line in the reconstruction represents the position of
tomogram 187, white arrows show ne sediment in body cavity; c transverse detail of
a sediment lled Bournonia right valve with angular bioclasts including a Distefanella left
valve (black arrow). (For interpretation of the references to colour in this gure legend,
the reader is referred to the web version of this article.)
Fig. 4. Volume rendering reconstructions under different colour spectrumsettings; a vol-
ume rendering reconstruction of the entire block (shells appear dark, sediment grey,
height 120 mm); b horizontal viewof inclined shells in the entire block at different geo-
graphic directions and under transparent sediment (block height 120 mm). (For interpre-
tation of the references to colour in this gure legend, the reader is referred to the web
version of this article.)
83 D.K. Hennhfer et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 403 (2014) 8087
Fig. 5. Isosurface reconstructions of a single Distefanella (D0) and adjacent specimens (D1, D2, D3, D4, D5) fromdifferent angles; a detail of juvenile settlers (D3, D4, D5) on D0 showing
close growth of juvenile rudists from above; b growth of D0 from attachment on D1, growth along D2 followed by fractured shell stage, to sub-horizontal stage and attachment of
new generation (view from W); c growth of D0 (view from S) shows initial vertical growth of D3, D4 and D5, subsequent horizontal growth parallel to D0 and close vertical growth;
d attachment of D4 between costae of D0 (detail view from S); e attachment of D0 between costae of D1, change of growth direction towards D2 followed by parallel growth and
direction change after outgrowing D2. Total height of D0 is 74 mm at maximum dorso-ventral thickness of 10.5 mm.
p
o
p
u
l
a
t
i
o
n

d
e
n
s
i
t
y

(
r
u
d
i
s
t
s
/
d
m

)
biostrome height (in cm)
a
r
e
a

c
o
v
e
r
a
g
e

(
%
)
80
20
40
60
0
20
40
60
2 0 3 4 5 6 7 8 9 10 1 11 12
0
Distefanella
Bournonia
Pseudopolyconites
a
b
total
Fig. 6. Area coverage (horizontal area claimed by all present rudist genera) and population density (living rudist specimens per area) during the rst 120 mmof vertical biostrome accu-
mulation; a area coverage is about 40% over the lower 105 mmand increases to N60%over the nal 15 mm; b overall populationdensity increases from29 specimens to 78 per 100 cm
2
,
Bournonia and Pseudopolyconites remain at low but stable values, while Distefanella increases population density from 22 to 55 specimens to 78 per 100 cm
2
.
84 D.K. Hennhfer et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 403 (2014) 8087
vertical growth. D4 is attached to a central position on top of D0 in be-
tween the costae (Fig. 5d) and does not showan initial vertical growth.
D3, D4 and D5 grew on top and parallel to D0 until reaching the nal
sub-vertical growing stage of D0 (Fig. 5a, d). Ultimately, settlers D3,
D4 and D5 followed the growth direction of D0 and continue growing
closely beside eachother (Fig. 5a). The three juvenile rudists grewclose-
ly, remaining within a single quadrant around D0. The scan images
show that in the sediment surrounding this bouquet, no other rudist
bivalve grew that inuenced growth within the biostrome.
4.3. Quantitative analysis
Three rudist species provided the biogenic buildup of the analysed
sample. D. radoicicae is the most prominent species (74% of all counted
rudist specimens), other main representatives are Pseudopolyconites sp.
(15%) and B. excavata (11%). Kuehnia sp. (K. braciana?) is an accessory
constituent in the biostrome; however, this genus only played a minor
role within the bioconstruction.
The horizontal area that is claimed by shells of different rudist gen-
era (area coverage) was evaluated for each layer and shows an increase
of area occupied by shells of living rudists during the rst 120 mm of
vertical carbonate accumulation (Fig. 6a). At the base of the analysed
limestone block, rudists occupied 33% of total horizontal area. Over
the rst 45 mm of growth, coverage steadily increases to N40% and
remains stable at an average of 45% for the overlying 50 mm. This
50 mm plateau phase is marked by minor uctuations. Between
90 mm and 105 mm, a decrease in coverage occurs from 45% to 33%.
Within the nal 15 mm (105 mm to 120 mm) values of coverage
increase again to N60%.
Over the examined vertical distance of 120 mm the overall
number of living rudist specimens per area (using a standardised
area of 100 100 mm) increases from initially 29 specimens to
78 per 100 cm
2
(Fig. 6b). However, this increasing tendency is
marked by three uctuations with a peak at 10 mm, 50 mm, and
100 mm height. Bournonia maintains an almost constant number of
specimens (min. 3, max. 10) over the analysed interval (Fig. 6b, long
dashed line), reaching a maximum number at 105 mm height. 11% of
all evaluated individuals belong to this species.
Pseudopolyconites shows a slight but constant increase in the num-
ber of specimens (Fig. 6b, dotted line) ranging from two at the base of
the biostrome to 18 near the top (95 to 100 mm). Pseudopolyconites rep-
resents the second strongest population within the biostrome rising to
approximately 15%. Distefanella (Fig. 6b, short dashed line) shows a
strong increase in the lowermost 10 mm, followed by stagnation in in-
dividuals between 10 and 30 mm height, and a second strong increase
between 30 and 50 mm in the biostrome. In the uppermost two thirds
of the examined block the populationdensity for D. radoicicae maintains
at stable levels of between 50 and 60 specimens per 100 100 mmarea,
except for a short phase of slightly reduced population density at about
15 mmbelowthe top of the block. All taxa examined showa decrease in
the number of specimens roughly between 50 mm and 90 mm. This
phase is characterised by the absence of uctuations. Synchronous in-
crease of population density and area coverage is observed at the top
of the analysed limestone block.
5. Discussion
5.1. Depositional environment
Growth of the biostrome initiated on a ne peloidal packstone
to grainstone (Fig. 2df) in a shallow open marine environment.
Centimetre scale changes between peloidal packstone and peloidal
grainstone (Fig. 2e) indicate episodically higher water energy, presum-
ably caused by winds or storms or tidal currents. The presence of larger
benthic imperforate foraminifers such as S. mediterranea (Fig. 2g)
indicates stenohaline conditions (Cvetko Teovi et al., 2001) as well
as water depths of b30 m. Cestari and Pons (2007) relate the genus
Distefanella to moderate to high-energy conditions and high sedimenta-
tion rates.
A general coarsening trend of bioclasts is observed from base to top
of the biostrome andinterpreted as an indicator for increasing water en-
ergy. In addition, free valves (left valves) are exclusively disarticulated
and very rare in the interstitial sediment. We therefore suggest that
shallowing water level or an increase in hydrodynamic conditions
favoured the settlement and growth of Distefanella associations. Alter-
natively, the size increase observed in bioclasts may result from
mechanical and biological erosion associated with a breakdown of
surrounding shell material.
Bioerosion was caused by clionid boring sponges and may have
played a major role in the detachment of free valves. However, the
absence of a ligament and the straight and thin teeth of Distefanella con-
tributed to an easy dislocation and transport of the free valve. Although
effects on shell orientation during early compaction cannot be fully ex-
cluded, the predominant shell inclination towards ENE suggests a major
current direction either towards ENE or WSW within the analysed
interval. In contrast, NS toppled cylindrical Distefanella right valves
identied on the top of the biostrome bed indicate a different current
direction. The discrepancy could derive from different causes: the ob-
served shell inclination may be a response to constant tidal currents,
whereas the toppling on top may have been storm related.
5.2. Settlement and growth
The absence of a stable substrate forced juveniles of Distefanella,
Bournonia, and Pseudopolyconites to follow different settlement strate-
gies. Within the sample examined, successful settlement was exclusive-
ly restricted to a hard substrate, such as the shell of an adult rudist, or,
on rare occasions, another biogenic hard substrate (toppledright valves,
shell fragments). Distefanella juveniles settled on vertical conspecic
right valves resulting in bouquets. Bournonia and Pseudopolyconites
juveniles, on the other hand, do not showa preferred substrate and set-
tled on shell fragments in the sediment as well as any adult rudist shell
available. As a result, no bouquets or other monospecic associations
have been recognised for these taxa (Fig. 3a). Their solitary growth
strategy and apparent independence from other specimens in the
analysed block suggests that Bournonia and Pseudopolyconites were
able to colonise medium to high-energetic environments. Distefanella,
on the other hand, is not considered to be a pioneer rudist that initiat-
ed settlement and gave opportunity for the settlement of other rudist
taxa. Rather, Distefanella juveniles preferably settled on secure locations
such as between two costae of conspecic rudist right valves. This strat-
egy resulted in monospecic bouquets of b10 individuals (Figs. 3b, 5d).
An active choice of substrate for juvenile attachment is therefore
assumed and was likely linked to biochemical markers. For instance,
bacterial lms on the surface of Mytilus edulis (Satuito et al., 1995), or
neurotransmitters and ammonia on Crassostrea (Fitt and Coon, 1992)
are known to attract juveniles in modern bivalve communities. The NS
alignment observed in the outcrop for toppled shells of Distefanella
and horizontally lying right valves served as a hard substrate for new
settlers of this taxon (Fig. 2c). Limited by the availability of hard sub-
strates within the ecosystemthis alignment may have led to a preferred
growth direction by lined up and vertically growing shells. However,
successful settlement of Distefanella juveniles on toppled adults of the
same species instead of other toppled rudist shells cannot be explained
by biochemical markers because the host shells were most likely not
alive as recovery growth of fallen shells has not been observed (e.g. de-
scribed in Skelton et al., 1995). The mesh reconstructions (Fig. 5) illus-
trate an isolated single specimen of D. radoicicae from settlement on
an adult right valve (D1) and its role as a settling ground (hard sub-
strate) for new juvenile (D3, D4, D5). They further demonstrate the
combination of both vertical growth and secure anchorage in a soft sub-
strate environment. Initial settlement in between the costae proved to
85 D.K. Hennhfer et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 403 (2014) 8087
be successful as it provided protection from strong currents. During
later growthstages the juvenile shell appears to have reactedto the like-
ly energetic environment by changing growth direction and searching
for contact to D2 (Fig. 5e). We interpret this second shell attachment
as an approach to increase stability. Interestingly, settling juveniles D3,
D4 andD5 showcomparable growth behaviour (Fig. 5a, b). After attach-
ment in depressions between costae these juveniles stay in close
vicinity or in direct contact with the adult shell, probably to secure
anchorage. Two laterally attached juveniles, D3 and D5, grew sub-
vertically at rst (Fig. 5b, c), probably to avoid turbid water close to
the sediment surface. Eventhoughthis wouldimply proximity to the sed-
iment surface, it seems plausible that at the time of attachment of D3 and
D5, the right valve of D
0
could not have been entirely embedded in sedi-
ment. Fig. 5bd illustrates that juveniles tried to reach the upper side of
the sub-horizontally growing right valve of D0, but are anchored to an
adult shell. During the latest stages of growth of D0, settlers D3, D4 and
D5 almost reached maximum diameters, but grew close to each other.
This growth strategy contrasts with that of Biradiolites mooretownensis
from the Campanian of Jamaica (Hennhfer et al., 2012), a rudist in
close taxonomic and morphologic relation to the present species. At
rst, juvenile settlers of bothtaxa growas vertical as possible, toavoidtur-
bid water by creating maximum distances to the substrate. However,
juveniles of Distefanella grew as dense small bouquets within a generally
loose associationwhile juveniles of B. mooretownensis avoidedcontact be-
tween individuals. The reason for this different growth behaviour of the
Jamaican rudist is seen in its decreased water energy and nutrient avail-
ability (Hennhfer et al., 2012). Bouquet or cluster growth reects the
availability of a secure attachment (Skelton et al., 1995). This interpreta-
tion is conrmed by the present data; however, the concept of growth
in high energetic regimes should be expanded. Thinly walled elevators
such as Distefanella grew as small multigenerational bouquets (Figs. 3b,
5) of few specimens. The sketch drawing (Fig. 7) is an approach to illus-
trate different recorded attachment localities. This framework increased
stability by deep anchorage in the sediment using previous generations
of the same species as a substrate, also during episodes of higher water
energy, in which the sediment could have been partially remobilised.
This radiolitid rudist association is therefore not entirely reliant on sedi-
ment supporting its vertical growth, as previously suggested for other
rudist genera (Ross and Skelton, 1993; Gili et al., 1995; Gili and Skelton,
2000) and could have allowed low superstratal growth. At present, we
cannot provide absolute data on growth rates and sedimentation rates;
however, vertical carbonate accumulation was presumably in the range
of a few centimetres (Steuber, 2000; Hennhfer et al., 2012) per year.
5.3. Population dynamics
Area coverage and population density provide valuable information
on ecologic pressure within an ecosystem(Gtz and Stinnesbeck, 2003;
Gtz, 2007; Hennhfer et al., 2012). For instance, the general increase of
area coverage and population density of the present paucispecic rudist
community illustrates a gradual increased use of available space. From
bottom to top, more successfully attached juvenile shells occupy more
space. Only Bournonia and Pseudopolyconites do not provide reliable
results as both are rare taxa in the association, opportunistically using
the framework, or space between bouquets of Distefanella for attach-
ment. The graph of overall population density (Fig. 6b) shows three
major peaks of similar character and spacing (10 mm, 50 mm,
100 mm). The coarse inter-shell sediment of the sample does not
allow explicit spat counting, but juveniles successfully attached to
adult right valves episodically and started growth. Comparable patterns
of population density and episodically increased numbers of rudists per
area have been documented for Biradiolites (Hennhfer et al., 2012) and
were there interpreted as seasonal attachment of spat in a monospecic
rudist population. Area coverage (Fig. 5a) in the present sample is sig-
nicantly lower than in other examined associations (about 40% as
compared to N85% in ?Radiolites, Gtz, 2007). This difference may result
fromadaptation to a different environment (i.e. substrate, water energy,
or food supply). Protected settings (e.g., B. mooretownensis biostrome,
Hennhfer et al., 2012) allowed for a denser growth and consequently
broader bioconstructions (thickets). The nal massive increase of area
coverage identied at 105 mmheight in the present sample is associat-
ed with an increase of population density which implies an increased
survival of attached recruits accompanied with a doubling of horizontal
area occupation. Changes in ecologic conditions may provide a possible
explanation for this juvenile bloom, possibly due to increased nutrient
supply and/or higher water energy. Alternatively, an increase in shell
fragments led to a higher availability of hard substrate for attachment.
The increase of available settling spots will therefore reect in the quan-
tity of successfully attached juveniles. Towards the top of the rudist-rich
limestone layer coverage further increases while other radiolitids de-
crease in number and disappear. As a result, a monospecic Distefanella
association is present towards the top of the unit as described in the
Fig. 7. Interpretation of Distefanella settlement (sketch) as multigenerational bouquets of less than 10 specimens; juveniles attach to conspecic right valves, either toppled or in life
position; the multigenerational bouquets offer sufcient support against wave energy and allow vertical growth into the water column.
86 D.K. Hennhfer et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 403 (2014) 8087
literature (Fig. 12 in Korbar, 2007). The nal massive increase of area
coverage (Fig. 6a) therefore marks the end of an ecological succession,
exemplied by the local demise of competing radiolitid genera
Bournonia and Pseudopolyconites.
6. Conclusions
The Late Cretaceous radiolitid biostrome from Bra, Croatia, was
deposited in a tropical and shallow marine setting under moderate hy-
drodynamic conditions. A shallowing trend is indicated by coarsening-
upward of bioclasts that favoured the initiation of paucispecic rudist
biostromes dominated by Distefanella, Bournonia and Pseudopolyconites.
The shells are preserved in growth position, nevertheless, water energy
and bioerosion most probably caused fracturing of a few valves. The
majority of these right valves are slightly inclined towards the ENE,
which is either the result of a predominant water or wave direction or
of different mechanical behaviour during early compaction.
Distefanella, Bournonia, and Pseudopolyconites followed different
strategies to compensate the paucity of stable substrate to allow secure
settlement. Distefanella formed bouquets of less than 10 specimens,
whichgrewon vertically growing adult right valves of a previous gener-
ation. Individuals of Bournonia and Pseudopolyconites are larger-sized
and occur solitarily. Multigenerational anchoring of bouquets signi-
cantly increased the stability of each specimen and allowed the rudists
to securely grow into the water column. Distefanella juveniles actively
chose their location for settlement. They attached to costae of adult con-
specic right valves, probably to secure attachment in early growth
stages. After attachment, juvenile Distefanella settlers were able to
change growth direction and to further attach to other adult shells of
the same taxon. This strategy of being attached to multiple individuals
certainly increased the stability and resulted in monospecic bouquets.
Quantitative analysis of the sampled bouquet from the Bra biostrome
revealed a loose association of individuals at the base, probably related
to the higher energetic regime. The simultaneous increase of population
density and coverage suggests a change in the ecologic conditions
favouring Distefanella over both Bournonia and Pseudopolyconites, ulti-
mately resulting in a monospecic and more dense rudist association.
The method of serial grinding and serial scanning used herein, com-
bined with digital 3D reconstructions, thus provides a valuable tool to
assist palaeoenvironmental reconstructions in rudist biostromes and
helps to reveal detailed insight in settlement strategies, growth, and
population dynamics.
Acknowledgements
We wish to commemorate our colleague and friend Stefan Gtz,
who sadly passed away on 30 July 2012. We are deeply grateful to
Peter W. Skelton (Milton Keynes) and an anonymous reviewer, who
both provided very helpful reviews and comments on an earlier version
of the manuscript. We would like to acknowledge the nancial support
given by Deutsche Forschungsgemeinschaft (Projekt GO 1021/3-2), by
Heidelberg University (Global Change and Globalization, Frontier
Innovationsfonds), by Ministerium fr Wissenschaft, Forschung und
Kunst BadenWrttemberg (Research Seed Capital), and the Croatian
Geological Survey.
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