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Geobios 44 (2011) 543–548

Original article
Entrapment, preservation and incipient fossilization of benthic predatory
molluscs within deep-water coral frames in the Mediterranean Sea§
Piégeage, préservation et début de fossilisation de Mollusques prédateurs benthiques
au sein d’un système corallien d’eau profonde en mer méditerranée
Lorenzo Angeletti *, Marco Taviani
ISMAR-CNR, UOS Bologna, 101, via Gobetti, Bologna, Italy
Received 1 September 2010; accepted 4 February 2011
Available online 12 October 2011

Abstract
Holocene Madrepora-Lophelia subfossil frames recovered at ca. 690 m from deep-water coral grounds south of Malta (Strait of Sicily, Central
Mediterranean Sea) were found to entrap shells of cnidarian predatory gastropods such as Babelomurex sentix, ‘‘Coralliophila’’ squamosa
(morphotype ruderatus) and architectonicids. This finding documents the capability of deep-water coral reefs to serve as taphonomic traps
eventually promoting the preservation of rare components of their original biota.
# 2011 Elsevier Masson SAS. All rights reserved.

Keywords: Deep-water corals; Taphonomy; Gastropods; Muricidae-Coralliophilinae; Architectonicidae; Mediterranean Sea

Résumé
Nous montrons que des systèmes coralliens d’eau profonde subfossiles (Holocènes) à Madrepora-Lophelia découverts à environ 690 m de
profondeur au sud de Malte (Détroit de Sicile, Mer Méditerranée centrale) ont capturé des coquilles de gastéropodes prédateurs de cnidaires, tels
que Babelomurex sentix, « Coralliophila » squamosa (morphotype ruderatus) et des architectonicidés. Cette observation illustre la capacité des
récifs coralliens d’eau profonde à servir de piège taphonomique, favorisant éventuellement la préservation d’éléments rares de leurs biotopes
d’origines.
# 2011 Elsevier Masson SAS. Tous droits réservés.

Mots clés : Coraux d’eau profonde ; Taphonomie ; Gastéropodes ; Muricidae-Coralliophilinae ; Architectonicidae ; Mer méditerranée

1. Introduction
Cold- (or deep-) water coral reefs (CWCs) are marine
habitats characterized by high levels of biodiversity (Freiwald,
2002; Freiwald et al., 2004; Roberts et al., 2006; Hovland,
2008). They can contribute to making spectacular carbonate
mounds, such as those along the NE European margin (e.g.,
Wheeler et al., 2007). CWCs occur almost worldwide,
including the semi-enclosed Mediterranean basin, which boasts
a superb, albeit discontinuous, paleontological documentation
§ submarine cementation is conducive to the lithification of the
Corresponding editor: Davide Olivero.
* Corresponding author.
E-mail address: lorenzo.angeletti@bo.ismar.cnr.it (L. Angeletti).
from the Miocene onwards (Taviani et al., 2005, and references
therein). In this basin, most of the extant CWC-bearing
assemblages are of the Pleistocene age and either crop out in
southern Italy and Rhodes (Di Geronimo et al., 2005; Titschack
and Freiwald, 2005) or are still submerged (Taviani et al.,
2005).
Both living and dead coral structures can provide niches to a
variety of benthic organisms (Freiwald et al., 1997, 2004;
Freiwald and Wilson, 1998). Biohermal frame growth and
related biostromal production may optimise entrapment, and
ultimately the preservation of vagile visitors or stable exploiter
visitors of such coral habitats. The recurrent process of early
intracoral fine-grained matrix resulting in carbonate rocks
(framestones) which encase both coral frames and skeletal

0016-6995/$ – see front matter # 2011 Elsevier Masson SAS. All rights reserved.
doi:10.1016/j.geobios.2011.02.004
544 L. Angeletti, M. Taviani / Geobios 44 (2011) 543–548

remains of associated macrofauna, mainly molluscs, followed

by brachiopods, echinoids and decapods.
Collections of pre-Modern coral frames and coral hard-
grounds obtained from various Mediterranean submerged
locations show that this taphonomic process is common.
Although the main coral entrappers are basically the same
basin-wide (i.e., the colonial scleractinians Lophelia pertusa
(Linnaeus, 1758) and Madrepora oculata Linnaeus, 1758, and
more rarely the solitary Desmophyllum dianthus (Esper, 1794)),
the entrapped organisms may differ from site to site (e.g.,
Remia and Taviani, 2005; Taviani et al., 2005, 2011). Benthic
gastropod shells belonging to Emarginula, Putzeysia, Alvania
and Amphissa as well as holoplanktonic thecosomatous
pteropods and heteropods are almost ubiquitous. Last glacial
Desmophyllum-bearing micritic hardgrounds dredged at
depths > 2000 m from the Malta-Siracusa escarpment encase
articulated Delectopecten vitreus (Gmelin, 1791) shells, a
bathyal pectinid, inside the coral calices (Taviani and
Colantoni, 1979). Buried latest Pleistocene Madrepora mounds
at ca. 350–400 m in the NE Tyrrhenian sea have been
documented by Remia and Taviani (2005) to enclose in their
weakly-cemented frames articulate brachiopods (Gryphus,
Megerlia) and vagile gastropods (Emarginula, Coralliophila).
By large, most of the ‘fossilized’ shells are not uniquely
related to deep-water corals. Cnidarian predatory-gastropods
Fig. 1. Study area. A. Map showing the location of RV Urania MEDCOR 25
(such as Muricidae-Coralliophilinae, Ovulidae, Epitoniidae,
station in the Strait of Sicily. B. 3D reconstruction of the escarpment based on
Architectonicidae; Robertson, 1967, 1970; Sabelli and Taviani, swath bathymetry data marking the South Malta CWC province from which the
1984; Bieler, 1993; Oliverio et al., 1997; Gittenberger, 2006; material examined in this article was obtained (white dot marks station
Kokshoorn et al., 2007; Reijnen et al., 2010) occur much more MEDCOR 25).
rarely in fossil taphonomic coral traps.
A spectacular case in point is discussed here, related to
Madrepora-Lophelia subfossil frames from deep-water coral BP, and as calendar year estimates (cal. years BP) after
grounds south of Malta (Strait of Sicily) entrapping shells of calibration; BP refers to 1950 AD.
extremely rare cnidarian-predatory gastropods. The area is the
CWC province recently described by Schembri et al. (2007) 3. Deep water coral taphonomic traps
and Freiwald et al. (2009). Location and main topographic
characteristics of the study area are presented in Fig. 1. The south Malta CWC province is a site where live
scleractinians (Lophelia, Madrepora, Desmophyllum, Dendro-
2. Material and methods phyllia, Javania, Caryophyllia) and other sessile invertebrates
such as octocorals (including Corallium), antipatharians
The study material is sourced from CNR cruise MEDCOR (Leiopathes), barnacles (Pachylasma) and hexactinellid
with RV Urania (December 2009), more precisely from sponges contributing to make an important and diverse hotspot
MEDCOR Station 25 (358 30.7960 Lat N-148 11.0760 Long E, settling a deep-water escarpment (Freiwald et al., 2009; Taviani
depth 690 m). Swath bathymetry data were acquired using a et al., 2011). Although lush live coral communities are well-
RESON 8160 multibeam echo-sounder with nominal sonar documented here, dead subfossil corals and coral hardgrounds
frequency of 30 kHz and angular coverage sector of 135 beams are equally common. During cruise MEDCOR we sampled
per ping at 18. Sub-bottom Chirpsonar profiles were obtained various examples of framestones and wackestones mostly
using a hull-mounted 16-transducer source with a sweep consisting of the colonial scleractinians Madrepora oculata
modulated 2–7 kHz outgoing signal equivalent to a 3.5 kHz and, subordinately, Lophelia pertusa, showing various degrees
profiler. Sampling was achieved using various equipment of patination by Fe-Mn oxides, bioerosion and cementation.
(corers, grabs, epibenthic hauls), depending upon the nature of MEDCOR Station 25 was particularly prolific in providing
the sea bottom. Samples were washed on-board in freshwater, samples of such carbonates. The screening of framestones and
dried and examined with the aid of magnifying lenses and hardgrounds produced various examples of benthic organisms
optical microscopes. One Babelomurex sentix (Bayer, 1971) entombed in these degraded coral frames often affected by early
shell and one Lophelia pertusa coral were 14C-AMS dated at diagenetic processes. Although some of the coral frames are
the Poznan Radiometric Laboratory, Poland. Radiocarbon only weakly cemented and retain no intracoral matrix, others
results (Table 1) are reported as raw 14C-ages given in 14C-yrs show more advanced lithification also at the expense of the
 L. Angeletti, M. Taviani / Geobios 44 (2011) 543–548 545

Table 1
Radiocarbon Lab-code ‘‘Poz’’ refers to the Poznan Radiocarbon Laboratory in Poznan, Poland. Radiocarbon years (14C-yrs) correspond to the true half-life.
Calibrated age ranges reflect the 95% probability window (2s-error) and were calculated with Calib 5.10, using the surface marine calibration curve MARINE 04 by
Hughen et al. (2004).
Sample Species Lab-code Radiocarbon age [14C-yrs BP] Calibrated age range (2s) [yrs cal BP]
MEDCOR25 Babelomurex sentix Poz-37811 732  73 659–804
MEDCOR25 Lophelia pertusa Poz-37812 628  68 560–696

muddy matrix (Fig. 2(A)). This process likely started during the
latter hundreds or thousands of years in the Holocene and is still
continuing. These potential fossils can be conveniently
categorized as occasional visitors or more rarely ecologi-
cally-consistent associates, i.e., basically ‘freezing’ in situ part
of the original ecosystem. The first group includes taxa such as
the gastropod Amphissa acutecostata (Philippi, 1844) and
decapod calcified remains (Fig. 2(B)). The second group
includes examples of vagile organisms inhabiting the coral reef
itself. In the specific case-study treated here we document the
inclusion in coral frames and hardgrounds of some rare
predatory gastropods (Fig. 3(A–J)). Coral frames can often trap
pelagic tests (Fig. 3(K)).
Blackened shells of the elusive Muricidae-Coralliophilinae
neogastropods Babelomurex sentix and ‘‘Coralliophila’’
squamosa (the deep-sea morphotype ruderatus Sturany,
1896, of the common subtidal-circalittoral nominal taxon
‘‘Coralliophila’’ squamosa (Bivona Ant. in Bivona And.,

Fig. 2. Coral framestone-hardground from MEDCOR 25. A. Large slab of coral framestone-hardground largely made up by Madrepora oculata and Lophelia
pertusa branches (and subordinate Stenocyathus vermiformis and Corallium rubrum) displaying different degrees of sediment infilling and lithification from absent to
advanced; note the patination by Fe-Mn oxides; scale bar = 5 cm. B. Detail of the same slab showing a blackened shell of Amphissa acutecostata (white arrow) almost
completely embedded within the lithified matrix); scale bar = 1 cm. C. Same slab showing a coarser fabric with encased shells of Heliacus alleryi (black arrow) and of
blackened ‘‘Coralliophila’’ squamosa (morphotype ruderatus: white arrow); scale bar = 1 cm.
546 L. Angeletti, M. Taviani / Geobios 44 (2011) 543–548

Fig. 3. Examples of shells encased in coral framestones and hardgrounds, all from MEDCOR 25. A, B. Architectonicid shell (cf. Discotectonica discus) cemented to
coral (Lophelia pertusa) framestone, white arrow points to the broken base of an unidentified scleractinian growing over the gastropod shell, black arrow points to the
broken base of Corallium rubrum. C, F, G. Shell of Babelomurex sentix (Muricidae-Coralliophilinae) welded to a Lophelia branch; note degradation, fouling and
blackening of the shell. D, E. Architectonicid shell (Heliacus alleryi) associated with a lithified matrix hosting pelagic molluscs and foraminifers. H. Loose
Babelomurex sentix shell. I, J. Shell of ‘‘Coralliophila’’ squamosa (morphotype ruderatus) intensely fouled by various Lophelia corallites. K. Degraded and
blackened ootheca of the epipelagic cephalopod Argonauta argo, infilled by lithified sediment. Scale bars = 5 mm.
 L. Angeletti, M. Taviani / Geobios 44 (2011) 543–548
1838): see Taviani et al., 2009) were observed cemented inside
Madrepora-dominated framestones and subordinated Lophelia
(Fig. 3(C, F–J)). Both taxa, unknown as fossils, are seldom
recorded in the literature (e.g., Oliverio, 1989; Giannuzzi-
Savelli et al., 2003; Oliverio and Gofas, 2006) and have been
recently recorded alive from these same CWC grounds by
Taviani et al. (2009). It is likely that they feed upon some
cnidarians associated with the CWC reefs but the precise
host(s) is unknown at present. One shell of B. sentix from
MEDCOR 25 provided a 14C-AMS age of 732  73 years BP
while one Lophelia branch from the same station held an age of
628  68 years BP (Table 1). Therefore, the present finding
documents that these rare gastropods consistently inhabit the
South Malta CWC province at least since the latest Holocene,
co-occurring with another member of the subfamily Corallio-
philinae, ‘Coralliophila’ richardi (P. Fisher, 1882) (predatory
on Madrepora and Lophelia; Taviani et al., 2009) of which
some dead shells were also found at MEDCOR 25 but not
entrapped in coral framestones or hardgrounds. Some of the
shells served as a substrate for coral settlement.
Besides Coralliophilinae, other predators of cnidarians have
been detected in our samples. Shells belonging to at least two
different species in the family Architectonicidae (Gastropoda,
Heterobranchia) were detected welded to lithified matrix in
coral framestones and hardgrounds (Fig. 3(A, B, D, E)). One
such shell is poorly preserved and its base is embedded in the
lithified matrix not allowing one to ascertain with confidence its
taxonomy, but possibly referable to Discotectonica discus
(Philippi, 1844). As for some of the Coralliophilinae discussed
above, the shell was also functional to later coral settlement.
Two shells belonging to Heliacus alleryi (Seguenza, 1876)
were also detected in the coral hardgrounds (Figs. 2(C) and
3(D, E)). Loose shells of these uncommon Mediterranean
Architectonicidae (Melone and Taviani, 1984) were relatively
common in MEDCOR 25, but their precise cnidarian preys are
unknown at present.
4. Conclusion
Deep water coral reefs may serve as efficient taphonomic
traps which may help the post-mortem preservation of
skeletonized organisms, potentially ensuring their fossilization.
A remarkable example of such is offered by the finding of pre-
modern Lophelia-Madrepora coral frames and hardgrounds at
the recently discovered South Malta CWC province. These
carbonates were found entrapping shells of exceedingly rare
deep-water Coralliophilinae (Babelomurex sentix, ‘‘Corallio-
phila’’ squamosa – morphotype ruderatus) and Architectoni-
cidae (cf. Discotectonica discus, Heliacus alleryi). This record
is a rare case documenting the incipient fossilization of
ecologically consistent CWC predator-host assemblages.
Acknowledgements
We are grateful to Captain, Crew and Colleagues onboard
RV Urania during MEDCOR mission. We thank A. Beu and an
anonymous referee for their constructive criticism. English
547
kindly revised by Derek Jones. Funding was partially provided
by CNR and HERMIONE project (contract no 226354) within
the 7th Framework Program of the EU. This is ISMAR-
Bologna scientific contribution no 1679.
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