Sclerobionts on Upper Famennian Cephalopods from the Holy Cross Mountains, Poland
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CORE Metadata, citation and similar papers at core.ac.uk Provided by Springer - Publisher Connector Palaeobio Palaeoenv (2011) 91:63–73 DOI 10.1007/s12549-010-0045-x ORIGINAL PAPER Sclerobionts on upper Famennian cephalopods from the Holy Cross Mountains, Poland Michał Rakociński Received: 16 June 2010 /Revised: 7 September 2010 /Accepted: 23 September 2010 /Published online: 27 October 2010 # The Author(s) 2010. This article is published with open access at Springerlink.com Abstract Encrustation by diverse sclerobionts is found on These oldest encrusters include tabulate corals (Galle and about 4% of upper Famennian cephalopods from the active Parsley 2005), bryozoans, edrioasteroids (Baird et al. 1989; Kowala Quarry, Holy Cross Mountains, central Poland. Frey 1989; Kácha and Šarič 2009), crinoids, cystoids These infested cephalopods are represented mostly by (Ganss 1937, vide Rakús and Zítt 1993), inarticulate clymeniid ammonites, but also include goniatitids and brachiopods (Gabbott 1999; Lockley and Antia 1980) and nautiloids. Sclerobionts on cephalopod shells include cornulitids (Gabbott 1999; Morris and Rollins 1971) found crinoids (represented by their holdfasts, about 57% of on nautiloids (mainly orthoconic). Silurian epibionts utilising encrusters), moulds of problematic worm tubes (less cephalopod hosts (orthoconic nautiloids) were "spirorbids" common, about 28.5%), bryozoans, microconchids, possi- (Watkins 1981) belonging to the microconchids (Taylor and ble cornulitids and organisms of uncertain affinities. All of Vinn 2006; Vinn and Taylor 2007; Zatoń and Taylor 2009), the sclerobionts likely utilised shells of dead cephalopods crinoids (Prokop and Turek 1983) and inarticulate brachio- as a hard substrate for their settlement. However, most of pods (Lockley and Antia 1980). Numerous authors have the infestation appears on internal moulds because the reported diverse assemblages of encrusters on cephalopod cephalopod aragonite was dissolved during diagenesis. It is shells (nautiloids, goniatitids and clymeniids) during the possible that some of the sclerobionts encrusted exhumed Devonian, and many examples from various stratigraphic lithified internal moulds. These isolated cephalopod shells levels (Emsian to Famennian) are known. These epibionts and internal moulds likely served as benthic islands for include corals, bryozoans, crinoids, cystoids, brachiopods various encrusters on a Devonian muddy sea-floor. and bivalves, as well as problematic worm tubes or foraminiferans (e.g. Baird et al. 1989;Chlupáč and Turek Keywords Sclerobionts . Hard substrate . Cephalopods . 1983; Davis et al. 1999; Grimm 1998; Klug and Korn 2001; Devonian . Holy Cross Mountains . Poland Nagel 2006;Thayer1974). Moreover, colonisation of early Carboniferous orthoconic nautiloids by problematic worm tubes has been documented by Klug and Korn (2001). Many Introduction encrusters on ammonoid or nautiloid shells and belemnite rostra are known from the Mesozoic. These include Many modern and ancient invertebrates encrusting cepha- foraminiferans, annelids, corals, bryozoans, bivalves, bra- lopod shells have been documented by numerous authors. chiopods, crinoids, barnacles, and oysters (e.g. Cope 1968; The earliest examples of cephalopod shell colonisation by Heptonstall 1970;Kauffman1978; Klug and Lehmkuhl hard substrate biota are noted in the Upper Ordovician. 2004; Lukeneder 2008; Macchioni 2000; Manni et al. 1991; Meischner 1968; Nicosia 1986;Pugaczewska1965; Rakús and Zítt 1993;Seilacher1960; Schmid-Röhl and Röhl 2003; M. Rakociński (*) Wilson et al. 1998). Landman et al. (1987)documented Faculty of Earth Sciences, University of Silesia, numerous examples of modern cephalopods infested by Będzińska Str. 60, 41-200 Sosnowiec, Poland various organisms utilising the shells of Recent Nautilus. e-mail: [email protected] These include foraminiferans, serpulids, bryozoans, corals, 64 Palaeobio Palaeoenv (2011) 91:63–73 barnacles, scyphozoans, bivalves, and sponges (see also 2007), and one black shale horizon denoted as the Kowala Donovan 1989; Maeda and Seilacher 1996;Reyment Black Shale by Marynowski and Filipiak (2007). This 2008; Taylor and Wilson 2003 and references therein). succession is located on the northeastern and northern walls Donovan (1989) illustrated an interesting discovery of a of the Kowala Quarry (Fig. 1). The stratigraphic setting of modern coleoid shell of Spirula spirula that remained this unit has been determined using ammonoids (Rakociński covered by soft tissue while the cephalopod was alive, 2007, 2009) and includes the zones ranging from Clymenia subsequently being encrusted by the barnacle Lepas laevigata to Wocklumeria sphaeroides, which correspond to anatifera after death when soft tissues of the cephalopod the following units of conodont zones: a considerable part of decayed. While numerous occurrences of sclerobionts on expansa and the lower and middle parts of praesulcata.This Devonian cephalopods have been described, very little setting corresponds to the P. jugosus and D. trigonica of research on this subject has been done in Poland. Dzik (2006). Berkowski (2002, Pl. 8, fig. 3) illustrated one specimen of the Famennian goniatite Sporadoceras infested by the coral Neaxon tenuiseptatus Różkowska from the Kowala Material and methods Quarry. Epibionts on cephalopod hosts from the upper- most Famennian of the Kowala Quarry and their palae- Material oecologic and taphonomic implications are described here. The material for this investigation was collected between 2003 and 2009 from the northern and northeastern walls Geological setting (mainly in rubble) of the active Kowala Quarry (Fig. 1). About 1200 specimens (represented by complete specimens The area investigated in this study is located in the southern and fragments) were examined. Most of the cephalopods limb of the Gałęzice–Kowala syncline, in the southern part are represented by internal moulds, occasionally with shell of the Kielce region of the Holy Cross Mountains, Poland, fragments. In addition, some specimens were partly approximately 10 km southwest of Kielce (Fig. 1). The surrounded by rock matrix. In 2008–2009, specimens of succession starts with carbonate deposits, which are partly Famennian cephalopods from the Kowala Quarry were reef limestones of Frasnian age. They are overlain by studied to find epibionts. Only 50 specimens of cephalo- Famennian rocks represented by thin-bedded rhythmic pods displayed encrustation by various sclerobionts. Before successions of dark-grey and black limestone and marly being photographed, the specimens were coated with shale in the lower and middle parts of the Famennian ammonium chloride (NH4Cl). Images of some sclerobionts succession (Berkowski 2002; Racki et al. 2002;Szulczewski were also prepared using an environmental scanning 1971). In the uppermost part of the Famennian section electron microscope (model XL30 ESEM/TMP; Philips, exposed at the Kowala Quarry, green and red nodular marly- Eindhoven, the Netherlands), which made it possible to limestone with abundant cephalopods intercalate with marly scan the specimens without coating. All specimens are shale and several black shale horizons and tuffites. An housed at the Faculty of Earth Sciences, University of uppermost black shale horizon corresponds with the Silesia, Sosnowiec, under the collection number GIUS-4. Hangenberg event (Marynowski and Filipiak 2007). The Famennian section in the Kowala Quarry is the most Terminology complete and thickest succession of the series in the Holy Cross Mountains (e.g. Berkowski 2002;Dzik2006). The Many terms have been used to categorise organisms total thickness of Famennian deposits here is about 200 m inhabiting marine hard substrates (for review see Davis et (e.g. Bond and Zatoń 2003). The Famennian part of the al. 1999; Taylor and Wilson 2002, 2003). In the field of Kowala section has been subdivided into informal lithological palaeontology, a commonly encountered problem is that of sets from H-3 to L (Berkowski 2002; Racki and Szulczewski establishing whether organisms attached to a shell while the 1996). The specimens described in this report were found in cephalopod was alive, while the shell was floating in the the uppermost part of the Famennian succession, known as sea after the death of the cephalopod (necroplanctonic)or unit L (sensu Berkowski 2002). The strata investigated while the shell was lying on the sea floor as a hard consist of green and red marly cephalopod-bearing lime- substrate. However, several criteria can be used as stone, intercalating with marly shale (“Wocklumeria Lime- indicators of whether an organism settled on a cephalopod stone”; for a more detailed description, see, for example,, syn-vivo or post-mortem (e.g. Davis et al. 1999; Klug and Berkowski 2002; Dzik 2006; Marynowski and Filipiak Korn 2001). Palaeobio Palaeoenv (2011) 91:63–73 65 Fig. 1 a Simplified geological map of the western and central part of (state on October 2005; photograph by M. Lewandowski), with the Holy Cross Mountains (after Marynowski and Filipiak 2007). b informal lithological set L of Berkowski (2002); KBS Kowala black Aerial view of the Kowala quarry (from Google Maps: http://maps. shale. d Rubble on northern wall of the Kowala Quarry (state on google.com). The walls shown in c and d are indicated by white October 2005; photograph by M. Lewandowski) arrows. c View of the northeastern wall of the active Kowala Quarry 66 Palaeobio Palaeoenv (2011) 91:63–73 Davis et al. (1999) used the term epizoa for