Rivista Italians di Palcontologia e Stratigrafia volume 110 no. 1 2 pis. pp. 3C3-309 April 2004

THE NUSPLINGEN LITHOGRAPHIC LIMESTONE - A "FOSSIL LAGERSTAETTE" OF LATE KIMMERIDGIAN AGE FROM THE SWABIAN ALB (GERMANY)

GERD DIETL 1 & GUNTER SCHWEIGERT :

Received October 10, 2002; accepted October I, 2003

Keywords: Fossil lagerstaette, protected excavation area, Up­ dle of the 19,h century (O. Fraas 1855; Quenstedt 1843, per Jurassic, Kimmeridgian, lithographic limestone, Swabian Alb, S\V 1855; Dietl et al. 2000). Several excavation campaigns took Germany. place, at first with commercial aspects, prospecting for

Abstract. A short overview is given on the actual excavations limestones to be used for lithography or for roofs, but within a fossiliferous lithographic limestone site near Nusphngen (west­ later restricted to scientific interests. A bibliography on ern Swabian Alb, SW Germany). In contrast to the better known Ear­ the Nusplingen Lithographic Limestone with all refer­ ly Tithonian Solnbofen Lithographic Limestone, it is much richer in ences until 1997 was published by Schweigert (1997). fossils and 0.5 my older (Late Kimmeridgian, Beckeri Zone, Ulmense Sub/one). Many of the fossils exhibit an excellent preservation, some­ In contrast to Solnhofen, the Nusplingen Litho­ times even of the organic matter. More than 7.000 specimens have been graphic Limestone is of Late Kimmeridgian age (Fig. 2). recorded belonging to almost 300 taxa (plants, microlossils, inverte­ It represents a stratigraphical equivalent to the "Liegcnde brates, iclmofossils, and vertebrates). Bankkalk" Formation which belongs to the Ulmense Sub- zone, Beckeri Zone (Fig. 3), thus about 0.5 my older than Riassunto. Vienc fornita una breve rassegna degli attuali scavi in tin sito di calcare litografico fossilifero vicino a Nusplingen (Alb Svevo occidental, Germania sudoccideniale). In contrasto con il piu nolo Calcare Litografico di Solnholen del Tithoniano inferiore, que- sto e molio piu ricco di fossili, ed e piu vecchio di 0.5 milioni di anni (Kimmeridgiano superiore, Zona a Beckeri, Sottozona ad Ulmense). Moln lossili mostrano una conservazione ecccllente, a volte perfino dell.i materia organica. Sono stati docunientati piu di 7000 esemplari appanenenti a quasi 300 taxa (piante, microfossili, invertebrati, icno- (ossili e vertebrati).

Introduction

The Nusplingen Lithographic Limestone is a fos­ sil lagerstaette similar to the famous Tithonian Solnhofen Lithographic Limestone of southern Franconia in Bavar­ ia. It is located in the western part of the Swabian Alb, few kilometers north of the Upper Danube Valley (SW Germany) (Fig. 1). The first fossils from the Nusplin­ Fig. I - Map showing the location of the fossil site Nusplingen, a gen Lithographic Limestone were reported since the mid­ few km N of the River Danube, SW Germany.

Staailiches Museum fur Naturkunde, Rosenstein 1, D-70I9I Stuttgart, Germany. 1 g.dietl.snins(("naturkundemuseum-bw.de, 2 schweigert smns(«lnaturkundemuseum-bw.de 304 G. Died & G. Schweigert

Fig. 3 - The general stratigraphical position of the Nusplingen Litho­ graphic Limestone (Nusp. PI. Pm.) within the Upper Jurassic of the southwestern Swabian Alb/SW Germany.

Environment

The laminated limestones were deposited in a more or less anoxic environment of a lagoon surrounded by sponge/microbial bioherms, some of them partly tecton- ically uplifted over the sea level as small islands (Fig. 4). The minimum depth of the lagoon was approximately 80-100 m. At the seafloor of the lagoon living conditions Pig. 2 - Detailed section of the Nusplingen Lithographic Lime­ were unfavourable for most organisms apart from very stone and its stratigraphical position within the Upper Kim­ meridgian. The thickness of the Nusplingen Lithographic few exclusions. Hence, an exceptional preservation was Limestone in the figured section is of about 10.5 m. possible which is normally prevented by bioturbation. Within the succession of laminated limestones, several turbidites with graded bedding ("allodapic limestones") and few thicker bioturbated beds indicating a better ox­ the Solnhofen Lithographic Limestone famous for the ygenation are intercalated. The turbidites comprise an­ oldest bird, Archaeopteryx. This is confirmed by the am­ gular lithoclasts and other components like calcareous monite fauna, which allows a high resolution biostratigra- ooids, sponge spicules, and even some fragments of her- phy (Schweigert 1998b; Schweigert & Zeiss 1999) .Today, matypic corals from shallow areas surrounding the la­ the outcrop of the Nusplingen Lithographic Limestone goon. It is possible to correlate the outcrops in the quar­ is not larger than 1.5 km2; its thickness reaches a maxi­ ries with the help of these thicker beds. Besides, some mum of 17 m. Several sections from drillings, from two silica layers also occur throughout the basin. The depo­ small quarries and from natural outcrops were reported sition of laminated limestones ended abruptly. From the by Dietl et al. (1998) and by Bantel et al. (1999). Since nearby sponge/microbial reefs, a lot of boulders had been 1983 the whole area has become a Protected Excavation broken off, gliding into the central parts of the lagoon, Area ("Grabungsschutzgebiet") by the Monument Pro­ where they were deposited directly above the laminated tection Law (see Bloos, this volume) of the "Land" Baden- limestones with a thickness of several meters. The allo- Wuerttemberg because of its extraordinary fossils. The chthonous nature of these boulders is indicated by ge- Nusplingen Lithographic Limestone is actually exposed opetal fillings in hollow fossils and several finds of am­ in two small quarries. For the public it is not allowed to monites significantly older in age than the underlying excavate or to take away fossils from the two quarries laminites. Below the boulder beds, the original stratifi­ or from everywhere in the protected area. To visitors lo­ cation of the laminites is strongly disturbed, with fold­ cal instructions about the importance of this fossil site ing, boudinage, lateral compression, and repetition by and about the above mentioned prohibitions are given stapling. We presume that a strong seaquake was the rea­ directly at the quarries. Visitors and excursions can visit son for this dramatic event. No data are available from at any time the quarries of the Nusplingen Lithographic the younger Upper Jurassic in the closer area because Limestone. of Cenozoic erosion. Nusplingen Lithographic Limestone 305

Fig. 4 - Reconstruction and palaeoge- ographic setting of the Nus­ plingen lagoon, without scale.

and Baiera), which are derived from near­ by uplifted islands, are still preserved with their organic tissue showing finest details like cell-structures and stomata. Recent­ ly, we were even able to identify amber (Schweigert & Dietl 2003). This amber occurs still in situ within the resin vessels of some araucariacean cone scales. Marine algae are restricted to bituminous layers. They closely resemble recent brown algae like Cystoseira. Calcareous algae have not been recorded at all.

Invertebrates Completely preserved specimens of non-rigid sponges like the Rhax-bezr- ing Codites (formerly often misidentified as brown algae and better known as "Pbyl- lothallus") are rather common (PI. 2, fig. The fossils from the Nusplingen Lithographic Limestone 1). Bivalves are mostly documented by different species of pectinids, limids, and oysters. Apart from a single layer with small nuculids, all bi­ New excavations by the Natural History Museum valves are allochthonous. They were brought into the laminated facies by predators or passively living on drifting ammonites before they final­ of Stuttgart (SMNS) in two small quarries accompanied by ly sank to the sea floor. The most frequent and typical macrofossils of several drillings proliferate new data on the sedimentology, the Nusplingen Lithographic Limestone are ammonites and aptychi. The palaeoecology, taphonomy, biostratinomy, and especially ammonite fauna comprises Lithacoceras ulmense (Oppel), Silicispbmctes the fossil content of the Nusplingen Lithographic Lime­ boelderi (Sapunov), Silicisphinctes russi (Schweigert), Hybonoticeras sp., stone (Dietl & Schweigert 1999b, 2001). Since 1993 more Physodoceras nattheimense Schweigert, Sutneria cf. rebholzi Berckhemer, Ochetoceras zio (Oppel), Glochiceras lens Berckhemer, Lingulaticeraspseu- than 7.000 fossils were recovered and are going to be re­ dopercevali Schweigert etc., characteristic of the boelderi faunal horizon gistered in an electronic database. They belong to almost of the Ulmense Subzone, Late Kimmeridgian (Schweigert 1998b; Sch­ 300 taxa, many of them new. Some of the fossils which are weigert & Zeiss 1999). Some ammonites are very spectacular because embedded in bituminous layers occurring in the upper part they are preserved with their complete jaw apparatus and stomach content of the succession exhibit an extraordinary preservation of (Schweigert & Dietl 1999a, 2001). The lower jaw is represented by the calcitic Aptychus, whereas the upper jaw which was originally chitineous organic matter. They tell us interesting stories about sea life is preserved in carbon. Also nautilids (Pseudaganides) have been found in or around the Nusplingen lagoon and in the surround­ with both jaws (rhyncolithes = upper jaw and conchorhynchus = lower ing shallow sea of the Upper Jurassic in SW Germany, far jaw) still in the body chamber, together with stomach-contents consisting away from larger continental islands like the Ardennian of decapod or ophiuran remains (Dietl & Schweigert 1999a) (PI. 2, fig. and Bohemian Massifs in the north and in the north-east. 3). Belemnites occur frequently in all ontogenetic stages of a single spe­ cies, Hibolithes semisulcatus (Miinster) (Schweigert 1999). The guards are very often bitten by predators. Teuthids (Trachyteuthis, Plesioteuthis, Lep- Microfossils totheuthis etc.) are also mostly bitten but often preserved with their inksacs. The Nusplingen Lithographic Limestone yields a significant amount of debris from the nearby reefs and from the water column. A Among the arthropods, marine polychaetes sometimes exhibit high percentage is made up of coccolithophorids and their debris char­ a soft-part preservation (Schweigert & Dietl 2000). A very well known acterizes the laminites as marine alginites (Bantel et al. 1999). Surface fossil of Upper Jurassic lithographic limestones is the limulid Mesolimu- structures of some bedding plains point to a temporary development of lus walchi (Desmarest), but in the Nusplingen Lithographic Limestone microbial mats. More than 60 species of radiolarians were identified in it is quite rare. The newly excavated fossils include the first record of a thin sections or by dissolving the limestone with acid (Ziigel et al. 1998; centipede, Eogeopbilus jurassicus Schweigert & Dietl, from the Jurassic Bantel et al. 1999). Together with the ammonites, the radiolarians allow (Schweigert & Dietl 1997). Other remarkable fossils are several insects an integrated biostratigraphical approach in the latest Kimmeridgian. such as beetles and dragonflies (Urogomphus, Stenopblebia,Aeschmdium, Other common microfossils are Saccocoma skeleton elements and vari­ Cymatophlebia) (PI. 1, fig. 2). They are also partly preserved in organic ous sponge spicules, namely of the Rhax type. matter (Schweigert et al. 1996; Bechly 1998). Many well-preserved de­ capods are typical of the Nusplingen Lithographic Limestone and more

Plants than 20 taxa have been recorded: Antrimpos, Cycleryon, Hefriga, Aeger, Bylgia, Dusa, Coleia, Eryma, Glyphea, Palaeopentacheles, Pustulina etc. In bituminous strata occurring in the higher part of the succes­ (Schweigert et al. 2000; Schweigert 2001b) (PI. 2, fig. 2). Coleia longipes sion land plants (PI. 1, fig. 1) (e.g. Cycadopteris, Brachyplyyllum, Zamites, 306 G. Died & G. Schweigert

(O. Fraas), a primitive eryonid, only known from Nusphngcn, still shows is much more diverse, although restricted to specific layers. Its compris­

its compound eyes (Schweigert & Dietl 1999b) (PI. 2, fig.4). es leeding burrows like Gbondrites, Tliieyif/.$cbelinuiy Pdrabaeiitzscbeltmti, Brachiopods are present with rare specimens of the genera Gbei- Rbr/.oamdlmm, and moving trails such as Serperiticbnordcs, Iclsowchiits, rvtbyris, Lacunosella, Ortiitbella, and Torqnirhyncbici (Dietl & Schweigert Undichiia (Schweigert 1998a; 2001a). Coprolites are generally frequent 2000). Other benrhic organisms such as echinoids (Stomucbimis, Rbab- in the laminites indicating that there was marine life in the water column docidaris, Polycittaris, Plegiocidaris, Paracidaris, Nenoticidaris, Diplopo- above the anoxic sea floor. Some phosphoritic ones are attributed to hshes diii, and Diplocidnris), ophiurans (Sinosura), and crinoids (Ptcrocmna, and reptiles, whereas the very common Limtbrtcana is interpreted as the Sohinocriniles, and Phcatoerinus) also occur only sporadically (Grawe- faeces of aspidoceratid ammonites (Dietl & Schweigert 2001). Baumeister ct al. 2000). In contrast, the planctonic crinoid Saccocoma tenelLi (Coldfuss) is rather frequent on some bedding planes, but much less conspicuous than in the Tithonian of Solnhofcn. Remarks

Vertebrates Both the diversity of the fossil fauna and flora and Among the vertebrate launa the fishes are predominant. They are represented by elasmobraiichii (Sr/uatiria, Eonotidanus, 1 leterodon- its exceptional preservation make the Upper Jurassic Nus­ [Ks, Spberiodus etc.) (Schwcizcr 1964; Bottcher & Duffin 2000), holo- plingen Lithographic Limestone an outstanding fossil la- cephali (licbyodus, Elasmodectes) (Heimberg 1949; Schwcizer 1964), gerstaette which opens a window into the past ol the crossopterygn (Coccoderma, Undina), and numerous actinopterygii earth. Some interesting parts of the section, however, have like Allotbrissops, Amictbtdioji, Aspidorbyncbtts, Gaturtis, turyconntts, been exploited only for few m:, which provides hardly any Euro, Gyrodus, Tepto/epides, Ophiopsis, Siemensichtbys, Sotnhofenamia, Thirstf etc (Hcincke 1906). The most typical fish of the Nusplingen statistical database. Thus, we hope the excavations can be fossil site, however, is the angel shark Squatirnt ucantboderma (Fig. 5), continued successfully in the following years. typically preserved with its complete skeleton and skin outline. Dur­ ing the new excavations, 10 well preserved specimens, juveniles, males, and lemales, were discovered. Acknowledgements. The enormously enlarged knowledge on this Reptiles rarely occur, but this may be due to the small size o! the fossil lagerstaette would have been impossible without the engaged excavations. They are represented bv marine crocodiles (Geosaurus, Daco- work of our technicians, namely M. Kapitzke, M. Rioter, O. Maass (.ill uturus) (F. Fraas 1902) and 3 genera of pterosaurs, Pterodactylus, Gallodac- SMNS), and of our enthusiastic volunteers R. Hugger (Albstadt-On- tylus, and Rhwipborbyncbtn (Qucnstedt 1855; Plieninger 1907). During the stmettingen), B. Russ (Nusplingen) and O. Dietl (Stuttgart). The ex­ new excavations, onlv several isolated bones and teeth were lound. cavations and the contemporaneous scientific work were generouslv funded by the Deutsche Forschungsgemeinschalt (Dl 680/1). Mam- Ichnofossils thanks should also go to all others who engaged themselves m our In contrast to the typical Lithographic Limestones of the Sol- project. We arc grateful to K. Page and U. Nicosia for their comments nholen area, the ichnolauna of the Nusphngcn Lithographic Limestone and suggestions. Nusplingen Lithographic Limestone 307

PI. AT I' I lij;. 1 - I'lendospcrm - Cycddopicvn jurcnsis (Kurr) Ilirmer, Nusplingen Lithographic Limestone (Irom bituminous bed). Nusplingen quarry, length 40 cm, SMNS PI875. I'll;. 2 - Dragonllv - Urognmpbiis iiitsplnigaisrs Bcchlv, Nusplingen Lithographic Limestone (Irom bittiminous bed). Nusplingen quarrv, span ol wings 15.5 cm, SMNS 62602. 308 G. Dietl & G. Schurcigcri

PI.ATI-, 2

Pig. 1 - Non-rigid sponge - Coililes serpemiiium (Sternberg), Nusplingen Lithographic Limestone, Nusplingen quarrv, height 1 I cm, SMNS 64402. Pig. 2 - Eryonid decapod - Cyvleryon prophie/tms (Schlotheim) O (= spinimumis Germar), Nusplingen Lithographic Limestone, Nusplingen quarry, length 3272. Pig. 4 - Eryonid decapod - Coleia longipes (O. Fraas), Nusplingen Lithographic Limestone. Nusplingen quarrv, length 9.3 cm, SMNS 63744. Nusplingen Lithographic Limestone 309

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