Cent. Eur. J. Geosci. • 4(1) • 2012 • 9-46 DOI: 10.2478/s13533-011-0060-0

Central European Journal of Geosciences

The Middle Triassic marine reptile biodiversity in the Germanic Basin,in the centre of the Pangaean world

Research article

Cajus G. Diedrich1∗,

1 PaleoLogic, Private Research Institute Halle/Westph., GERMANY

Received 12 September 2011; accepted 16 December 2011

Abstract: The Middle Triassic fossil reptile localities near Bayreuth (Bavaria, southern Germany) consist of shallow marine autochthonous glauconitic marls and terebratulid-rich tempestite carbonates of the newly defined Bindlach and Hegnabrunn formations. Single bones and incomplete skeletons of marine reptiles have been recorded in bone beds within in the Illyrian and Fassanian stages. These include the remains of the sauropterygians Neusticosaurus sp., Lariosaurus cf. buzzii [1], Nothosaurus mirabilis [2], Paranothosaurus giganteus [2], Placodus gigas [3], Cyamodus rostratus [4], Cyamodus münsteri [5], Pistosaurus longaevus [6], and ichthyosaurs Omphalosaurus sp., and Shastasaurus sp. or proterosaur Tanystrophaeus conspicuus [7]. New skeletal reconstructions are based on the osteological analysis of three dimensionally preserved bones and skeletal remains. The large number of marine endemic placodont macroalgae feeders (P. gigas) in the Bayreuth sites coincides with the presence of invertebrate palaeocommunities that are characteristic of macroalgae meadow paleoenvironments. Most of the reptile species and genera from the Bayreuth localities also occur in beds of similar ages from the Monte San Giorgio (Switzerland/Italy) or Perledo (Italy) lagoonal areas. Ichthyosaurs and pistosaurs were adapted for open marine conditions, and may have migrated from the Panthalassa Oceans into the shallow marine Germanic Basin to reproduce, whereas placodonts and many other sauropterygians seem to have lived permanently in those shallow marine habitats, with large squamates and thecodont or smaller archosaurs in coastal areas. Keywords: Reptile biodiversity, skeletal reconstructions, Illyrian/Fassanian (Middle Triassic), palaeobiology, palaeobiogeog- raphy, reproduction zones, Germanic Basin, global faunal interchange © Versita Sp. z o.o.

1. Introduction Muschelkalk” limestone, which outcrops in the vicinity of Bayreuth, in Bavaria (southern Germany) - see Figure 1B-C; [5,7, 12, 13]. These localities occur near Heg- nabrunn, Bindlach (Bindlacher Berg), and Laineck (Os- chersberg). Four of these exposures remain accessible Several important Middle Triassic (Illyrian to Fassanian) Fig. 1C and are valuable for the recovery of missing or marine vertebrate fossil localities are known from Pan- lost information relating to historically collected material. gaea Fig.1A; [7–11]. One such area with several bone- These small quarries were already famous at the begin- rich sites is situated in the intracratonic Germanic Basin, ning of the 19th century, with the first Placodus tooth in the world famous and historically important “Upper finds reported in 1800 by Graf zu Münster [5, 14]. Inten- sive collecting during the 19th century, mainly by manual ∗E-mail: [email protected] workers in quarries on six outcrops, resulted in the col-

9 The Middle Triassic marine reptile biodiversity in the Germanic Basin,in the centre of the Pangaean world

Figure 1. The investigated historical and recent Upper Muschelkalk (Middle Triassic) outcrops along the Lainecker Höhenzug, near Bayreuth (Bavaria, southern Germany), and vertebrate bone-rich or skeleton localities discussed in the text (Middle Triassic Globe after [26]; intertidals and Germanic Basin palaeogeography from [28]; Middle Triassic localities after [1, 7, 8, 22, 23, 30, 31, 42, 101, 113, 122, 158, 159, 161]).

10 Cajus G. Diedrich,

Figure 2. Stratigraphy of the Lainecker Höhenzug Upper Muschelkalk (Middle Triassic) carbonates (about 70 metres thick), with the new formation subdivision, compiled from the three main localities at Laineck, Hegnabrunn and Bindlach (Bavaria). A-D. Main facies and sediment types of the Lainecker Höhenzug Upper Muschelkalk carbonates: A. Glauconitic nodular marls with infauna (here two Pleuromya) and terebratulid-dominated tempestite shell beds of the Bindlach Formation at Bindlach 1. B. Boundary between the platy limestone (Tonplatten) facies and the glauconite marl/tempestite beds at Bindlach 1. C. Variable glauconitic nodular marls and terebratulid-rich tempestites of the Hegnabrunn Formation, at Hegnabrunn. D. Variable dolomites and dark bone beds at top of the Meißner Formation / base of the Warburg Formation, in the Laineck road cutting.

11 The Middle Triassic marine reptile biodiversity in the Germanic Basin,in the centre of the Pangaean world

lection of thousands of, mainly marine, reptilian teeth and bioherm colonies [31]. The Muschelkalkmuseum at In- bones, as well as the remains of two skeletons (the first gelfingen houses an overview selection of these world- Nothosaurus:[7]; and the first Pistosaurus:[14]). The famous Muschelkalk fossils, and is also famous for its late most famous “Triassic bone monograph”, which compiled Upper Muschelkalk reptile skeletons of the thecodont ar- material mainly from the Bayreuth sites, was published chosaur Batrachotomus kupferzellensis, as well as many by Meyer [7] including illustrated skeletons, skulls, and skeletal remains of amphibians from southern Germany individual postcranial bones. Most of these specimens [31]. were forgotten for a long time, but fortunately only a few Of worldwide importance are the holotypes and originals appear to have been lost during the two world wars. of fish from Bayreuth [32, 33] and, in particular, the rep- The material illustrated by Meyer [7], as well as other tile remains such as the placodont holotype and origi- previously undescribed material, is revised herein in a nal skulls of Placodus and Cyamodus [3, 4, 6, 7, 10, 12, new overview monograph, using multidisciplinary methods 13, 20, 34–43]. Cyamodus remains from Germany have including stratigraphic studies, facies analyses, osteol- recently been revised, including a full osteological and ogy, actuopalaeontology, and palaeoecology. This review palaeoecological review [44], and the main Placodus gi- has included all of the “forgotten” collections, which are gas material from Germany has also now been revised spread across several German museums. These collections in a separate monograph [17]. Not only have individual even include skeletons and mounted skeletal casts from a skulls, teeth and bones been collected from the Bayreuth number of Middle Triassic marine reptiles of the Germanic area, but also articulated skeletons of the first described Basin. Amongst these skeletons is the holotype skeleton nothosaur Nothosaurus mirabilis in the world, and the of the world’s only known placodont Placodus gigas [15], only known skeleton of the pistosaur Pistosaurus longae- the original being in the Senckenbergmuseum in Frank- vus [6, 7, 21, 45, 46]. Several bones from the famous “long- furt [15–17], with casts exhibited in the Urwelt-Museum necked lizard“ squamate Tanystrophaeus conspicuus [7] of Oberfranken at Bayreuth and at the University of Tuebin- the Germanic Basin (mainly from Bayreuth sites), have gen [18]. Also included in these collections is the holo- also been re-examined [47]. type skeleton of the placodont Henodus chelydrops [19], the paratype skeleton of Pistosaurus longaevus [20, 21], The German Middle Triassic fossil collections cover a skeletal casts of Simosaurus gaillardoti (cf. [22]), several longer history and stratigraphic range than the Monte original skeletons of the pachypleurosaur Neusticosaurus San Giorgio Swiss/Italian collections and have at least pusillus [23], as well as other skeletons such as holo- as much international value. Compared to the tectoni- types and originals from the amphibians Mastodonsaurus cally deformed and compressed bones of the Monte San giganteus and Gerrothorax pustuloglomeratus [24, 25]. Giorgio skeletons (for example [10, 48]), the Germanic Muschelkalk bones are mostly preserved in three dimen- Several German collections also contain many Middle Tri- sions (see [7], and the new ”bone atlas” presented herein assic reptile skeletons from the Germanic Basin that have ) and provide further information and interpretation pos- never been recorded from Monte San Giorgio (for example sibilities with regard to the precise bone osteology and Placodus gigas). However, this research focuses on the histology, skeletal reconstructions, and the locomotion of Middle Triassic marine and coastal species from the Up- these reptiles. The reconstructions of reptile skeletons per Muschelkalk of the Bayreuth area only and provides from the Germanic Basin presented in Diedrich [49] are an overview of the palaeobiodoversity of this rich reptile updated with few modifications based on combined inter- faunal assemblage. This study does not attempt a global pretations from skeletal material and individual bones. revision of all Middle Triassic reptiles, despite the need for such a revision, which was started [10]. In this con- The Bayreuth region of Germany represents an important tribution some species which were belived to be endemic part of the geological and palaeontological history of the are revised to occur with similar species in the Monte San Upper Muschelkalk in the Germanic Basin [27, 31, 50–55]. Giorgio and the Germanic Basin. It has provided information that allows palaeogeograph- ical, biogeographical, and palaeoenvironmental maps to The historical Bayreuth reptile discoveries are from late be compiled showing the development of the Germanic Illyrian to Fassanian and new material was collected from Basin during the Middle Triassic [28], as well as the es- a road cut from the early Longobardian layers (the mid- tablishment of basin-wide facies correlations using Ger- dle of the Middle Triassic), a period in which major ex- manic ceratite biozones. pansion of the marine Germanic Basin took place [26– 29]. The famous Muschelkalk invertebrate and vertebrate The similarities between the faunas of Monte San Giorgio “Muschelkalk-Fossillagerstätten” fossil deposits [30] in- and the Germanic Basin, together with an improved un- clude abundant cephalopod ceratites and extensive crinoid derstanding of the global palaeobiology of Middle Trias-

12 Cajus G. Diedrich,

sic reptiles, mean that earlier models proposed by Rieppel the Paläontologische Museum of the University of Zürich and Hagdorn [42] now need to be considerably modified (PIUMZ). and updated with respect to the habitats, palaeoecology, The stratigraphy, sedimentology and facies of all the ac- and biogeography of the placodonts. In particular, the cessible Bayreuth exposures have now been studied in theory that “Triassic Sea Cows” (not only Placodus gi- comprehensive investigations based on recent fieldwork. gas) were macroalgae feeders and exhibited convergent This was considered necessary because neither the strati- development with Sirenian seagrass feeders [43, 56] will graphic ages nor the facies relationships of the fossil dis- be supported herein through the review of the Bayreuth coveries from the various quarries were previously well localities, their facies, and analysis of their qualitative understood. The Lainecker Höhenzug ridge was explored and quantitative reptile faunal assamblages. The analy- to the north of Bayreuth, between Hegnabrunn and Lai- sis of facies-relationships, which have not been discussed neck, in order to relocate those historical outcrops that in any detail by previous authors such as Rieppel and remained accessible (Fig. 1). The Hegnabrunn quarry, Hagdorn [42] or Schoch and Wild [22], is the key for un- with its middle Upper Muschelkalk section, forms one of derstanding the global distribution and palaeobiology of the largest and best exposed sites and has not been back- marine reptiles in Pangea, its intracratonic shallow marine filled. Some of the other sites have, however, been at least basins and the coasts of the Tethys. Cladistic analyses partially backfilled while those at Laineck-Oschenberg of the marine reptiles, such as have now been performed and Laineck-Rodersberg have been completely backfilled. many times on the Bayreuth material [57–62], only con- Only the upper parts of the Bindlach 1 and 2 localities are tribute to a partial understanding of their phylogenetic now accessible, the lower parts being covered by gravel relationships, but do not offer any explanations for their dumps. The stratigraphy and sedimentology of three of palaeobiology or their distribution, and have even led to these historical quarries was studied. In 2010 a new road misidentifications, discussed in this paper for Lariosaurus cut between Bindlacher Berg and Oschersberg, northeast or Ceresiosaurus. of Laineck, exposed the uppermost Upper Muschelkalk for a short time, thus completing the section part of the late Upper Muschelkalk to early Keuper. The ceratite occur- 2. Materials and Methods rences in particular were compared in order to understand the biostratigraphic range of each section and bone bed The four main historical vertebrate and invertebrate ages. New ceratite material was collected from the gravel collections (Bindlach-Bindlacher Berg I-IV, Laineck- dumps or taken directly from the sections, allowing a first Oschenberg, Laineck-Rodersberg, and Hegnabrunn) from biostratigraphic subdivision of the sections (Fig. 2). Sed- six quarry localities along the Lainecker Höhenzug re- iments attached to bones in the collections often provided gion near Bayreuth have been studied using an interdis- supporting evidence for stratigraphic reconstructions of ciplinary approach (for localities see Figs. 1C and 2). the source horizon. Terebratulid shell-bed material is very often the main carbonate rock-type attached to bones, or Included in the study were all the holotypes, original glauconitic limestones, both of which are typical lower to specimens, and other unpublished bone material from the middle Upper Muschelkalk sediments of Bayreuth. Bayreuth quarries (Figs. 2B-C) in an attempt to un- derstand their context in terms of facies and palaeoen- Using the two skeletons of Nothosaurus mirabilis (coll. vironment. These specimens are housed in the Sencken- UM-O) and Pistosaurus longaevus (coll. SMF), it was bergmuseum, Frankfurt (SMF: mainly coll. Strunz), the possible to identify many single bones for the first Urwelt-Museum Oberfranken, Bayreuth (UM-O: mainly time, which resulted in the separate publication of about coll. Münster, Meyer and Strunz), the State Museum 200 individual Pistosaurus bones, recovered mainly from of Natural History (Museum Staatliche Naturhistorische the Bayreuth outcrops [21]. The Nothosaurus mirabilis Sammlungen) in Stuttgart (SNSD: mainly coll. Wild), postcranial skeleton is currently under preparation; only the Bayrische Staatssammlung at the University of Mu- certain important bones prepared from the skeleton are nich (BSP: parts of the coll. Meyer and Münster), the figured herein which were important for distinguishing sin- University of Tübingen (UT), and the Natural History Mu- gle bones from other reptiles (as well as for the new recon- seum of the Humboldt-University, Berlin (MB: mainly coll struction of the skeleton). This skeleton has been impor- Strunz, and Meyer). A small quantity of material is also tant for the successful identification of several hundreds stored in various other museums such as the Natural His- of individual bones from this species. The identification of tory Museum, Erfurt (ME), the Naturkundemuseum, Bam- this material also assisted in the identification of bone ma- berg (NMB), and the Natural History Museum in London terial from Paranothosaurus giganteus, the largest of all (NHMM). Skeletons used for comparisons are housed in the Bayreuth reptiles, which was then compared with the

13 The Middle Triassic marine reptile biodiversity in the Germanic Basin,in the centre of the Pangaean world

skeleton from Monte San Giorgio (P. amsleri, coll PIUMZ). within the marl beds (coll. UM-O). All three species were These single bones are also presented herein. Finally, the collected from the gravel dump, but a few specimens also Cyamodus and Placodus remains were identified, making came from within the section itself (Fig. 2), allowing the special use of the Placodus gigas skeleton from Steins- first biostratigraphic zonation and dating into the Illyr- furt (coll. SMF R1035) and the Cyamodus kuhnschnyderi ian of this quarry section and the neighbouring Bindlach skeleton from Monte San Giorgio (coll. PIUMZ). Follow- quarries. These quarries occupy a similar elevation and ing new determinations it was possible to identify several cover a similar stratigraphic range due to horizontal strat- hundred single bones from the Bayreuth material, includ- ification along the Bayreuther outcrops. ing some from the previously overlooked or mis-identified Lithology: At the bottom of the Bindlach quarry are repet- sauropterygian Lariosaurus. A cast of a large individual itive yellowish, dolomitic, 2 cm to 5 cm thin to 20 cm skeleton of Lariosaurus balsami from Comer Lake (Italy) thick-bedded platy limestones. Above the Middle/Upper and an original young individual specimen from Perledo Muschelkalk boundary first glauconitic and terebratulid (Italy) were used for the comparisons (cast of an older in- dominated massive limestone beds start the lithiologi- dividual skeleton in coll. BSP with no I.D. number, and cal change. About 6-8 metres of the following low- a younger individual original skeleton in coll. SMF, no. ermost Upper Muschelkalk are covered by the gravel R.13B). dump. The upper parts of the Bindlach Formation are exposed and consist of several parasequence series to- talling about 22 metres in thickness. The lithology 3. Geology comprises cyclic repetitions of similar bioturbated grey- greenish glauconitic marlstones (5-50 cm thick), green- 3.1. Stratigraphy and sedimentology ish glauconitic marlstones with carbonate nodules, grey- green marlstones, grey wavy marlstones (= “Wellenkalke” in Lower Muschelkalk facies type), and 5-30 cm thick Three main sections have provided a preliminary overview terebratulid dominated and fossil-rich bioclasts and oo- of the Illyrian to Longobardian (Upper Muschelkalk) lime- lites containing limestone beds, which later also include stones (Fig. 2), which have a total thickness of about bone beds, and finally crystalline grey-white thin lime- 70 m along the western margin of the Bohemian Mas- stone beds. The last thick glauconite marl bed in the sif. The Hegnabrunn section, which is the most northerly, compressus biozone below the Spiriferina Bed is defined the Bindlach 1 section, which is similar in age but has as the top of the Bindlach Formation. much less accessible outcrops, and the Laineck road cut- ting, which is the most southerly of the three, combine Bone bed layers: Three main bone beds can be identified to provide an almost complete Upper Muschelkalk marine in the Bindlach Formation from the abundance of fish- sequence. Marked lithological and faunal differences can scale remains (larger bones are rarer and not possible be seen between the sections, as well as from other basin to excavate in the sections): the pulcher, robustus and or marginal facies sections in Germany (Fig. 3). These compressus bone beds. The presence of several bone bed lithological and facies differences have allowed the iden- layers within this section explains the large quantity of tification of two new formations: the Bindlach Formation vertebrate remains recovered in historical times. These and the Hegnabrunn Formation. can be re-dated into the atavus to compressus (or possi- bly to the evolutus) ceratite biozones of the basal Upper Bindlach 1 (stratotype Bindlach Formation): The section Muschelkalk (late Illyrian to late Anisian). The vertebrate (GPS coordinates: 49◦59’41.03”N; 11◦36’58.77”E; Fig.2 reptile fauna in this facies is dominated by placodonts – part Bindlach 1) is at the Bindlach quarry 1 on the (mainly Placodus, with fewer Cyamodus and only rare plateau margin. Paraplacodus), pistosaurs (Pistosaurus), and nothosaurs Litho- and chronostratigraphy: The 22 metres exposed (rare Simosaurus, abundant Nothosaurus, and a small Bindlach section starts at the mm/mo boundary (Mid- proportion of Lariosaurus), but also includes pachypleu- dle/Upper Muschelkalk): only 50 cm thickness is exposed rosaurs (Neusticosaurus) and extremely rare ichthyosaurs of the top of the uppermost Middle Muschelkalk Diemel (Shastasaurus, Omphalmosaurus) or terrestrial reptiles Formation up to the basal Hegnabrunn formation (Illyr- (Tanystrophaeus; cf. Fig. 3 - see faunal analyses). ian). Hegnabrunn (stratotype Hegnabrunn Formation): This Biostratigraphy: Ceratites including Ceratites (Dolocer- section (GPS coordinates: 50◦ 5’9.40”N; 11◦33’11.56”E; atites) pulcher, Ceratites (Doloceratites) robustus, and Fig.2 – part Hegnabrunn, and GPS coordinates: Ceratites (Opheoceratites) compressus were found in the 49◦59’41.03”N; 11◦36’58.77”E; upper part Bindlach) is accessible part of the Bindlach 1 and 2 quarries, mainly an old quarry.

14 Cajus G. Diedrich,

Figure 3. Main Upper Muschelkalk marine benthic palaeocommunities and facies types in the Germanic Basin of central Europe. The formations are diachronous, and the ceratites disappear at various levels in the Warburg Formation (ceratites not proven in brackets). Bissendorf and Lamerden sections after [49, 62]; palaeocommunities compiled and redrawn from Aigner [31, 52], Diedrich, [130].

Litho- and chronostratigraphy: It comprises the middle postspinosus or ?C.(C) penndorfi/sublaevigatus were col- part of the Upper Muschelkalk it is about 19 metres thick, lected from the gravel stockpiles and from the upper part Fassanian in age. of the section. Together, all the ceratites found in Heg- nabrunn have a maximum range from the compressus to Biostratigraphy: The ceratites C. (A.) spinosus and C. (A.)

15 The Middle Triassic marine reptile biodiversity in the Germanic Basin,in the centre of the Pangaean world

the sublaevigatus ceratite biozones and are thus younger the base of this second part of the section. The section than in the section at Bindlach 1. reaches even higher into the early Keuper of the early La- dinian (semipartitus biozone) of the Warburg Formation, Lithology: The Formation starts with the top of the Bind- but no large ceratites were found above the nodosus zone lach section and consists of two thick, terebratulid-rich during this field work, and might be absent in this already shell layers, in which the presence of small, 1-2 millime- strong terrestrial influenced facies. tre glauconite sand particles is unique within the tem- pestite shell beds. These beds are similar in glauconite Lithology: After several metres of interbedded dark-grey content and tempestite origin to the south German well marl layers and thin micritic limestone layers (= Tonplat- distributed Spiriferina Bed. At Hegnabrunn the section ten) the “Alberti Bed” is possibly represented as a thick continues the Hegnabrunn Formation with two relatively bioclastic bed. This bivalve and terebratulid shell-rich bed thin sequences comprising terebratulid limestone bioclas- contains few vertebrate bones or teeth and includes some tic layers and marlstones, including limestone beds, which hardground layers. Following a brief hiatus the section appear to fall within the praecursor zone (Fig. 2). The se- continues with a marked lithlogical change to bioclastic quences of the following praespinosus zone are rhythmic limestone beds dominated by smaller terebratulid shells and smaller scaled (10-40 cm thick), consisting predom- and one of those beds seems to be the “Beds of small Ter- inantly of terebratulid limestone bioclastic beds that de- ebratulids”. Finally, following these sedimentary cycles veloped into hardgrounds with Balanoglossites/Trypanites and their marine deposits, grey dolomitic marls from the burrows. Enantiostreon oysters are commonly attached terrestrial influenced Warburg Formation occur at the top to the hardgrounds. These tempestites are overlain by of the section (Figs. 2D, 3), which explains the absence glauconitic, nodular, marly limestones. The spinosus zone of ceratites in this region. has two sequences, starting with several metres of mas- Bone bed layers: The first one is the nodosus bone bed. sive terebratulid bioclastic beds and followed by thick au- Placodonts appear to be absent above the nodosus zone, tochthonous, glauconitic, nodular, marly limestones (Fig. where limnic amphibians and lagoon-adapted reptiles 2). The postspinosus zone exhibits a change to yellowish such as askeptosaurs appear. Overlying the last tem- platy limestones and variable massive terebratulid bio- pestite layers several thin bone beds are developed, but clastic limestone beds (Fig. 2). In the penndorfi zone also the three terebratulid beds that are well distributed two 40 cm thick massive terebratulid bioclastic limestone in southern Germany. These are in turn overlain by the beds are developed, which are again overlain by green- boundary bone beds, which contain richer vertebrate re- ish, glauconitic, nodular, marly limestones that form the mains including bone elements from Blezingeria sp. (ver- top of the quarry section, but probably not the top of the tebra centrum), Mastodonsaurus sp. (vertebra centrum), Formation, which remains unexposed. and Nothosaurus giganteus (vertebra centrum), as well Bone bed layers: One of the tempestite shell layers is as coprolites. Actinopterygian scales or teeth and shark a bone bed - the evolutus bone bed. Another bone bed teeth (mainly Acrodus) are abundant. appears at the boundary between the spinosus and post- spinosus zones; the last bone bed is the postspinosus bone bed. 3.2. Facies and palaeocommunities Laineck-road cut: The section (GPS coordinates: There are several palaeocommunities in the Upper 49◦59’5.54”N; 11◦38’57.67”E; Fig.2 – part Laineck) was Muschelkalk facies types of the Germanic Basin (Fig. 2), exposed on the Oschersberg. four of which have been found within the Bayreuth sec- Litho- and chronostratigraphy: It comprises the middle tions. Some apparent variations in the palaeocommunities part of the Upper Muschelkalk of the Meißner and War- within particular Muschelkalk facies types (cf. Figs. 2-3) burg Formations (Fassanian to early Longobardian) over is simply due to variations in the invertebrate taphon- a total height of about 21 metres, but a section between omy. Eigth main facies dependant palaeocommunities are the lower and upper parts was not accessible. A 10 cm known in the Upper Muschelkalk of Germany: thick dark claystone bed within the sublaevigatus zone is The intertidal biolaminate community (Fig. 2A) of the possibly useful as a marker bed. Diemel Formation (top of the Middle Muschelkalk) is Biostratigraphy: Several ceratites were found mainly well defined stratigraphically in Bachmann and Kozur within the Tonplatten facies, including two well preserved [54] and from the facies and paleoenvironment of the bi- C. (C.) sublaevigatus steinkerns. Within the beds of small olaminates of the basal Diemel Formation (= mm1 sub- terebratulids (which often exhibit colour preservation) a stage) in Diedrich [62] it is not yet or anymore exposed steinkern of C. (C.) nodosus (in the (“Albertii Bed”) dates in the Bayreuth localities, but must be present below

16 Cajus G. Diedrich,

the dolomites of the basal exposed section in, for ex- abundant [70]. ample, the Bindlach 1 quarry. This is indicated by a The terebratulid hardground community (Fig. 3E): In the small horseshoe crab find [63], which is recently identi- Upper Muschelkalk, the terebratulid Coenothyris vulgaris fied to have come from those intertidal deposits of the can form massive limestone beds (10 metres in northwest- Bayreuther region [62]. Autochthonous macrofauna is ab- ern Germany along the Teutoburger Wald mountain chain, sent from such intertidal biolaminates, except for limulid [71], for example at Bissendorf there are more then 10 me- trackways ([28, 62]) and rare limulid body fossils [62], trees of terebratulid floatstones/wackestones in the middle which have been reported from the Bayreuth area (Limu- section of the Upper Muschelkalk (“Hauptterebratelbank”, lus agnotus [63]) and other German sites, together with in the evolutus biozone of the Osnabrück Formation, [49]), juvenile horseshoe crabs which are indicative of repro- which is diachronous and similar to the thinner and strati- duction carbonate mud flats [62]. Most common are verte- graphically younger “Hauptterebratelbank” (in the no- brate tracks Procolophonichnium, Rhynchosauroides, Chi- dosus biozone of the Meißner Formation, Fig. 2) in south- rotherium, Isochirotherium and Synaptichnium [28, 64–66] ern Germany [31]. Within some layers of the terebratulid from five terrestrial coast-dwelling reptiles. Tracks would beds these brachiopods are still articulated and have un- be expected in the Lainecker Höhenzug intertidal deposits dergone little transport. In most layers they occur as sin- because of their marginal position, but there is a lack of gle valves, as is commonly the case in the Bindlach, Lai- outcrops. neck and Hegnabrunn tempestites, indicating reworking The lagoonal hypersaline community (Fig. 2B) defined and/or transport and suggesting that this is not the origi- for the mm1 Substage of the Diemal Formation in Hagdorn nal habitat for these terebratulids. A few hardground beds and Simon [67] appears at the base and also at the top of occur within thicker series of shell-beds. These are pene- the Middle Muschelkalk, as well as in part of the Diemel trated by Trypanites and Balanoglossites drilling/boring Formation, but is poorly exposed at Bindlach 1. The fauna types of ichnites. The oyster Enantiostreon spondyloides in the platy dolomitic limestones of the Germanic Basin is is found attached to these hardground surfaces [31], in the typically strongly reduced and dominated by the infaunal positions in which they lived, as seen abundantly in the Neoschizodus orbicularis bivalve, which is mostly found Bindlach and Hegnabrunn sections. These hardgrounds concentrated in tempestite deposits together with bones were also most probably the original habitat and attach- and rare skeletons of the pachypleurosaur Anarosaurus ment substrate for the terebratulids, which were washed pumilio or Phygosaurus sp., and Nothosaurus marchicus by coastal storm events into the deeper water palaeocom- remains (cf.[64, 67–69]), in shallow subtidal channels. munity habitats described below. An oolitic sand bar community (Fig. 2C) of the basal Up- Glauconitic marl softground community (Fig. 3D): The per Muschelkalk has been recognised and described in greenish glauconitic nodular marl layers are 5-30 cm thick the Germanic Basin (cf. [31, 52]) and at Bad Sulza is and contain many faunal remains. These sediments have associated with a bone bed [44], but has not yet been infilled burrows of Thalassinoides, Planolites and Rhi- documented in the Bayreuth quarries. It can, however, zocorallium, which are the most abundant in the Upper be expected in the unexposed base of the Bindlach quar- Muschelkalk shallow marine sediments of the Germanic ries. At Bad Sulza (in central Germany) these carbonate Basin [70]. These marls have a mixed benthic mollusc and sands contain a very important marine vertebrate fauna pelagic ceratite fauna, which is similar to that which has that includes fish and shark remains but is mainly made been illustrated for the lower Upper Muschelkalk Has- up of sauropterygian reptiles such as Nothosaurus juve- smersheim Marls (atavus Biozone) of southern Germany nilis, Simosaurus sp., Neusticosaurus sp., the lariosaur [31], except that these do not include the Pleuronectites Lariosaurus cf. buzzii, placodonts Placodus gigas, Para- laevigatus with radial colour stripes that can be found at placodus sp., Cyamodus sp., and pistosaurs Pistosaurus Bindlach. The bivalve mud-stickers, such as the abundant longaevus [44]. The benthic invertebrates in the Germanic Pleuromya muscoloides and the less abundant Myophoria Basin are the bivalves Astartellopsis nuda, Neoschizodus vulgaris, are often found double-valved and buried in life ovatus, Neoschizodus laevigatus, Neoschizodus german- position (Fig. 2A). This is also the case for for double- icus, Elegantinia elegans, Parallelodon beyrichi, Pseu- valved epifauna such as Hoernesia socialis and Bakevel- domyoconcha mülleri, and Entolium discites, while the lia costata, which are found all over southern Germany scaphopods are represented by Plagyoglyptaf sp. [31]. [31]. This palaeocommunity, which is mainly present in Gastropods tend to be large forms including Undularia the Bindlach Formation, is important for providing indi- scalata, nautilids are rare (Germanonautilus sp.), and cer- rect proof of macroalgae especially by the presence of atites are absent. In some intercalating hardgrounds, the the gastropod Neritaria [31], whereas a detailed inverte- ichnites Balanoglossites and Trypanites are common or brate palaeocommunity assemblage study is lacking. Tem-

17 The Middle Triassic marine reptile biodiversity in the Germanic Basin,in the centre of the Pangaean world

Figure 4. Middle Triassic fish remains from the studied sections at the Lainecker Höhenzug, near Bayreuth (Bavaria, southern Germany). 1. Hybodus longiconus (Agassiz, 1843) fin spine (SMF No. P419a), lateral. 2. H. longiconus (Agassiz, 1843) anterior tooth (SMF No. P415a), labial. 3. H. longiconus (Agassiz, 1843) lateral tooth (SMF No. P416f), labial. 4. Acrodus gaillardoti (Agassiz, 1837) lateral tooth (SMF No. P413c), occlusal. 5. Acrodus lateralis (Agassiz, 1837) tooth (SMF No. P414b), occlusal. 6. Polyacrodus polycyphus (Agassiz, 1837) lateral tooth (SMF No. P425), lateral. 7. Colobodus maximus Quenstedt, 1835 skull (SMF No. P1658, original to [89]), a. Lateral, b. Teeth enlarged. 8. Birgeria mougeoti (Agassiz, 1844) (SMF No. P431; original to [33]), a. Lateral, b. Teeth enlarged.

pestite beds are intercalated with the marls, and con- stones or are attached either to larger shell remains of sist mainly of the shells of terebratulids that originally bivalves, or to each other at their bases by holdfasts. The lived in a hardground palaeoenvironment (see terebrat- habitats of the crinoidal bioherms are not the crinoidal ulid hardground community described above). Many ver- limestone facies found at lower levels, which are bar de- tebrate remains have been found in, or directly below, the posits [52]. Between the crinoids, epifaunistic bivalves Coenothyris terebratulid shell-beds of these hardground settled on the firm- and hardgrounds [31], together with communities (Fig. 3), including skeletons and many holo- the oysters Newaagia noetlingi and Enantiostreon dif- types. forme. The small oyster Placunopsis ostracina, which is found in various facies types and on the shells of other Crinoidal bioherm communities (Fig. 2F) are not known organisms, is found in Bayreuth attached to other bivalve from the Bayreuth facies, but are very common on the shells. Mytilus eduliformis, Myalina blezingeri, and Pleu- steeper carbonate ramps of the Germanic Basin that are ronectites laevigatus often had a byssate mode of fixa- found in south-western and central Germany [31, 52, 72], tion. Spirorbis valvata microconchids were also cemented and are well presented in bioherm and facies models onto other shells, while regular short-spined “bioherm- [31, 52]. The Encrinus liliiformis crinoids mostly form adapted” sea urchins such as Serpianotiarias coaeva were colonies up to one metre in diametre on crinoidal float- vagile benthos. Plagiostoma striatum was an epifaunis-

18 Cajus G. Diedrich,

Figure 5. Placodus gigas (Agassiz, 1833) remains from the Bayreuth quarries (skeleton reconstruction from [49]). 1. Skull from Laineck, Oschen- berg (BSP no. A5 VII 1208; holotype of P. gigas in [3], original to [5, 34]), a. dorsal, b. lateral, c. ventral. 2. Lower jaw (cast) from Bindlach or Leineck (U-MO BT 5067.00), a. lateral, b. dorsal. 3. Skull of a large individual from the Bindlach Formation (Illyrian) of Bindlach (U-MO BT 13, syntype to [10], a. dorsal, b. lateral, ventral. 4. Lower jaw fragment from Bindlach or Leineck (SMF no. R.4110; original to [19]), a. lateral, b. dorsal. 5. Interclavicle from the Bayreuth quarries (SMF no. R29), dorsal. 6. Scapula from Bindlach, Bindlacher Berg (SNSD no. 16254a), dorsal. 7. Scapula from Bindlach, Bindlacher Berg (NMB without no.), dorsal. 8. Humerus from Hegnabrunn (SNSD no. 54583), ventral. 8. Femur of a middle aged individual from Bindlach, Bindlacher Berg (U-MO BT 672.00, original to [7]), ventral. 9. Anterior cervical vertebra from the Bayreuth quarries (SMF no. R2001a), a. lateral, b. cranial, c. dorsal. 10. Middle cervical vertebra from the Bayreuth quarries (SMF no. R2001), a. lateral, b. cranial, c. dorsal. 11. Last cervical to first dorsal vertebra from the Bayreuth quarries (SMF no. R275), a. lateral, b. cranial, c. dorsal. 12. Anterior dorsal vertebra (SMF no. R2001b), a. lateral, b. cranial, c. dorsal. 13. Middle dorsal vertebra from Bindlach, Bindlacher Berg (SNSD no. 54552), cranial. 14. Middle dorsal vertebra from Hegnabrunn (SNSD no. 59825), a. lateral, b. cranial. 15. Middle dorsal vertebra (SMF no. R2003), a. lateral, b. cranial, c. dorsal. 16. Middle dorsal vertebra centrum from Hegnabrunn (SMF no. R2004), a. cranial, b. dorsal, c. lateral. 17. Dorsal osteoderm bone from the Bayreuth quarries (SMF no. R2006), lateral. 18. Dorsal osteoderm bone from the Bayreuth quarries (SMF no. R2005), lateral. 19. Ilium from the Bayreuth quarries (SMF no. R2020), lateral. 20. Femur from Bindlach, Bindlacher Berg (U-MO BT 5067.00, original to [7]), ventral. 21. Femur from Bindlach, Bindlacher Berg (U-MO no Bayr 73), ventral.

19 The Middle Triassic marine reptile biodiversity in the Germanic Basin,in the centre of the Pangaean world

tic bivalve, as was the gastropod Naticella triadica [31]. this study the assemblage is placed above the crinoidal Brachiopods also settled on hardgrounds, but in smaller ramp facies in a more shallow environment, as in the Up- amounts. Within the crinoidal facies vertebrates are ex- per Muschelkalk model of Duchrow and Groetzner [71] for tremely rare in the fossil record. the terebratulid facies, or the Lower Muschelkalk model Tonplatten soft ground community (Fig. 2G): The marl- of Lukas [80]. stone and thin platy to banked limestone layers known There are several arguments in support of this latter inter- as “Tonplatten Facies” (for example [52]), which are par- pretation. Terebratulids needed clean and turbulent water ticularly evident in the Lainecker road cut section (Fig. (for example [71, 80]), a requirement that was best satisfied 2), are made up of lumachelle accumulations or micritic in the bar facies hardground areas and hence some ter- limestones - facies types that are very common all over ebratulids appear together with the crinoidal community the central parts of the Germanic Basin [52, 73]. The re- (Fig. 3F). Remains would have been washed in both direc- worked infauna of the underlying sediments consists of tions from the bar facies which has led to confusion in the the infaunistic Palaeonucula sp., Palaeoneilo elliptica, reconstruction of their real habitat, but the main drift was and Myophoria vulgaris, and the epifaunistic Plagios- towards the coast, which is documented in the terebratulid toma striata, Hoernesia socialis, Bakevillia costata and shell accumulations and terebratulid beds (= Terebratel- Entolium discites [31, 74] the gastropods Ampullina sp. bank”, 10 metres in thickness) especially in northwest- and Loxonema sp. probably fed on the substrate or on ern Germany where crinoid bioherms are missing, in the shell accumulations. All benthic and pelagic animals are Bissendorf shallow marine western Germanic Basin re- mixed together in the storm beds, along with ceratites that gion (cf. [71]). In those very shallow regions crinoids are form ceratite accumulations within the Germanic Basin nearly absent (only one trochitic bed in Bissendorf, see [31, 75–78]. Skeletons of vertebrates have been (rarely) [49]) being the result of a more shallow position of ter- found within such facies, including the famous skeleton of ebratulid colony settlements (3-5 metres, after [71]) such Placodus gigas [16, 62]. as crinoid colonies (5-10 metres, after [52]). The sections Placunopsis bioherm communities: These communities compared (Fig. 3) support this facies zonation – more described by Hagdorn [79] are typical of the uppermost abundant Terebratelkalke which are even more thick oc- Upper Muschelkalk in some regions, especially in the cur towards the coasts (Fig. 3, section Bayreuther locali- southern Germanic Basin. The small oyster Placunopsis ties and Bisendorf), while more crinoidal dominated Tro- ostracina is abundant in some beds (the ostracina Bed), chitenkalke and Tonplatten towards the basin (Fig. 3, sec- and can form stromatolite-like bioherm structures about tion Lamerden). Terebratulids are therefore more coastal, one metre in diametre (cf. [31]), but such bioherms have shallow habitat settlers such as crinoids. The sections at not been found in any of the Bayreuth sections. Bissendorf, Bindlach, and Hegnabrunn (Bayreuther sites) also provide evidence that the primary terebratulid habi- tat was above the crinoid “forests”, in less than about 5 4. Palaeontology metres of water depth (cf. [71], see Fig. 3), while in the Lower Muschelkalk even shallower conditions are inter- preted for terebratulid growth areas [80]. The brachiopods 4.1. Vertebrate taphonomy eventually accumulated in the tempestite shell deposits of the shallow marine carbonate ramp, as can be seen at The suggestion that the appearance of marine reptiles the Bayreuth sites, but also at Bissendorf, where those (predominantly facies-specific placodonts such as Pla- form massive coastal bar deposits [71]. In summary, the codus or Cyamodus) is indicative of macroalgae-rich terebratulid palaeocommunities are interpreted herein to palaeoenvironments [43, 56] is supported by evidence from have existed mainly above the crinoidal ramp on exten- the Germanic Basin vertebrate analyses (Fig. 15) and sive hardgrounds that formed in shallow water conditions, palaeoenvironmental models described herein, and in par- where crinoids were no longer able to exist. The brachio- ticular by the the Bindlach and Hegnabrunn palaeocom- pod shells were then transported during storm events and munities (Figs. 2-3). complete specimens and single valves formed secondary The placodonts appear mainly within the terebratulid- (allochthonous) accumulations, together with the verte- shell tempestites which are made up of brachiopods that brate bones of marine reptiles and actinopterygian and originated from the shallow marine hardgrounds of the shark remains, between the bar and the coastal palaeoen- terebratulid hardground community. This fanual assam- vironment (also Bissendorf, [49]). In some cases complete blage was placed in former times into the deep subtidal reptile carcasses were covered by such accumulations (for zone below the crinoidal ramp facies [31, 52]. However, in example, N. mirabilis and P. longaevus skeletons).

20 Cajus G. Diedrich,

Figure 6. Cyamodus remains from the Bayreuth quarries (skeleton reconstruction from [56]). 1. Cyamodus rostratus (Münster, 1839) skull from Bindlach, Bindlacher Berg (U-MO no. BT 748, holotype to [4]), a. dorsal, b. ventral, c. lateral. 2. Half lower jaw from Bindlach, Bindlacher Berg (SBMF no. 4040, original to [37]), a. lateral inner view with large posterior replacement tooth, b. dorsal, c. lateral outer view. 3. Cyamodus münsteri (Münster, 1830) skull of a young individual with false implanted teeth from Bindlach, Bindlacher Berg (UM-O BT 1210a, holotype to [5], a. dorsal, b. lateral, c. ventral. 4. C. münsteri [5] of an adult individual, with completed cast part from Bindlach, Bindlacher Berg (NHM no. R.1644, holotype to C. laticeps [35]), a. dorsal, b. ventral, c. lateral. 5. C. münsteri [5] lower jaw from Bindlach, Bindlacher Berg (UM-O BT 2172, original to [37]), a-b. photo and redrawing, dorsal, c. photo lateral. 6. Carapax fragment from Hegnabrunn (SMNS no. 81600), dorsal.

4.2. Vertebrate biodiversity

Reptile remains: The historical vertebrate bone collec- The Bayreuth sections produced many fish scales and tions mostly comprise of reptile remains (about 95%). For shark teeth, and although these can be easily found today qualitative analyses, the most important published (and in the bone bed outcrops they are poorly represented in also many unpublished) postcranial specimens is pre- the historical collections, making up only about 5% of the sented herein (Figs. 5-14). Many incorrect identifica- vertebrate remains. A selection of the species represented tions of postcranial bones are corrected, and other previ- at Bayreuth is described below and compared to material ously un-identified bones have now been identified. After from Monte San Giorgio, and anterior or lateral teeth from more than 150 years the Bayreuth bone collection and Hybodus longiconus [3] (Figs. 4.1-3), an Acrodus gaillar- the Meyer [8] monograph have thus been fully revised, doti [3] lateral tooth (Fig. 4.4), an Acrodus lateralis [3] including the following material. lateral tooth (Fig. 4.5), a Polyacrodus polycyphus [3] lat- eral tooth (Fig. 4.6), and Palaeobates angustissimus [3] Placodus gigas (Agassiz, 1833): Skeletons and many sin- teeth. gle bones from Bayreuth and other German sites have been reviewed [17] and material from the Bayreuth sites Fish remains: Some actinopterygian fish remains are compared to the only known, (almost) complete skeleton original specimens and holotypes, such as Birgeria of P. gigas in the SMF [16]. The holotype skull (Fig. mougeoti [33]; Fig. 4.8, which were not redescribed and 5.1, BSP no. A5 VII 1208) has subsequently been re- included in a recent revision of the Monte San Giorgio illustrated by Broili [91], Sues [46], and Rieppel [10]. More Birgeria specimens [88], or Colobodus frequens [32, 89, 90] then 25 known skulls and many lower jaws (illustrated in Fig. 4.7). There are also some more jaw fragments (sple- [17]), more then 600 individual teeth, and some hundreds nial, vomer) from C. frequenz, in various different collec- of postcranial bones, are known from this species in Ger- tions. many, mainly from the Bayreuth exposures. A selection of

21 The Middle Triassic marine reptile biodiversity in the Germanic Basin,in the centre of the Pangaean world

Figure 7. Lariosaurus and Neusticosaurus remains from the Bayreuth quarries (Neusticosaurus skeleton reconstructions from [49], Lariosaurus buzli [1] after a skeleton cast from the Comer Lake of Perledo, Italy, in the BSP). 1. Lariosaurus cf. buzli [1] skull from Bindlach, Bindlacher Berg (SMF No. R4572, original to [7]), a. dorsal, b. lateral, ventral. 2. L. cf. balsami [1] humerus from Bindlach, Bindlacher Berg (SMF no. R784), ventral. 3. L. cf. buzli [1] coracoid Bindlach, Bindlacher Berg (UM-O without no.), ventral. 4. Lariosaurus cf. balsami [1] ischium from Bindlach, Bindlacher Berg (UM-O no. BT-670; original to [7]), dorsal. 5. L. cf. buzli [1] femur from Bindlach, Bindlacher Berg (UM-O no. BT-722), dorsal. 6. L. cf. buzli [1] middle dorsal vertebra centrum (SMF no. R951), a. cranial, b. dorsal, c. lateral. 7. L. cf. bazii [1] posterior dorsal vertebra centrum (SMF without no.), a. cranial, b. dorsal, c. lateral. 8. Neusticosaurus sp. from Bindlach, Bindlacher Berg (NMB without no.), a. cranial, b. lateral. 9. Neusticosaurus sp. femur from Bindlach, Bindlacher Berg (NMB without no.), cranial.

cranial and postcranial bones from the Bayreuth localities have been the case in the Bayreuth region. have been illustrated herein (Fig. 5) in order to demon- strate their anatomical differences from the bones of other Cyamodus rostratus (Münster, 1839) and Cyamodus reptiles. Very characteristic among the postcranial bones muensteri (Agassiz, 1839: Several skulls of Cyamodus are the pectoral (interclavicula: Fig. 5. 5, scapula: Figs. (known as “Schnabelplacodus” in German) have been de- 5.6-7) and pelvic elements (ilium: Fig. 5.19), as well as scribed from the Bindlach localities (compilation in [56]), the procoel vertebrae centrae that are typical only in pla- of which the most important are illustrated in Figure 6. All codonts (Fig. 5.16), with their two large lateral processi of these skulls from the earlier species (atavus to evolutus on the cervical vertebrae (Figs. 5.9-10) and up to 6 cm zones) of Cyamodus rostratus [4] (U-MO no. BT 748) are long lateral processi on the dorsal vertebrae (Figs. 5.11- holotypes and originals described by Owen [35] (NHM 15). Also characteristic are the triangular-shaped osteo- no. R.1644, holotype to “P. laticeps”), or Drevermann [37] derm bones (Figs. 5.17-18), which are above the verte- lower jaw, UM-O BT 2172). The younger species (evolu- bral column. The humeri are distally extended (Fig. 5.8), tus to spinosus zones) of Cyamodus münsteri [5] is repre- whereas the femorae have a well developed trochanter sented by an old animal (Fig. 6.4, UM-O BT 1210a) and a (Fig. 5.20-21). Different sizes are all well represented in solitary juvenile skull (Fig. 6.3, NHM no. R.1644). Only the Bayreuth material, indicating the presence of a large postcranial bones are known from Bayreuth, including a population and providing material suitable for ontogenetic carapace fragment from Hegnabrunn (Fig. 6.6, SMNS no. studies [17]. These diving reptiles have been referred to 81600) that has also previously been used in various de- as a “Triassic sea cows” because they appear to have scriptions of the placodont armour ([38], [92]). fed on macroalgae meadows [49, 56] in shallow marine Neusticosaurus sp.: The remains of a small pachypleu- macroalgae-rich palaeoenvironments, as also appears to rosaur have previously been illustrated by [7], but only

22 Cajus G. Diedrich,

Figure 8. Nothosaurus mirabilis Meyer, 1839 remains from the Bayreuth quarries (N. mirabilis skeleton reconstructions from Diedrich [49], modified here in the extremities, pelvic and pectoral girdle; skeleton based after the N. mirabilis holotype skeleton from Meyer [34], see. Fig. 30). 1. Skull from Bindlach, Bindlacher Berg (UM-O without no.), a. dorsal, b. lateral, c. ventral. 2. Skull from Bindlach, Bindlacher Berg (UM-O without no.), a. dorsal, b. lateral. 3. Skull from Hegnabrunn (SNSD no. 59074, original [60]), a. dorsal, b. ventral. 4. Lower jaw from Bindlach, Bindlacher Berg (UM-O without no.), dorsal. 5. Pectoral girle (interclavicle, clavicles, scapulae) from Bindlach, Bindlacher Berg (UM-O without no.), ventral. 6. Coracoid from Bindlach, Bindlacher Berg (SMF no. R31), ventral. 7. Humerus of a young animal from a Bayreuth quarry (SMF no. R676), dorsal. 8. Humerus of the skeleton from Laineck, Oschersberg (UM-O BT 1000a, N. mirabilis holotype to Meyer [34]), dorsal. 9. Humerus of the skeleton from Bindlach, Bindlacher Berg (UM-O without no.), dorsal. 10. Radius from Bindlach, Bindlacher Berg (UM-O without no.), lateral. 11. Ulna from Bindlach, Bindlacher Berg (NMB without no.), lateral. 12. Ulna of the skeleton from Laineck, Oschersberg (UM-O BT 1000b, N. mirabilis holotype to Meyer [34]), lateral. 13. Anterior cervical vertebra from Hegnabrunn (SNSD no. 84752), a. cranial, b. lateral. 14. Middle dorsal vertebrae of the skeleton from Laineck, Oschersberg (UM-O BT 1000c, N. mirabilis holotype to Meyer [34]), lateral. 15. Middle dorsal vertebra centrum from Hegnabrunn (SNSD no. 83899), a. cranial, b. dorsal. 16. Posterior dorsal vertebra centrum from Hegnabrunn (SNSD no. 83887), a. cranial, b. dorsal. 17. Sacral vertebra from Hegnabrunn (SMF no. R1064), a. cranial, b. lateral. 18. Anterior caudal vertebra centrum from a Bayreuth quarry (SMF no. R765), ventral. 19. Middle caudal vertebra from Bindlach, Bindlacher Berg (SNSD no. 54070), a. cranial, b. lateral. 20. Posterior caudal vertebra from Bindlach, Bindlacher Berg (SNSD no. 54069), a. cranial, b. lateral. 21. Femur of the skeleton from Laineck, Oschersberg (UM-O BT 1000d, N. mirabilis holotype to Meyer [34]), lateral. 22. Nothosaurus mirabilis [4] femur from Bindlach, Bindlacher Berg (UM-O no. BT 679), cranial. 23. Tibia from a Bayreuth quarry (SMF no. R1066, original to [7]), ventral. 24. Tibia from a Bayreuth quarry (NMB without no.), cranial. 25. Fibula, five metatarals and two phalanx in articulation of the skeleton from Laineck, Oschersberg (UM-O BT 1000e, N. mirabilis holotype to Meyer [34]). 26. Pubis from Bindlach, Bindlacher Berg (UM-O without no.), ventral. 27. Ischium of the skeleton from Laineck, Oschersberg (UM-O BT 1000f, N. marchicus holotype to Meyer 23 [34]), ventral. 28. Ischium from Bindlach, Bindlacher Berg (UM-O without no.), ventral. 29. Skeleton from Laineck, Oschersberg (UM-O BT 1000, N. mirabilis holotype to Meyer [34]) which has a composed skull and one larger incorrect humerus of other individuals. The Middle Triassic marine reptile biodiversity in the Germanic Basin,in the centre of the Pangaean world

Figure 9. Paranothosaurus giganteus (Münster, 1934) skulls from the Bayreuth quarries (P. giganteus skeleton reconstructions from [49], modified here in the extremities; skeleton based after “P.amsleri Peyer, 1939” (P.giganteus) skeleton from the Monte San Giorgio). 1. P.giganteus [2] skull (cast) of a very large specimen from Bindlach, Bindlacher Berg (SMF no. R471, ”Nothosaurus baruthicus” holotype to [45]), a. dorsal, b. original ventral, c. original lower jaw. 2. P. giganteus [2] skull of a medium sized individual from Bindlach, Bindlacher Berg (UM-O without no., “Nothosaurus andriani” holotype to Meyer [34]), ventral.

a small amount of material is present in the collections direct comparisons with two mounted skeleton casts in the from the Bayreuth sites. Nearly all of this material con- SMF and SNSD exhibitions. Simosaurus appears to be sists of cervical to caudal vertebra centra (Fig. 7.8), to rarer in the Bayreuth localities resulting from the missing which can now be added a single femur (Fig. 7.9). A outcropping upper parts of the Upper Muschelkalk (Bind- clear assignation to a particular species remains prob- lach and Hegnabrunn sites). Following comparisons with lematic, but skeletons described from Monte San Giorgio the listed collections most of the remains, such as the lost ([93–97]) and Germany [87] need to be taken into account, holotype described by Meyer [99] from Luneville (France, especially those of N. pusillus [93]. see [58]), appear to be from the upper part of the Upper Muschelkalk (Longobardian). Two humeri (Figs. 11.1-2), Simosaurus cf. gaillardoti Meyer, 1842: Only a relatively an ulna and a single radius (Figs. 11.3-4), a femur (Fig. small number of postcranial bones have been identified 11.6), and a fibula and tibia (Figs. 11.7-8) are the extrem- (Fig. 11) within the large SMF, UM-O and MB collec- ity bone elements identified from Bayreuth sites. Even tions, using the original skeleton finds [98] and descrip- vertebrae are rare, and only one typical dorsal vertebra tions of cranial and postcranial elements [58], as well as

24 Cajus G. Diedrich,

Figure 10. Paranothosaurus giganteus (Münster, 1934) postcranial remains from the Bayreuth quarries 1. Coracoid of a large animal from a Bayreuth quarry (SMF no. R33), ventral. 2. Coracoid of a medium-sized animal from Bindlach, Bindlacher Berg (UM-O without no., original to [7]), dorsal. 3. Scapula from a Bayreuth quarry (SMF no. R36), dorsal. 4. Humerus of a large animal from a Bayreuth quarry (UM-O no. BT 687, original to [7]), dorsal. 5. Humerus of a medium-sized animal from a Bayreuth quarry (SMF no. R24), dorsal. 6. Humerus of a medium-sized animal from a Bayreuth quarry (SMF no. R27), dorsal. 7. Ulna of a large animal from Bindlach, Bindlacher Berg (UM-O no. BT 673, original to [7]), ventral. 8. Anterior cervical vertebra centrum from a Bayreuth quarry (SMF no. R514), a. dorsal, b. cranial, c. lateral. 9. Anterior cervical vertebra from a Bayreuth quarry (SMF no. R47), a. dorsal, b. cranial, c. lateral. 10. Posterior cervical to anterior dorsal vertebra centrum from a Bayreuth quarry (SMF no. R514), a. dorsal, b. cranial, c. lateral. 11. Posterior cervical to anterior dorsal vertebra from a Bayreuth quarry (SMF no. R871), a. cranial b. lateral, c. dorsal. 12. Dorsal vertebra from Hegnabrunn (SNSD no. 1874), a. cranial, b. dorsal. 13. Dorsal vertebra from a Bayreuth quarry (SMF no. R4556), a. cranial, b. dorsal. 14. Dorsal vertebra neural arch from a Bayreuth quarry (SMF no. R938), a. cranial, b. dorsal. 15. Dorsal vertebra centrum from Hegnabrunn (SNSD without no.), a. dorsal, b. cranial. 16. Ilium from a Bayreuth quarry (SMF no. R45), lateral. 17. Pubis from Bindlach, Bindlacher Berg (UM-O without no., original to [7]), dorsal. 18. Femur from Bindlach, Bindlacher Berg (UM-O no. BT 677, original to [7]), lateral. 19. Tibia from a Bayreuth quarry (SMF no. R1041), ventral.

can be illustrated (Fig. 11.5). The Simosaurus skeleton the first time (Fig.8). A third, too large, humerus and an reconstruction used was taken from Diedrich [49]. The incorrectly identified femur were originally included in a swimming style of the animal is reported to be a combi- plaster jacket (Fig. 8.28). It was possible to determine nation of axial swimming supported by a rowing action of several single bones during the preparation process (es- the limb extremities [58]. pecially in the UM-O coll.), which allowed them to be separated out from the Paranothosaurus giganteus (Figs. Nothosaurus mirabilis Münster, 1834: Remains from this 9-10), Pistosaurus longaevus (see details and bone list large nothosaur are presented herein, including the redis- in [21]), Simosaurus gaillardoti (Fig. 11) and placodont covered paratype skeleton from the Oschersberg location (Figs. 5-6) material, all of these being larger animals. at Laineck (Fig. 8.28, UM-O with no number). Some The vertebral column of the N. mirabilis holotype skele- important single newly-prepared bones are illustrated for

25 The Middle Triassic marine reptile biodiversity in the Germanic Basin,in the centre of the Pangaean world

(a) (b)

Figure 11. Simosaurus gaillardoti Meyer, 1842 remains from the Bayreuth quarries (S. gaillardoti skeleton reconstruction from [49]). 1. Humerus from a Bayreuth quarry (SMF no. R26, original to [7]), ventral. 2. Humerus from Bindlach, Bindlacher Berg (UM-O no. BT 689, original to [7]), ventral. 3. Ulna from a Bayreuth quarry (MB no. I.039.17, original to [7]), ventral. 4. Radius from Bindlach, Bindlacher Berg (UM-O without no.), ventral. 5. Dorsal vertebra centrum from Bayreuth quarry (MB no. I.092.01), a. cranial, b. dorsal, c. lateral. 6. Femur from a Bayreuth quarry (SMF no. R872), ventral. 7. Fibula from a Bayreuth quarry (SMF no. R85), ventral. 8. Tibia from a Bayreuth quarry (SMF no. R95), ventral.

ton starts with the first cervical vertebra, and includes genera are synonymous, which would be a similar situa- all of the dorsal (middle dorsal, Fig. 20.13) and half of tion to that of Lariosaurus buzzii and Paranothosaurus gi- the caudal vertebra. Pectoral bones, pelvic elements (is- ganteus. It seems likely that Nothosaurus mirabilis might chium, Fig. 8.26), forelimb bones (humerus, Fig. 8.8) and be present in the Monte San Giorgio fauna because it is hind limb bones (femur, Fig. 8.20) have been preserved, also known from elsewhere in the north-western Tethys, allowing minor modifications or corrections to be made at Siles in Spain [101]. Following this new revision of the to the skeleton reconstruction presented previously [49]. Illyrian-Fassanian reptiles most of the marine reptile gen- Anatomical comparisons between this skeleton reconstruc- era found in Monte San Giorgio can now be described as tion and the “Ceresiosaurus calcagnii [100] of Monte San present also in the Germanic Basin, which was connected Giorgio appear to support the suggestion that these two to the north-western Tethys thus allowing an easy faunal

26 Cajus G. Diedrich,

Figure 12. Pistosaurus longaevus Meyer, 1939 remains from the Bayreuth quarries. 1. Skull (cast) from Bindlach, Bindlacher Berg (MB, no. R.47, holotype to “Pistosaurus grandaevus Meyer, 1847-55). a. dorsal, b. lateral left, c. ventral. 2. Skull from Bindlach, Bindlacher Berg (UM-O no. BT-682; holotype to [7]), a. dorsal, b. lateral left, c. ventral. 3. Scapula of the paratype skeleton from Bindlach, Bindlacher Berg (SMF no. 4041-4), ventral. 4. Coracoid of the paratype skeleton remain from Bindlach, Bindlacher Berg (SMF no. 4041-1), ventral. 5. Humerus from Bindlach, Bindlacher Berg (SMF no.R.2011), lateral and joint views. 6. Humerus of the skeleton from the Bindlach Formation (Illyrian) of Bindlach (SMF no. 4041-5), dorsal. 7. Radius of the paratype skeleton from Bindlach, Bindlacher Berg (SMF no. 4041-9), ventral. 8. Ulna of the paratype skeleton from Bindlach, Bindlacher Berg (SMF no. 4041-7), dorsal. 9. Ulnare of the paratype skeleton from Bindlach, Bindlacher Berg (SMF no. 4041-10). 10. Carpalia of the paratype skeleton from Bindlach, Bindlacher Berg (SMF no. 4041-12). 11. Carpalia of the paratype skeleton from Bindlach, Bindlacher Berg (SMF no. 4041-13). 12. Anterior cervical vertebra from Bindlach, Bindlacher Berg (SMF no.875.f), a. cranial, b. lateral. 13. Middle cervical vertebra from Bindlach, Bindlacher Berg (SMF no. 875.b), a. dorsal, b. cranial, b. lateral. 14. Dorsal vertebra from Hegnabrunn (SMNS no. 84827). a. cranial, b. lateral.15. Sacral vertebra from Bindlach, Bindlacher Berg (SMF no. R276), cranial. 16. Middle caudal vertebra from the Bindlach Formation (Illyrian) of Bindlach, Germany (SMF no. R 4611), a. cranial. b. lateral. 17. Articulated ilium and ischium from Bindlach, Bindlacher Berg (SMF no. R76), dorsal. 18. Ilium from Bindlach, Bindlacher Berg (NMB BT-003065.01), ventral. 19. Pubis from Bindlach, Bindlacher Berg (UM-O No. BT 675, original to [7, 12], holotype to “Delphinosaurus” in [36]). 20. Femur from Bindlach, Bindlacher Berg (U-MO no. BT 5375). 21. Tibia from Bindlach, Bindlacher Berg (UM-O No. BT 683, original to [7]), ventral. 22. Fibula from Bindlach, Bindlacher Berg (SMF no. R 71), ventral. 23. P. longaevus [6] skeleton from Bindlach, Bindlacher Berg (SMF no. 4041, original to [45]).

27 The Middle Triassic marine reptile biodiversity in the Germanic Basin,in the centre of the Pangaean world

Figure 13. Ichthyosaur remains from the Bayreuth quarries. 1. ?Ichthyosauria indet. coracoid from Bindlach, Bindlacher Berg (UM-O without no.), dorsal. 2. Shastasaurus sp. ischium from Bindlach, Bindlacher Berg (UM-O no. 709; original to ”Placodus” in [7]), ventral. 3. Shastasaurus sp. dorsal vertebra (SMF no. R284, holotype to “Cymbospondylus” in [36]), a. cranial, b. lateral. 4. Ichthyosauria indet. radius from Leineck, Oschersberg (UM-O no. 6777), dorsal. 5. Omphalmosaurus sp. dorsal vertebra (UM-O no. Bayr-67), a. cranial, b. lateral.

Figure 14. Tanystrophaeus conspicuus Meyer, 1847 remains from the Bayreuth quarries (skeleton reconstruction from [49]). 1. Anterior tooth from Bindlach, Bindlacher Berg (UM-O no. Bayr-65), lateral. 2. Third cervical vertebra of a younger animal from Bindlach, Bindlacher Berg (UM-O no. BT 732, Holotype to [7], original to [47]), lateral. 3. Fifth cervical vertrebra from Bindlach, Bindlacher Berg (UM-O without no., original to [47]), lateral. 4. Ninth cervical vertebra from Bindlach, Bindlacher Berg (UM-O without no., original to [47]), lateral. 5. Tenth cervical vertebra from Bindlach, Bindlacher Berg (UM-O no. BT 736, original to Huene [47, 162]), lateral. 6. Eleventh cervical vertebra from Bindlach, Bindlacher Berg (SMF no. R285, original to Huene [47, 162]), lateral. 7. Anterior thoracic vertebra from Bindlach, Bindlacher Berg (UM-O no. BT 741, original to Huene [162]; [47]), lateral-mirrored. 8. Middle thoracic vertebra from Bindlach, Bindlacher Berg (SMF no. R820, original to Huene [47, 163]), lateral. 9. Middle thoracic vertebra from Bindlach, Bindlacher Berg (SNSD no. 54690, original to [47]), lateral-mirrored.10. Anterior caudal vertebra from Bindlach, Bindlacher Berg (SMF no. R4092, original to [47]), lateral. 11. Middle caudal vertebra from Bindlach, Bindlacher Berg (UM-O no. BT 2171, Original to [47]), a. cranial, b. lateral, c. dorsal. 12. Middle caudal vertebra from Bindlach, Bindlacher Berg (SMF no. R282, holotype to Huene [162]= Thecodontosaurus latespinatus,[47]), lateral-mirrored. 13. Distal caudal vertebra from Bindlach, Bindlacher Berg (SNSD no. R287, original to [47]), lateral. 14. Left humerus of a medium-sized animal from Bindlach, Bindlacher Berg (UM-O No. Bayr-66), cranial. 15. Left humerus from Bindlach, Bindlacher Berg (SMF no. R4034, original to Huene [47, 163]), cranial.

28 Cajus G. Diedrich,

Figure 15. Palaeoenvironmental reptile group assemblage analyses of ichthyosaurs (open marine), pistosaurs (open marine), placodonts (shallow marine macroalgae-rich), nothosaurs (shallow marine to lagoonal), pachypleurosaurs (lagoonal), askeptosaurs (lagoonal), amphibians (fluvial), terrestrial (beach to land). A. Bissendorf after [49]), B. Lamerden, and C. Bayreuth, from new results.

29 The Middle Triassic marine reptile biodiversity in the Germanic Basin,in the centre of the Pangaean world

interchange. of similar age from Bad Sulza that was mis-identified as “N. marchicus” in Rieppel and Wild [60], which is in fact Paranothosaurus giganteus (Münster, 1834): a similar Lariosaurus skull [107]. Also from Bad Sulza, This species is based on the holotype skull fragment from several postcranial bones with similar proportions, from Bindlach (from one of the Bindlacher Berg quarries, UM- all parts of the body, again prove the presence of a large O no. BT 685). The largest skull ever found with its lower Lariosaurus (cf. buzzii[1]) in the Germanic Basin during jaw intact is described by Geissler [45], ”Nothosaurus the late Illyrian to Fassanian. In contrast, N. marchi- baruthicus”, SMF no. R471; Fig.9.1), which was also cus has only been positively identified from the Bithy- found at the Bindlacher Berg near Bayreuth. Many sin- nian to the early Illyrian in the Germanic Basin, during gle postcranial bones in the UM-O and SMF collections the upper Lower/basal Middle Muschelkalk (late Pelso- were re-sorted and some from other collections were also nian). With the subsequent transgression Nothosaurus identified during the course of this research, mainly using appeared as a more highly evolved form at the end of the skeleton from Monte San Giorgio Paranothosaurus the Middle Muschelkalk [104], as has recently been de- amsleri [102] for comparison. This revision of the sin- scribed from a large quantity of cranial and postcranial gle bones and skeletons has resulted in a first ”bone material from Bad Sulza [107]. Without details of the element overview” for this, the largest of the nothosaur stratigraphic occurrences, the skull determinations, or the species (Figs. 9-10). The history of this skeleton from study of all postcranial bone remains from the Bad Sulza Monte San Giorgio has again revealed the use of invalid and Bayreuth sites, previous authors have not been able and synonymous species names at the Monte San Giorgio recognize this genus [10], which was in fact widely dis- Swiss/Italian site, which has previously been discussed for tributed in the north-western Tethys and the Germanic the skulls by Rieppel and Wild [60]. However, rather than Basin from the Illyrian to the Longobardian. The claims following the suggestion by Rieppel and Wild [60] to use of Monte San Giorgio endemism [10] must therefore now the genus name “Nothosaurus”, the Paranothosaurus [102] be revised. genus name is preferred because of the very strong cra- nial and postcranial anatomic differences between those Pistosaurus longaevus Meyer, 1839: The previously il- two, as again demonstrated herein for all body parts, re- lustrated skeleton reconstruction [49] has been modified vealing very different bone and skeleton anatomies (Figs. following a new and comprehensive review [21]. As well 8-10). The main difference is in the vertebrae, which have as the holotype skull from Bindlach (Fig. 12.2, [7]), many short spines in P. giganteus [58] including short cervical to single bones have been re-identified during the study of dorsal vertebra centra with extended zygapophyses (Figs. the paratype skeleton [87], which was found in the SMF 9.8-15), which would only have allowed a paraxial type of collection under the name “Nothosaurus strunzi” (SMF locomotion and prevented any rowing action of the limb no. 4041), as described by Geissler [45] and, in some se- extremities. Bones from all parts of the body are repre- lected parts, by Sues [92]. More then 200 single bones, sented from the Bayreuth sites, and an articulated skele- mostly from the Bayreuth localities were included with ton from Bayreuth remains unpublished (coll. SNSD); the the study of this postcranial skeleton (Fig. 12.23), allow- skeleton reconstruction based on “P. amsleri [103]” in [49] ing a detailed anatomical atlas to be prepared [21], from was re-used, but once again with slight modifications, in which only some selected material from the Bayreuth sites this case on the limbs and in particular the ulna/radius. is illustrated herein (Figs. 12). The locomotion of this open-marine adapted reptile was reconstructed from the Lariosaurus cf. buzzii Tschanz, 1989 : The skull (Fig. 7.1) skeletal anatomy, especially the primitive paddles which illustrated in Edinger [104] as Nothosaurus sp., and iden- allowed “primitive subaquatic flying” [21]. tified by Rieppel and Wild [60] as “N. marchicus” is revised here to Lariosaurus cf. buzzii [1] because the skull shape, Shastasaurus sp.: In contrast to the sauropterygian rep- nasar, orbit and parietal foramina shapes and measure- tilian diving forms and the rich ichthyosaur fauna from the ments match more closely to those of Lariosaurus than north-western Tethys of Monte San Giorgio (for example to those of Nothosaurus. In addition to the cranium, the [96, 97, 108–111], the very limited amount of German postcranial remains first identified herein, which include a ichthyosaur material present within the Bayreuth material humerus, coracoid, femur and ischium, and also possibly a [36] consists of just a few bones. One non-prepared lower dorsal vertebra centra (Figs. 7.2-7), are quite character- jaw was not possible to re-figure, instead the dorsal istic of Lariosaurus when compared to a skeleton cast in vertebra of the type specimen of “Cymbospondylus” is the BSP (from Perledo, Italy) and the descriptions of Cu- figured ([36], Fig. 13.3). The ischium described by rioni [1] and Ticli [105] or Tschanz [106], and thus support Huene [36] (Fig. 13.2) can now be compared to the first the presence of this genus, as would be expected, in the complete Shastasaurus skeleton from the Longobardian Germanic Basin. Other new evidence comes from a skull of China [112], and its attribution to Shastasaurus can

30 Cajus G. Diedrich,

thus be validated, but the species attribution of the gio) and shallow marine coastal subtidal deposits (for ex- single bone must remain open, even though it is almost ample at the Bayreuth sites). identical in shape. The “Delphinosaurus” holotype scapula determined by Huene [36] has been revised to be a pubis bone from Pistosaurus longaevus ([21], Fig. 12.19). The Shastasaurus species described globally from the Triassic are S. carinthiacus [108], nomen dubium) 5. Discussion from the Austrian Alps, S. neubigi ([113], nomen dubium) from Bavaria in Germany, S. pacificus from California and Mexico in North America [114], and S. tangae from 5.1. Stratigraphy Guizhou Province in China [109, 112]. Few discoveries in Germany do not yet allow a clear species attribution. The Upper Muschelkalk sections studied in the Bayreuth area have been subdivided according to the international chronostratigraphy proposed by Bachmann and Kozur [54], Omphalosaurus sp.: A small ichthyosaur radius (Fig. Kozur and Bachmann [125]. The lithostratigraphical sub- 13.4) and a dorsal vertebra (Fig. 13.5), which possibly division in part follows Hagdorn and Simon [126] and originated from Omphalosaurus sp. compared to skeletal Bachmann et al. [127], the fossil marker horizons mainly material from North America (cf. [115–117]), can here be after Hagdorn and Simon [67], and the ceratite stratig- added to the fauna. The rareness of ichthyosaurs in the raphy after Rose [128], updated in Hagdorn [79]. Two Germanic Basin supports the facies interpretations of gen- new formation names (stratotypes) are introduced herein erally shallow marine habitats, even in the basin centre, with the Bindlach and Hegnabrunn formations distin- during the Upper Muschelkalk. Omphalosaurus was dis- guished by their markedly different facies types. The tributed in the northern hemisphere [118, 119] and seems sections of Lainecker Höhenzug have been compared to to have only rarely migrated into the Germanic Basin. those from Lamerden in the more deep basinal position Tanystrophaeus conspicuus Meyer, 1847: The terrestrial [87], Bissendorf in the marginal western shallow basin lepidosaur Tanystrophaeus conspicuus [7] holotype was position [129], and Bad Sulza in similar position as the based on a cervical vertebra from Bindlach (Fig. 14.2, Bayreuther sites [44] in Germany, and with their facies UM-O no. BT 732, see [120]). No skulls for this ter- interpretations [67, 79, 126, 130, 131] using the ceratite restrial reptile have yet been found in the Germanic biozonations which are not restricted to particular facies Basin, but teeth found in the Bayreuth area are illus- [31, 47, 50, 128, 132–136]. trated herein (Fig. 14.1), together with an overview of selected postcranial material (Fig. 14.2-15) that was re- described by Huene [121] and also to a large extent by 5.2. Bone bed dating Wild ([47, 120, 122]), including a femur (Fig. 14.15) and a previously unidentified humerus from a young animal The Bayreuth bone beds at Bindlach can be dated through (Fig. 14.14). The neck and limb positions in former skele- comparisons with the above-mentioned ceratite biostratig- ton reconstructions [123, 124] have been modified in the raphy, mainly following Rhode [132], and Hagdorn [79], skeleton anatomy, neck and limb positions by Diedrich and with other recently studied and correlated sections [49], and the locomotion adapted to match Synaptichnium in Bissendorf and Lamerden ([28, 133]; Fig. 3). They fall footprints which have recently been interpreted to have into three ceratite biozones between the atavus, pulcher, been produced by Tanystrophaeus [28]. Tanystrophaeus robustus, and compressus zones, while the “Spiriferina is quite common in the Bayreuth sites and it is believed Bed” which is well known in southern Germany ([137]), to have been a coastal hunter, as was T. longibardicus is possibly present at the top of the Bindlach 1 section. [47] from Monte San Giorgio. New interpretations suggest The historical bone material was not all collected from that Tanystrophaeus may have been adapted for “beach one layer or from a single bone bed (there are at least fishing”, with the neck elongation trend within the genus three main bone beds, Figs. 2-3) and spans a period of (T. antiquus to T. longibardicus,[47]) being explained as a about 2.5 million years in the late Illyrian and Fassa- response to changes in the beach morphology from exten- nian (chronostratigraphy after [125]). These bone beds sive shallow intertidal flats during the Lower Muschel- are all much older than the famous Muschelkalk/Keuper kalk (T. antiquus) to steeper, less extensive beaches in bone beds of southern Germany, well known from Crail- the Upper Muschelkalk (T. conspicuus)[28]. The presence sheim, which are from the Fassanian/Longobardian bound- of coastal-dwelling animals resulted in their taphonomic ary [138, 139] which correspond herin to the Bayreuther record in coastal lagoons (for example at Monte San Gior- boundary bonebeds in the road cut section.

31 The Middle Triassic marine reptile biodiversity in the Germanic Basin,in the centre of the Pangaean world

Figure 16. A. Main reptile types in the Middle Triassic world during the Illyrian (Middle Triassic northwestern Tethys composed after: [26]; ichthyosaurs distribution from Sander and Mazin, [109]; intertidals Germanic Basin after [28]; other sauropterygian distributions from [28, 62]; pistosaur distribution from [21]; archosaur migrations from [66]). B. Horseshoe crab reproduction migrations into the Germanic Basin appear to have resulted in large-scale food-chain reactions for terrestrial and marine reptiles. Thecodont archosaurs appear to have migrated north-south on the land massive and ichthyosaurs and pistosaurs must have interchanged east-west, whereas pla- codonts were limited to the shallow marine coastal areas and intracratonic basins along the Pangaean-Tethys coastlines within the tropics (Middle Triassic globe and Europe after [26]; intertidal zones of the Germanic Basin after [28]).

The Hegnabrunn section includes the evolutus and kalk nodosus bone bed is the last marine layer with a spinosus bone beds and contains bone remains that are marine reptile fauna and is followed by several thin bone younger than those from Bindlach (and possibly Laineck). beds at the Muschelkalk/Keuper boundary (cf. Figs. 2- The evolutus bone bed is also distributed in the Weserber- 3) in whose brackish influenced and lagoonal facies the gland of north-western Germany, north-east of the Rhen- first limnic amphibian remains appear (cf. amphibians in: ish Massif [78]. This section at Hegnabrunn covers a pe- [24, 25, 31, 140]). Within about the following 1 million riod of about 1.0 million years (after [125]). years more bone beds developed because of the regres- sion of the Germanic Basin and more common repetitions In the Laineck road cut the uppermost Upper Muschel- of marginal facies conditions (for example [31, 129]. These

32 Cajus G. Diedrich,

Figure 17. Palaeoenvironmental change and reptile exchange in the Middle Triassic Germanic Basin. A. With the new lower Upper Muschelkalk transgression the placodonts and nothosaurs appeared, together with macroalgae meadows, in the central Germanic Basin. B. With the deepening and maximum flooding of the middle Upper Muschelkalk crinoid bioherms typically developed along steep ramps in central and southern Germany, but in the Bayreuth region macroalgae meadows of a shallow marginal facies were still present, providing perfect feeding areas for placodonts. C. In the late Upper Muschelkalk the crinoid bioherms and macroalgae meadows disappeared, as did the adapted reptile fauna, to be replaced by forms adapted to lagoonal conditions, together with limnic amphibians. D. Brackish lagoons in the relict Germanic Basin / Hesse Depression allowed only a few adapted turtle-like placodonts and some other sauropterygian reptiles to migrate and interchange through the southern Burgundian Gate connection to the Tethys Ocean.

bone beds are both thicker and more condensed, form- zone in the Lainecker Höhenzug area (Fig. 3). ing “bone bed gravels” comprising the remains of animals adapted to brackish conditions (for example Blezingeria) and even amphibians (for example Mastodonsaurus, Pla- giosuchus, Gerrothorax), thus indicating the influence of fresh water [133] and the disappearance of marine condi- tions along the basin margins, starting above the nodosus

33 The Middle Triassic marine reptile biodiversity in the Germanic Basin,in the centre of the Pangaean world

Figure 18. In the intracontinental Germanic Basin the presence of extensive macroalgae meadows seem to have resulted in large numbers of Placodus gigas “Triassic seacows”. Meanwhile, Pistosaurus longaevus was the earliest of the ”subaquatic flying reptiles” and was adapted to open marine conditions, together with the ichthyosaurs. Nothosaurus giganteus was a shallow marine “Triassic marine alligator”, while the smaller Nothosaurus mirabilis can be seen as “Triassic marine gavial” (Illustration G. “Rinaldino” Teichmann).

5.3. Facies differences in the Bayreuther sec- Germany (for example [129]); these alternate rhythmically tions with terebratulid-dominated tempestites have sometimes become hardgrounds (cf. [31, 52]; Fig. 2C). The succeed- ing Meißner and Warburg Formations at the top of the Four main facies types can be distinguished in the Up- Upper Muschelkalk (Laineck road cut) consist of dolomitic per Muschelkalk of Lainecker Höhenzug, with different limestones, dolomites, dark-coloured clays, a bone bed, cycles of sedimentation (Figs. 2-3). The Bindlach For- and grey to brown marlstones, and are similar to northern mation is largely made up of glauconitic nodular marls Germany with ceratite absence above the nodosus biozone and terebratulid-dominated limestone beds which are of (for example [28]; Fig. 2D). tempestite origin (cf. [52]; Fig. 2A). In some parts of this section, and also over much of the Upper Muschel- While the chronostratigraphy follows Bachmann and kalk, platy limestones (= Tonplatten, cf. [52]) are repre- Kozur [54], and Kozur and Bachmann [125], the lithostrati- sented with some thicker claystone layers forming possi- graphical subdivisions of Hagdorn and Simon [126], and ble regional marker beds which correlate well in southern Bachmann et al., [127] were also used in part, because Germany such as the Spiriferina Bed, “Wellenkalk Bed”, there is an overlap in Bayreuth between the main facies or bed of small terebratulids (cf. [137]; Fig. 2B), but types of northern and south-western Germany (cf. Fig. 3), overlap with marker beds especially in the upper sec- resulting in variations in the general marginal facies. As tion which is more in northern German facies such as a result, two new formation names have been introduced the “Albertii-Bed” (for example [129]) or the best marker because of the absence of the trochitic crinoidal lime- beds at all – the partly facies breaking and basin dis- stones (Trochitenkalk Formation: [126, 127]; Fig. 3) and tributed bone beds. The overlying Hegnabrunn Forma- the near absence of the deeper shallow marine Tonplat- tion is characterized by nodular marlstones (Fig. 2C), ten facies (Meißner Formation; cf. [126, 127]). The Bind- which are also glauconitic and are absent in northern lach Formation consists of cyclic, autochthonous, glau-

34 Cajus G. Diedrich,

conitic marls and terebratulid-dominated tempestite beds south-western Bavarian Depression and to the west of (cf. [51]). The parasequences are mostly coarsening up the Rhenish Massif and Hessian Depression areas, where sequences - sensu Aigner [141] - within the general trans- they formed along the steep carbonate ramps as a result gressive trend of the middle Upper Muschelkalk. The au- of the basin morphology ([31, 52], Fig. 3). tochthonous marls are similar to the Hasmersheimer sub- With the facies development in the basal Upper Muschel- formation sediments, but they are stratigraphically and kalk (Diemel Formation) (Fig. 16, [28]) extensive oolitic palaeogeographically restricted, reaching only into the sand bar deposits accumulated in the central part of the atavus biozone, and are better developed in the south Germanic Basin. Placodont reptiles and some sauroptery- ([31]). These marls alternate with tempestites throughout gians migrated into the area with the new Muschelkalk the Bindlach Formation, extending from the atavus zone transgression, following the development of macroalgae into the compressus zone. The Hegnabrunn Formation, meadows (Fig. 16A). which follows the Bindlach Formation, is further dom- The shallow carbonate ramp intertidal flat zones subse- inated by tempestites, with intercalating nodular marl- quently disappeared and were replaced by steep ramps stones, and hence each of these sequences has been given whose facies types are reflected in the Trochitenkalk a separate formation name. Formation and Meißner Formation of the middle Upper The Coenothyris vulgaris terebratulids formed tempestite Muschelkalk at Bindlach and Hegnabrunn [28]. With layers that were only 5-25 cm thick in the Bayreuth lo- the development of deeper basin morphology and steeper calities, but at Bissendorf (northern Germany) these ter- coastal ramps the macroalgae zones migrated towards the ebratulids formed a 10 m thick massive terebratulid-shell coastlines and were replaced by crinoidal meadows (Fig. limestone bed (rudstone) in the Osnabrücker Formation 16B). The maximum flooding increased the reptile biodi- ([129], Fig.3). Coenothyris is almost completely absent versity of also ichthyosaurs which were adapted to fully from Lamerden, being present only in a single thin (10 marine conditions (Fig. 16B). The regression in the upper cm) tempestite layer in the atavus zone that also includes part of the Upper Muschelkalk (Warburg Formation/Erfurt double valved specimens of C. vulgaris (cf. Fig.3). Al- Formation) then produced drastic changes, with the dis- though at Lamerden the facies is mostly that of a steep appearance of macroalgae and crinoids. Only periodic basinal ramp, starting with a 10 m thickness of crinoidal interchanges through the Burgundian Gate allowed fur- limestones (= Trochitenkalk facies) and overlain by Ton- ther marine reptile migrations into the Bavarian/Hesse platten facies (Fig. 3), the Trochitenkalk facies is absent Depression (Fig. 6C). from the Lainecker Höhenzug outcrops and in Bissendorf This lagoonal area connected to the Northwest-Tethys is represented only by a 10 cm thick Trochitenkalk Bed in then became more hypersaline and restricted, and even- the pulcher zone (cf. Fig. 3). tually brackish (Fig. 16D) with bone beds containing a mixing with non-marine faunas (= Grenz bone beds, Crail- sheim, [142, 143]; Fig. 15 - Lamerden), followed by an in- 5.4. Facies related reptile occurrences and crease in marine/brackish reptiles (for example Kirchheim biogeography Anthrakonit Bed) and the dominance of pachypleurosaurs (cf. [23]), and finally, the presence of brackish/limnic am- The selected sections figured herein (Fig. 3) are quite phibians and thecodont archosaurs in the early Keuper well subdivided by their bio- and chronostratigraphy and (Kupferzell, Vellberg-Eschenau, Lower Grey Marls Bed: include the various bone beds [49, 129]i, show the main cf. [24, 25, 31, 140]). facies types, within a cross section of those facies types in the Upper Muschelkalk of the Germanic Basin. The lagoonal facies at the beginning of the Upper Muschel- kalk is mainly represented in the centre of the Germanic 5.5. Quantitative analyses of the Bayreuther Basin, while intertidal flats with mud-cracked biomats reptile fauna were present at the coasts [28]. The new Upper Muschel- kalk marine transgression oolitic bars (= oolite bar fa- At least 600 single bones have been newly identified, cies, Fig. 3) were distributed within the basin centre which form the basis for the preliminary statistics. One and on the western ramp. The steeper basin margins sur- problem appears to be the absence of smaller reptiles, es- rounding the Bohemian Massif developed into glauconitic pecially those such as the pachypleurosaurs, but the ma- marls and terebratulid tempestites (= marginal facies, Fig. terial from the larger reptiles nevertheless allows a com- 3). The crinoidal facies formed at about 7 metres water parison to be made with the previously examined bone depth, with Encrinus bioherms developing mainly in the bed assemblages from Bissendorf [49] and Lamerden [17].

35 The Middle Triassic marine reptile biodiversity in the Germanic Basin,in the centre of the Pangaean world

In this analyses of the Bayreuth reptile fauna only the Placodus within the reptile fauna of Monte San Gior- skull material was used for the minimum number of indi- gio, which remained unexplained in previous palaeobio- viduals (MNI) statistics, because the postcranial material geographical studies [42, 150], is therefore now clearly is not fully prepared and identified in all specimens. For explained. However, similar species of Paraplacodus and those species where there was no skull material available Cyamodus, which are also believed to have been macroal- (Neusticosaurus, Tanystrophaeus etc.) the number was gae feeders [43], are present in both regions, for example set on 1, which was added on the other species amounts Paraplacodus broili [151], and Cyamodus kuhnschnyderi. each. The results of this cranial-based analysis (Fig. These may have fed on different algae that were spread 15) support the quantitative analyses and reptile facies- along the Pangaean coastlines [56]. Another similarity relation (Fig. 3). These statistics, when compared to those between these two regions is the presence of pachypleu- from Lamerden and Bissendorf, support the compositional rosaurs such as Neusticosaurus pusillus [23, 152]. Lar- differences in the reptile fauna and the facies-related in- iosaurus balsami is also not endemic and has been iden- terpretations. Only in the macroalgae- and terebratulid- tified from Bad Sulza [107] as well as in the Bayreuth rich shallow marine facies of the atavus to evolutus bone localities to be a common smaller nothosaur; its remains beds are placodonts represented (predominantly by Pla- were previously incorrectly attributed to Lower Muschel- codus) at Bissendorf, whereas they are abundant (to- kalk species (such as “N. marchicus”- skulls, see [60]) gether with Cyamodus) within the reptile fauna at the and the postcranial material was completely overlooked. Bayreuth sites (Fig. 15C), and almost completely ab- The large Nothosaurus mirabilis, as well as Paranoth- sent from Monte San Giorgio [151] or Lamerden [133] la- osaurus giganteus, were also previously incorrectly iden- goons. They are well represented in the Bissendorf com- tified, but their skull identifications were subsequently re- pressus bone bed but fewer in number, although this is vised [60]. Simosaurus gaillardoti seemed to be restricted possibly a result of the limited amount of material avail- to the fossil record of the Germanic Basin, but more re- able (Fig. 15B). In Lamerden, the enodis/poseckeri bone cent discoveries at Siles, in Spain [101] prove that they bed assemblage is very different in its reptile composi- also existed in the north-western Tethys. While there tion and quantity (Fig. 15A), and the absence of any appear to be major differences in the the abundances of placodonts correlates with the absence of a macroalgae- ichthyosaurs in the Germanic Basin and Monte San Gior- palaeocommunity. Instead, amphibian remains result from gio (cf. [108, 110, 111, 113, 152]), the Shastasaurus and a limnic influence (Fig. 15A), although predominantly Omphalosaurus species are similar in both the Tethys and pachypleurosaur bones suggests a lagoonal paleoenviron- the Germanic Basin, and their abundance differences re- ment. Not only are the Muschelkalk invertebrate com- late to the open marine northwestern Tethys and shallow munities [31, 51, 74] important for facies analyses and intreacratonic Germanic Basin conditions. palaeoenvironmental reconstructions, but so to are the reptile or fish (cf. [144, 145]) faunas. Finally, the terrestrial reptile record is different between the Germanic Basin and Monte San Giorgio/Perledo, as is the track record attributed to these animals ([129]). At Monte San Giorgio the carcasses of several terres- 5.6. Global context of the Germanic Basin trial reptiles were washed into the lagoons, including those of Hescheleria ruebeli [153], Macrocnemus bas- reptiles and track records sani [154], Tanystrophaeus longibardicus [155], and Tici- nosuchus ferox [156]. These all appear to be the younger The endemism in north-western Tethys and Germanic records (Fassanian) of the older (Aegean-Illyrian) track Basin reptiles can now be seen to be far less common than records from the Germanic Basin of Procolophonichnium previously described (for example [10]). The main fau- (Hescheleria tracks), Rhynchosauroides (Macrocnemus nal differences between the Germanic Basin and Monte tracks), and Chirotherium, Isochirotherium (Ticinosuchus, San Giorgio marine reptile faunas are the presence or Arizonasaurus tracks) [66, 129]. The track record also absence of abundant Placodus gigas. Their abundance provides evidence that additional Middle Triasic reptiles in the Germanic Basin was because of the shallow ma- remain to be discovered at Monte San Giorgio because, rine macroalgae-rich habitats which were absent in the for example, the Isochirotherium tracks are most proba- Monte San Giorgio lagoons (cf. [145–147]). These “Tri- bly related to the Arizonasaurus skeletal remains found assic sea cows” [43] were not shell consumers (as has in North-America (Fig. 17, [64, 66]). The Germanic Basin commonly been claimed without any supporting evidence: provides a track and terrestrial fossil track record for the Westphal, [10, 148, 149], but fed on these macroalgae- Middle Triassic that includes behavioural details such as rich habitats of the Germanic Basin. This absence of the feeding of archosaurs on horseshoe crabs [64, 66], and

36 Cajus G. Diedrich,

palaeoenvironments that were unique during the Middle it would appear that these reptiles adapted to shallow ma- Triassic in the centre of the Pangaean World. The lagoons rine conditions (placodonts, nothosaurs) interchanged be- of Monte San Giorgio provide evidence of another aspect tween “Israel” and “China” along the north-western Tethys of the Middle Triassic biodiversity, together with ecolog- coastlines (Fig. 16). The material from Bayreuth is from ical information – and only a consideration of both areas the Illyrian to Fassanian and younger in age than the together can provide a complete picture of the Middle Tri- new skeletal material from China, which dates into the assic marine and coastal life, and in particular the reptiles Pelsonian [11]. The placodont occurrences correlate with which were eventually restricted to the warmer ocean re- the distribution of shallow marine carbonates. In contrast, gions of the globe [157]. This also supports the theory the open water marine pistosaurs must have interchanged of the “Triassic Sea Cows”, because the shallow marine between the east coast of Pangaea where their remains carbonates and macroalgae palaeocommunities that they have been found [113], and the west coast of Pangaea, fed on appear to have been restricted to the subtropics to which could only have happened more or less within the tropics marine regions (Fig. 16). tropics (Fig. 16). The fully-marine adapted ichthyosaurs, however, are found all over the world (Fig. 16, [113, 152]). 5.7. Horseshoe crab reproduction beaches Various reptile groups were already producing live off- and reptile food-chain reactions spring within marine environments, such as, for example, the pachypleurosaurs [160] and ichthyosaurs [110], and the sauropterygians were also possibly able to reproduce Within warmer regions horseshoe crab reproduction zones in this more evolved manner, which explains the absence have been identified as having been restricted to the ex- of marine reptile trackways on the surrounding intertidals tended intertidal and coastal zones of the Germanic Basin of the Germanic Basin coasts [64, 129], which would be during the Middle Triassic [62], and appear to have re- expected in beach reproduction zones, where the eggs of sulted in widespread food chain reactions among the ter- reptiles are likely to have been deposited. restrial archosaur populations, possibly even resulting in global migrations ([62, 66]). The large numbers of horse- shoe crabs in these reproduction zones (mainly docu- mented by the enormous amounts of tracks and track- ways: [62]) must also have attracted fish, and higher 6. Conclusions predators such as the shallow marine pachypleurosaurs and nothosaurs, explaining their presence in large num- bers within the Germanic Basin. The record of juvenile The famous Middle Triassic reptile outcrops near horseshoe crabs found in intertidal deposits of the upper- Bayreuth, in southern Germany, are in most cases still most Middle Muschelkalk at one of the Bayreuth local- accessible. Three main sections at Hegnabrunn, Bindlach ities [63, 158], Diemel Formation, cf. Fig. 3) supports and Laineck are composed of a 70 metre thick “complete the unique situation of the Germanic Basin in Pangaea Upper Muschelkalk” limestone series (ranging from be- with respect to its food chains. The horseshoe crab repro- low the pulcher to above the semipartitus biozones). This duction beaches are completely absent from the Monte can be subdivided into two new formations, the Bindlach San Giorgio lagoons, as are reptile tracks or intertidal Formation and Hegnabrunn Formation, which are in turn biolaminate sediments. In summary, the Monte San Gior- overlain by the existing Warburg Formation and Erfurt gio/Perledo lagoons and the Germanic Basin shallow ma- Formation. The section ranges from the top of the up- rine habitats are both of similar international importance permost Middle Muschelkalk (Diemel Formation, late Il- in understanding of invertebrate and vertebrate palaeo- lyrian) to the earliest Keuper (Erfurt Formation, middle communities of similar ages (Illyrian-Fassanian), in two Longobardian). Dating of these sediments is possible on different paleogeographical situations. the basis of ceratite finds, which range from the atavus to the nodosus biozones. The Upper Muschelkalk series contains several bone beds, which are named after the 5.8. Biodiversity, biogeography and repro- ceratites that they contain, which can be found mostly duction basin-wide as marker beds in several cases. Most of the historically collected bones are from the Bindlach sec- That the Monte San Giorgio/Perledo lagoons and the tions, which range from the atavus to the evolutus bio- Germanic Basin are very similar in their marine verte- zones (lower Upper Muschalkalk, late Illyrian to early brate faunas is not surprising, and following the discovery Fassanian). Sediment attached to un-prepared bones is of new placodont and nothosaur records in China [11, 159] similar to the main facies type of the Bindlach Formation,

37 The Middle Triassic marine reptile biodiversity in the Germanic Basin,in the centre of the Pangaean world

terebratulid-rich tempestite layers that intercalate with production on carbonate intertidals is best documented the autochthonous glauconitic marls. by very abundant horseshoe crab trackways; addition- These layers are the main bone horizons, from which have ally some reptile bones of Phygosaurus, Nothosaurus and come the famous marine reptile remains of Neusticosaurus shark fin spines were found washed into large Isochi- sp., Lariosaurus cf. balsamii, and the Nothosaurus rotherium and Chirotherium trackways. mirabilis Meyer, 1839 holotype skull and a skeleton, All of the special conditions of the Germanic Basin that the Paranothosaurus giganteus (Münster, 1834) holotype are absent from other parts of the Pangaean world, in- skull fragment, the Placodus gigas (Agassiz, 1833) holo- cluding its location northwest of the Tethys Ocean in type skull, the Cyamodus rostratus (Münster, 1839) holo- the warm climatic zone of the central Pangaean conti- type skull, the Cyamodus münsteri (Münster, 1830) holo- nent, the unique intertidal carbonate mud flat coastlines, type skull, the Tanystrophaeus conspicuus Meyer, 1847 and the shallow marine and macroalgae-rich palaeoenvi- holotype neck vertebra, the Pistosaurus longaevus Meyer, ronments, can best be compared to the analogous situa- 1839 holotype skull and a skeleton, and Shastasaurus sp. tion in the present-day Persian Gulf. Sea turtles (with jaw and or Omphalosaurus sp. dorsal vertebra originals. ethological convergences to placodonts), whales (etholog- Benthic invertebrate palaeocommunities at Bayreuth in ical convergences to pistosaurs), and in particular, large the Illyrian/Fassanian series are adapted to macroalgae populations (comprising several thousand individuals) of meadows and facies. Those algae seem to have been the seagrass-feeding dugongs (ethological convergences to main food source for Placodus gigas populations consist- Placodus) migrate into the Persian Gulf. ing of small young to large grown up individuals. The In some instances higher marine reptiles such as pis- reptiles occur mostly during the Bindlach Formation and tosaurs and ichthyosaurs may have only migrated sea- to a lesser extent in the Hegnabrunn Formation. The sonally, for reproduction purposes. At the highstand that “Triassic Sea Cows” formed a high percentage (45%) of occurred within the Bindlach and Hegnabrunn formations, the reptile assemblages in the Bayreuth region, where a very diverse invertebrate and vertebrate community lived these shallow marine divers were found abundantly only under normal saline conditions and warm water temper- in the intracratonic Germanic Basin. Large numbers of atures that prevailed during these higher-sea level con- the placodonts (Paraplacodus, Placodus, Cyamodus) are ditions. The placodonts disappeared from the Germanic useful for the identification of macroalgae habitats and Basin in response to the disappearance of macroalgae; the palaeoenvironments. In contrast, the ichthyosaurs, which disappearance of all placodonts is well documented within are abundant all over Pangaea, particularly in the north- the Bayreuth Upper Muschelkalk sections. This mas- west Tethys and the Monte San Giorgio/Perledo lagoons, sive environmental change in the Germanic Basin occurred are almost completely absent from the Bayreuth facies within the Warburg Formation (Longobardian), when the indicating a reptile group highly adapted to open marine Germanic Basin was uplifted and fluvial influences pro- conditions. Pistosaurs, which are common in the Bayreuth duced brackish lagoons that yielded bone beds completely facies are also open marine adapted “subaquatic flying devoid of placodont bones. The placodonts were mainly reptiles” and appear to have travelled over the globe, as replaced by large populations of small pachypleurosaurs, do modern whales; they possibly migrated into the spe- Blezingeria and much larger nothosaurs N. giganteus or cial shallow marine intracratonic Germanic Basin for re- S. gaillardoti, which appear to have adapted to Neusti- production purposes only, because more than 99% of the cosaurus predation. With the disappearance of the Ger- bone material recovered is from large grown up individu- manic Basin during the Erfurt Formation (Longobardian), als. amphibians (Mastodonsaurus, Gerrothorax, Plagiosuchus) Horseshoe crabs also used the Germanic Basin as a re- and archosaurs (Batrachotomus) became abundant in the production area during the Middle Triassic, which must brackish, shallow lagoon bone beds, often still mixed in have resulted in a wide-reaching food-chain reaction. Ev- some layers with the remains of pachypleurosaurs and gi- idence for this has already been provided in the form ant nothosaurs. of archosaur feeding traces on a horseshoe crab, but is Apart from their invertebrate palaeocommunities, the ver- also likely to have extended to marine reptiles such as tebrates and bone beds of the Middle Triassic are very pachypleurosaurs (Anarosaurus pumilio, Phygosaurus sp.) valuable for stratigraphic and palaeoenvironmental re- and small nothosaurs (Nothosaurus marchicus) that pos- constructions - a value that not only applies to the sibly fed seasonally the fish, which latter preyed upon Germanic Basin, but is also useful for global compar- the horseshoe crab eggs in the shallowest marine coastal isons. In this case, ichthyosaurs and pistosaurs were habitats, such as best demonstrated at the Bernburg ver- the only open marine reptile groups that could have tebrate and vertebrate track site. There horseshoe re- been distributed right across the Panthalassa Ocean

38 Cajus G. Diedrich,

(including the Tethys), possibly even migrating into ologic private research institution (www.paleologic.de). I marginal basins for reproduction purposes. Most other thank Dr. J. Szulc for the review of the former manuscripts. marine reptiles, such as the nothosaurs (Lariosaurus, Finally E. Manning provided support for language check- Nothosaurus, Simosaurus), for example, were shallow ma- ing, and with other critical remarks to the geology. rine coastal hunters. Pachypleurosaurs (Neusticosaurus) were adapted to lagoonal environments, and placodonts were restricted to macroalgae-rich shallow marine habi- tats, which were best developed in parts of the Germanic Basin, such as around Bayreuth, but also existed on tem- perate shelf margins along the Pangaean coasts. References Finally, along the beaches of Bayreuth and the en- [1] Curioni G.. Cenni sopra un nuovo saurio fossile die tire Germanic Basin coastline, terrestrial reptiles includ- monti di Perledo sul Lario e sul terreno che le rac- ing Tanystrophaeus appear to have used the intertidal chiude. Giornale del’ J.R. Instituto Lombardo di Sci- zones and beaches as habitats, leaving behind their foot- ence Lettre ed Arti NS, 1847,16, 159-170 prints, together with those of several other terrestrial rep- [2] Münster Graf zu G. Vorläufige Nachricht über einige tiles such as Macrocnemus (Rhynchosauroides tracks), neue Reptilien im Muschelkalke von Bayern. Neues Hescheleria (Procolophonichnium tracks), Ticinosuchus Jahrbuch für Mineralogie, Geognosie, Geologie und (Chirotherium tracks), and Arizonasaurus (Isochirotherium Petrefaktenkunde, 1834, 521-527. tracks), whose carcasses were washed into the coastal [3] Agassiz L. Recherches sur les Poissons fossiles. sediments as well as into the adjacent lagoons. Neuchâtel et Soleure, 1833-1843 [4] Münster Graf zu G. Beitrage zur Petrefaktenkunde, mit XVIII nach der Natur gezeichneten Tafeln der Herren Hermann v. Meyer und Professor Rudolph Acknowledgements Wagner. Buchner’sche Buchhandlung, Bayreuth, 1839 [5] Münster Graf zu G., Über einige ausgezeichnete fos- I am grateful to Dr. J.M. Rabold for facilitating the study sile Fischzähne aus dem Muschelkalk bei Bayreuth. of historical vertebrate and invertebrate collections from Birner, Bayreuth, 1830 the Bindlach, Laineck and Hegnabrunn quarries (mainly [6] von Meyer H., Mittheilung an Professor Bronn in the Meyer and Münster collections), in the Urwelt- gerichtet. Neues Jahrbuch für Mineralogie Geogosie Museum Oberfranken, Bayreuth, and for the integration und Geol Petrefaktenkunde, 1939, 559-560. of the fossil material from the 2010 fieldwork into the mu- [7] von Meyer H., Zur Fauna der Vorwelt. 2 Abt. Die seum’s collection. PD Dr. O. Rauhut kindly supported Saurier des Muschelkalkes mit Rücksicht auf die the study of material in the Bayrische Staatssammlung, Saurier aus Buntem Sandstein und Keuper. Frank- Munich (mainly Meyer material) and Dr. R. Brocke from furt a. Main, 1847-1855 the Senckenberg Natural History Museum, Frankfurt, pro- [8] Peyer B., Die Reptilien vom Monte San Giorgio. vided access to the historical bone collection from the Serie Zoologie, 1944, 78, 1-95 Bayreuth localities. The Triassic reptile collection from [9] McGowan C., A new species of Shastasaurus (Rep- the Lainecker Höhenzug of the Staatliche Museum für tilia: Ichthyosauria) from the Triassic of British Naturkunde in Stuttgart was accessible due to the kind Columbia: the most complete exemplar of the genus. support of Dr. R. Schoch. PD. Dr. O. Hampe supported Journal of Vertebrate Paleontology, 1994, 14, 168- the work on collections in the Natural History Museum of 179 the Humboldt-University, Berlin. Dr. M. Mäuser allowed [10] Rieppel O., Handbuch der Paläoherpetologie, Teil the study of the invertebrate and vertebrate material in the 12A. Sauropterygia I. Placodontia, Pachypleu- collections at the Naturkundemuseum, Bamberg. I would rosauria, Nothosauroidea, Pistosauroidea. Friedrich like to thank Dr. R. Riedel for supporting the work on Pfeil-Verlag, München, 2000 the collection in the Naturkundemuseum, Erfurt. Dr. S. [11] Motani R., Da-Yong Jiang D.Y., Tintori A., Sun Y.- Chapman kindly provided the Cyamodus skull photos of L., Hao W.C, Boyd A., Sanja H.F., Schmitz L., Shin the material in the Natural History Museum, London, (the J.Y., Sun, Z.Y., Horizons and Assemblages of Mid- original material from Owen). The sole sponsor for this re- dle Triassic Marine Reptiles from Panxian, Guizhou, search, as part of the “Middle Triassic Megatracksite and China. Journal of Vertebrate Paleontology, 2008, 28, Reptile Project of the Germanic Basin”, was the Palae- 900-903

39 The Middle Triassic marine reptile biodiversity in the Germanic Basin,in the centre of the Pangaean world

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