Open Geosci. 2015; 7:755–782

Research Article Open Access

Cajus G. Diedrich * The vertebrates from the Lower Ladinian (Middle Triassic) bonebed of Lamerden (Germany) as palaeoenvironment indicators in the Germanic Basin

DOI 10.1515/geo-2015-0062 sis; palaeoenvironment; palaeobiogeography; Germanic Basin; Central Europe Received January 28, 2013; accepted May 21, 2015

Abstract: A marine/limnic vertebrate fauna is described from the enodis/posseckeri Bonebed mixed in a bivalve shell-rich bioclastic carbonate rudstone at the eastern 1 Introduction coastal margin of the Rhenish Massif mainland at Lamer- den (Germany) within the western Germanic Basin (Cen- Middle Triassic vertebrate skeleton or bone localities and tral Europe). The condensation layer is of Fassanian (La- bonebed layers are known from all over the “Pangaean dinian, Middle Triassic) in age. The vertebrate biodiver- Globe”, especially Northern America, Europe, and China sity includes ve dierent shark, and several actinoptery- (Figure 1A) [e.g. [1–7]]. In north-western Germany, a higher gian sh species represented by teeth and scales. Abun- density of vertebrate sites (Figure 1B) is the result of dant isolated bones from a small- and a large-sized pachy- many active limestone quarries which expose bonebeds th pleurosaur Neusticosaurus species, which can be com- within dierent layers since the 19 century. The old- posed as incomplete skeletons, originate from dense pop- est described historical sites are in southern Germany ulations of dierent individual age stages. Important fa- near Bayreuth (Bavaria) and became famous with the cies indicator reptiles are from the thalattosaur Blezin- world-wide rst sauropterygian monograph published by geria ichthyospondyla which postcranial skeleton is re- Hermann von Meyer [8]. Those localities are known to- constructed hypothetically using additional postcranial day to contain several bonebed layers in the Illyrian- bones from similar aged various German localities. The Fassanian limestone series which were recently revised vertebrate biodiversity of the enodis/posseckeri bonebed in their stratigraphy, facies and vertebrate biodiversity for of Lamerden reect a limnic/uvial freshwater inuenced several still open old quarries [7]. After the rst historic fauna (amphibians/terrestrial and marine reptiles) with wave of Triassic reptile discoveries in southern Germany, dominance of normal saline marine inuences. Macro- the few younger “Muschelkalk/Keuper-Grenzbonebeds” algae meadow adapted placodont reptiles are absent in (= Boundary Bonebeds, Fassanian/Longobardian bound- Lamerden, as well as open marine-adapted ichthyosaurs, ary) became famous in more recent times. Those bonebeds supporting a lagoon with fresh water inuence position at contain single tooth and bone remains of a mixed ma- the Rhenish Massif mainland coast. In those contempo- rine/brackish/limnic vertebrate fauna which was never ex- ranous brackish lagoons, which seem to be isochronous cavated systematically (no square meter documentation) to northern Tethys lagoons of the Kalschieferzone at the at several southern German quarrys, such as in the most Monte San Giorgio (Switzerland/Italy), small pachypleu- famous locality Crailsheim [1–3]. Systematically excavated rosaurs were abundant prey in both regions for reptile bonebeds have been started to be analysed from dier- predators, especially large paraxial swimming alligator ent “Upper Muschelkalk” levels of northern Germany more habitus-like Paranothosaurus, which even contain stom- recently. At Bissendorf, the compressus Bonebed of the ach contents of pachypleurosaurs. Ilyrian-Fassanian boundary [9] is also important for the fa- cies/vertebrate fauna comparisons presented herein. The Keywords: Bonebed; vertebrate biodiversity; facies; Lon- pulcher/atavus Bonebed with its historical larger collec- gobardian (Middle Triassic); vertebrate taphocoeno- tions from Bad Sulza (central Germany) was also restudied

*Corresponding Author: Cajus G. Diedrich : PaleoLogic, Private Research Institute, Zdice, Czech Republic, website: www.paleologic.eu, E-mail: [email protected] © 2015 C.J. Diedrich, published by De Gruyter Open. This work is licensed under the Creative Commons Attribution-NonCommercial-NoDerivs 3.0 License. The article is published with open access at www.degruyter.com. Unauthenticated Download Date | 2/2/16 9:56 PM 756 Ë C.J. Diedrich

[10], and is compared to the new site presented herein, the tooth/scale over all excavation areas. One square metre enodis/posseckeri Bonebed from the central German site was mapped during each excavation campaign, which is Lamerden (Figure 1B). All the above mentioned bonebed quite well representative of the complete excavated area. localities are important for the understanding of verte- This bonebed does not show any clear scour trough, tidal brate assemblages in dierent facies, bathymetry and channel or ripple mark oriented concentrations or any salinity conditions which nally are useful for palaeoenvi- sorting. Statistical tests (= NISP) at two other squares (cf. ronmental reconstructions (palaeobiogeographical maps) Figure 3A), all chosen far from each others and during each and understanding of the palaeobiology of marine reptile excavation campaign, gave similar vertebrate (especially palaeopopulations of the Germanic Basin in Central Eu- sh remain) amounts. rope [cf. [7, 9]]. All bonebeds of the Germanic Basin are de- veloped in several dierent biostratigraphic (mostly at se- 2.2 Taphonomy of reptiles and amphibians quence boundaries, ceratite biozone boundaries) and geo- graphical (coastal reconstructions) positions and dierent The reptiles were mapped on surfaces of 4 and 160 square facies types [7, 9, 10], to which Lamerden contribute to a metres each (Figures 3-4). The problem of synsedimentary third type, besides the “shallow full marine/normal saline erosion by tempestitic channels overprinted the amount of bonebeds” of Bad Sulza, Bissendorf and Bayreuth. Finally nds in some areas (Figure 3A), which were not chosen for a discussion about Blezingeria can be added with very rare the sh/small reptile remain statistics. The large surface bone material from Lamerden and other sites of Germany. mapping (Figure 4) was similar chosen for an area, where the overlaying tempestite limestone bed did not reach and eroded into the bonebed. In some areas, the marls be- 2 Material and methods neath the bonebed were bioturbated by crustacean ich- nites of the type Thalassinoides (Figure 3A), which also in- 2.1 Excavations in Lamerden uenced the bonebed layers in some areas (cf. Figure 3). In total, 2,036 vertebrate remains, including teeth, scales and The lithostratigraphy, carbonate sedimentology and bones, were excavated, prepared and inventoried (Table 1). macro-invertebrate assemblage were studied for the A statistics about the bone element presence/absence Lamerden section to separate small-scale sequences for analysing a possible current or general taphonomic and to understand the palaeoenvironment development sorting is problematic, because the partly weathered change within the high order transgressive-regressive se- bonebed did not allow a sieving to obtain the small cranial quence (Figure 2). Furthermore, several ceratites have and appendicular bones or teeth, especially of the small been collected from the outcrop walls, which allow pre- pachypleurosaurs. Therefore, all amphibian and reptile senting an updated high resolution biostratigraphy (Fig- material (teeth, bones) larger then 3 mm was selective used ure 2). for the bone taphonomy analyses which allow subdividing Three bonebed excavation campaigns, each about four stages of bone fragmentation (Figure 5D): 1. None - four-weeks long, were performed in the active quarry bones show no breakage nor edge polishing, 2. Low - bones Lamerden (Kalksteinwerk Lamerden of the Fischer GMBH are rarely broken (mostly ribs and at bone elements) and and CoKG) in 2003, 2005 and 2006. The bonebed was have sometimes rounded corners, 3. Medium - fragmenta- excavated systematically on large surfaces, in total of tion and rounded corners is more common, 4. High - the about 600 square metres, representing the largest Ger- bones are all incomplete, fragmented, and after this even man bonebed excavation in Upper Muschelkalk bonebeds. rounded at the margins, joints are missing, and mainly In 2003, four square metres (Figure 3), and in 2005 (160 most stable and thick bone parts remained. square metres, Figure 4), were mapped for the vertebrate The “composite experiment” of most abundant pachy- taphonomy and vertebrate biodiversity analyses (NISP = pleurosaur reptile bones (Figure 5D) reects the pres- bone per species). Any surface mappings sadly did not re- ence/absence of the entire body bone elements (except the sult in palaeocurrent information (also due to short bone smallest ones as result of excavation technique or preser- elements only – no complete ribs e.g.). A further detailed vation) and dominance of the larger bone material. This mapping of nds was not performed again in 2006, also to is similar to large Paranothosaurus, and Blezingeria re- rescue as much as possible larger bone material for the ver- mains, but dierent from the more fragmentary material of tebrate biodiversity analyses. The large amount of sh re- Nothosaurus, Simosaurus and especially Pistosaurus (Fig- mains, especially selachian teeth and less of actinoptery- ure 5D), which latter had longer transport durations or dis- gian teeth and scales, did not allow the mapping of every tances.

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Figure 1: A. Globally important Midle Triassic marine reptile sites. B. The vertebrate locality Lamerden in North Hesse, Germany, an active limestone quarry in the Middle Triassic (Anisian/Ladinian) and other known important “Upper Muschelkalk” vertebrate bonebed sites [after [7, 9, 10, 64]]. The excavation surfaces were along the active quarry wall.

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Figure 2: Stratigraphy, environment and position of the enodis/posseckeri bone bed in the quarry of Bissendorf, NW-Germany (Middle Trias- sic). New ceratite nds allow a new precise dating of the section and the bone bed [Chronostratigraphy after [11]].

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2.3 Studied material and collections most complete marine “Upper Muschelkalk” sections in Germany, representing a full higher order trans- and re- The bone and invertebrate material from the excava- gressive cycle in the intracratonic Germanic Basin of cen- tions and collected cephalopods (ceratites) from the tral Europe (Figure 1A). stratigraphic work were prepared by the company Pa- The top of the Middle Muschelkalk (Heilbronn Forma- leoLogic and are deposited now in the Naturkunde- tion) consists of grey marls, dolomites and biolaminates. museum Ottoneum Kassel = NMOK). Compared mate- In Lamerden the footprints Rhynchosauroides and Procolo- rial is housed in the Senckenbergmuseum Frankfurt (= phonichnium were found in the biolaminates (= track bed SMF) including the only known Placodus gigas skeleton 21: [12]). Those deposits are overlain by several metre thick, (no. R1035), a Pistosaurus longaevus skeleton (no. 4041), yellowish dolomites of the Diemel Formation (= “Gelbe Ba- Neusticosaurus pusillus skeletons from Monte San Giorgio sisschichten”). (Kalkschieferzone), a Lariosaurus juvenile skeleton from Those dolomites are overlain by the approximately Perledo (Meride Limestones), as well as Cyamodus ma- 20 metre thick massive Trochitenkalk Formation. As a terial and many isolated bones of dierent marine rep- result of full marine conditions, these crinoid and less tiles. In the Urwelt-Museum Oberfranken Bayreuth (= UM- bivalve shell-dominated bioclastic rudstones/oatstones O), the Nothosaurus mirabilis skeleton (uncataloged), and can reach about 10 metres in thickness (= “Lower Tro- several skulls of Paranothosaurus, Nothosaurus, Placodus chitenkalk”, Figure 2). The Trochitenkalk limestone series with isolated material of dierent species were studied. In contains in the region a high amount (about 85%) of iso- the Natural History Museum Staatliche Naturhistorische lated remains of the crinoid Encrinus liliiformis. 90 cm Sammlungen Stuttgart (= SNSD), an undescribed Paran- above the “Upper Trochitenkalk”, a tsunamite layer with othosaurus giganteus skeleton (uncataloged in the exhi- complete crinoids was discovered [13] excavated and ana- bition) and many skulls and isolated bones of dierent lyzed [14]. About 40 cm below this allochthonous crinoid species including Tanystrophaeus material, mainly from layer, the autochthonous bioherm bed was found with Bayreuther sites, and from Monte San Giorgio, are de- about one metre wide and 40 cm high bioherms on large posited. The Bayrische Staatssammlung University Mu- scaled ripple marks on the top of the Upper Trochitenkalk nich (= BSP) has the Placodus holotype (no. BSP AS VII Bed. Those bioherms are in the pulcher Biozone, whereas 1208) and N. mirabilis skulls and a Lariosaurus balsamii the crinoid layer is already in the compressus Biozone [14] skeleton (uncataloged cast). In the University Tübingen (= (Figure 2). UT) the Simosaurus gaillardoti skeleton (uncataloged) was The following Meißner Formation is built of the “Ton- compared. The Natural History Museum of the Humboldt- platten” facies. It consists of changing marls and tem- University Berlin (= MB) collection contain isolated bone pestitic limestone layers, which contain many marine material, such as the Muschelkalkmuseum Ingelngen (= macrofossils and abundant cephalopods (ceratites). In MHI), and the Natural History Museum Erfurt (= ME). the penndor Biozone a larger ceratite accumulation was Compared Monte San Giorgio skeletons Neusticosaurus excavated [15]. Only the zones of the pulcher and en- pusillus and Neusticosaurus edwardsii or Serpianosaurus odis/posseckeri can be separated with their exact under- mirigiolensis are housed in the Paläontologische Museum and overlying boundaries. Two bone layers with few verte- der Universität Zürich (= PIUMZ). Other compared skele- brate remains are present in the Meißner Formation with tons of Anarosaurus pumilio or Nothosaurus marchicus the older compressus and evolutus Bonebeds (Figure 2). were studied in the Geologische Zentrum Göttingen (= The third and youngest bonebed is dated into the en- GZG). In the Naturkundemuseum Magdeburg (= NMM) the odis/posseckeri Zone (Upper Fassanian, Lower Ladinian; Serpianosaurus germanicus holotype skeleton was nally Figures 2-4). This bonebed is above the Großenberg Bed compared. and Schellroda Bed, both of which are shallow subtidal massive bioclastic limestone beds (Figure 2). The three mil- limetre thin bonebed layer is built dominantly of com- pressed bivalves (Bekevillia and Hoernesia epifaunistic 3 Geology, stratigraphy and facies types = partly “Tonplatten softground community” [16], of which only the periostracum remained, whereas all arag- The 70 metre thick carbonate series of Lamerden reach onite and calcite shells were dissolved during diagene- from the Middle Muschelkalk up to the Upper Muschel- sis. Remarkable are several cephalopod jaw nds of Rhyn- kalk/Keuper boundary (Erfurt Formation; see Formation cholithes hirundo being preserved three dimensionally. subdivision and aging according to [11]). It is one of the Those jaws seem to belong mainly to Germanonautilus,

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Figure 3: A. Bone mapping of four square metres for sh and small reptile remain statistics (NISP = bone per species) in the en- odis/posseckeri bone bed (Lower Ladinian) of Lamerden. B. Selachian, and actinopterygian remains from the enodis/posseckeri bone bed (Lower Ladinian) of Lamerden, Germany. Thalassinoides burrows indicate shallow subtidal conditions such as channels, which reworked the bonebed layers in larger areas. 1. Hybodus longiconus Agassiz 1843 symphyseal tooth (No. LAM-77), labial. 2. H. longiconus, anterior tooth (No. LAM-2), labial 3. H. longiconus, anterior lateral tooth (No. LAM-80), labial. 4. H. longiconus, middle lateral tooth (No. LAM-83), labial. 5. H. longiconus, posterior lateral tooth (No. LAM-74), labial. 6. ?Hybodus sp., n spine fragment (No. LAM-349), lateral. 7. Polyacro- dus polycyphus (Agassiz, 1837), anterior tooth crown (No. LAM-103), labial. 8. P. polycyphus, strongly used anterior tooth (No. LAM-102), labial. 9. P. polycyphus, lateral tooth crown (No. LAM-105), a. occlusal, b. labial. 10. P. polycyphus, strongly used lateral tooth (No. LAM- 104), labial. 11. Acrodus gaillardoti Agassiz, 1837, incomplete anterior tooth crown (No. LAM-99), occlusal. 12. A. gaillardoti, lateral tooth crown (No. LAM-101), occlusal. 13. Palaeobates angustissimus (Agassiz, 1838), anteriot tooth (No. LAM-111), occlusal. 14. P. angustissimus, lateral tooth (No. LAM-108), occlusal. 15. Acrodus lateralis Agassiz, 1837, anterior tooth crown (No. LAM-153), occlusal. 16. A. lateralis, lat- eral tooth crown (No. LAM-154), occlusal. 17. A. lateralis, posterior lateral tooth crown (No. LAM-152), occlusal. 18. Colobodus sp., scale (No. LAM-369). 19. Colobodus sp., scale (No. LAM-373). 20. Colobodus sp., scale (No. LAM-370). 21. Crenilepis sp., scale (No. LAM-385). 22. Gyrolepis sp., scale (No. LAM-381). 23. Actinopterygii indet., scale (No. LAM-394). 24. Dollopterus sp., scale (No. LAM-391). 25. Colobodus maximus Quenstedt 1835, tooth (No. LAM-357). 26. Colobodus frequens Dames, 1888, tooth (No. LAM-353). 27. C. frequens, splenial/vomer fegment with teeth (No. LAM-366). 28. C. maximus, vomer with teeth (No. LAM-368). 29. Saurichthys/Birgeria sp., tooth (No. LAM-276). 30. Saurichthys/Birgeria sp., tooth (No. LAM-286). 31. Jaw of an unidentied shark (LAM-1509).

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Figure 4: Bone mapping of 160 square metres of the excavation in 2005 for large reptile and sh remain statistics in the enodis/posseckeri bone bed (Lower Ladinian) of Lamerden. Elongated objects show mainly an N-S and NNW-SSO orientation. Important are the rare nds with single bones of amphibians Mastodonsaurus or Gerrothorax/Plagiosuchus and the terrestrial Tanystrophaeus.

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Figure 5: Marine sauropterygian pachypleurosaur reptile remains from the enodis/posseckeri bone bed of Lamerden, Germany. A. Anarosaurus sp. remains: 1. Dorsal vertebra centrum (LAM-590), a. cranial, b. dorsal, c. lateral. B. Neusticosaurus cf. pusillus (Seeley, 1882) remains: 2. Incomplete coracoid (LAM-564), ventral. 3. Ilium (LAM-5), ventral. 4. Femur (LAM-566), caudal. 5. Dorsal vertebra neural arch (LAM-1653), a. lateral left, b. caudal, c. dorsal. 6. Dorsal vertebra centrum (LAM-1001), a. lateral, b. cranial, c. dorsal. C. Neusticosaurus cf. edwardsii (Cornalia, 1854) remains: 7. Tooth (LAM-630), lateral. 8. Humerus shaft (LAM-1660). 9. Humerus shaft (LAM-1683), dorsal. 10. Il- ium (LAM-562), lateral. 11. Ischium (LAM-563), ventral. 12. Fibula (LAM-639), dorsal. 13. Cervical vertebra neural arch (LAM-1494), a. dorsal, b. lateral right. 14. Dorsal vertebra neural arch (LAM-1477), a. dorsal, b. lateral left. 15. Dorsal vertebra centrum (LAM-1499), a. cranial, b. dorsal, c. ventral. 16. Thoracic rib (LAM-575), lateral (Neusticosaurus pusillus (Seeley, 1882) and Neusticosaurus edwardsii (Cornalia, 1854) reconstructed after [4, 35, 42], and originals in the SNSD and SMF). D. Composed Neusticosaurus cf. pusillus and Neusticosaurus cf. edward- sii skeletons from the Lower Ladinian enodis/posseckeri bonebed of Lamerden, Germany. Besides the vertebral column (centra), the dorsal and caudal neural arches and some ribs. In red marked bones from adult and juvenile Neusticosaurus individual remains which were found in Lamerden.

Unauthenticated Download Date | 2/2/16 9:56 PM Middle Triassic vertebrate assamblage in Europe Ë 763 1839-1842, 1924-1932, 1963, 2012-2019 LAM-89-101, 889-911, 1445-1446, 1474, 1843-1845, 1936-1939 LAM-151-272, 874-888, 939-941, 987-995, 1846-1866, 1940-1950, 2020-2036 LAM-106-150, 841-864, 1448, 1933-1935 LAM-102-105, 865-873 LAM-353-356, 358-367, 930-938, 1973-1977 LAM-1540, 1562 LAM-1503, 1509 LAM-418-549, 550-561, 563-566, 568-574, 576-589, 598, 998-1246, 1475, 1661-1663, 1707-1708, 1714-1725, 1747-1749 bonebed of Lamerden (Germany), all coll. NMOK. 227 teeth from all jaw positions LAM-1-88, 349, 410-417, 733-840, 1755, 1757, 1761, 1764-1765, Two cranial or pectoral girdle bone fragments of unclear anatomic po- sition 54 teeth from dierent anteriorposterior to positions andold dierent individuals 303 teeth from dierent jawtions posi- 71 teeth from all jaw positions and one n spine 12 teeth from anterior to lateral po- sitions 20 jaw fragments and mainly teeth LAM-350-352, 357, 368,One 924-929, jaw 1463-1473, fragment 1972 with three teeth and isolated teeth and scales 140 teethOne large tooth fragment and der- mal skull roong element fragment 473 bones: one premaxillary frag- LAM-273-345,ment, 912-923, 1449-1462, one 1867-1873, humerus, 2003-2011 twofragments, coracoid two ileum bones,femora, two 403 vertebranearly centra, complete and 14 about 26 half neural arches, 23ments ribs and frag- ) Quen- Gigan- Dames pusillus Agassiz Agassiz Agassiz Birgeria enodis/posseckeri a. cf. sp. one thoracic vertebra centrum LAM-590 sp. or sp. 14 small scales LAM-386-393, 398-405 sp. 9 single scales LAM-375-384 Plagiosuchus (Jaeger 1828) SpeciesHybodus longiconus 1843 Material Coll.-no. Acrodus gaillardoti 1837 Acrodus lateralis 1837 Palaeobates angustissimus (Agassiz 1838) Polyacrodus(Agassiz 1837) polycyphus Gyrolepis Colobodus maximus 1888 Dollopterus Saurichthys sp. Plagiosaurinae indet. (?Ger- rothorax/ Anarosaurus Neusticosaurus (Seeley 1882) stedt 1835 Colobodus frequens Mastodonsaurus teus Vertebrate material from the Table 1:

Unauthenticated Download Date | 2/2/16 9:56 PM 764 Ë C.J. Diedrich LAM-626, 629-631, 633-634,1483, 1488, 1497, 1498, 696-721, 1495, 1498, 1500, 1502, 1477-1479, 1505,1518, 1512-1514, 1522, 1527-1529, 1533-1534, 1480-1481, 1539, 1541-1553, , 1555-1561, 1563-1571, 1573-1577-1582, 1650-1652, 1653, 1684-1685,1952-1954, 1706, 1955, 1957-1962, 1965-1970 LAM-636-637, 641, 1530, 1635, 2124, 2116 LAM-627, 632, 635, 643, 830,1512, 839, 1513, 1478, 1520, 1480, 1534, 1482, 1571, 1486, 1637, 1646, 1501, 1668, 2114, 2116 LAM-562, 630, 639, 1660 coll.1761-1763, 1821-1828, No. 1874-1875, LAM-575, 1913-1918, 1971 590, 637, 628, 1477, One lower jawment symphyses of frag- asome juvenile questionable animal,bones, and such as postcranial one enrolleda ulna, metapodial, one ileum, anrior ante- dorsal rib,and central cervical vertebra centrum gastralia, 9 Lower jaw fragments,one 64 incomplete teeth, humerus, ?one in- complete femur, several vertebra centra, partly half cervical todal cau- neural arches, 17caudal costae cervical and nally to 3 gastral ribs Some anterior andteeth, a large nasal bone lateral ofanimal a large, skull, old one lowertary jaw fragment, den- a?some carpalia/tarsalia, humerus one meta- shaft, pod, vertebrate centra and arches, ?dorsal, lumbal, and caudaland ?gastralia ribs, 43 bonesfragments, one including fanghumerus tooth, cranium shaft, one mur, incomplete one bula, fe- and ileum fragment completecentra, bone, neural arches, vertebrae questionable and ribs some Mün- edward- Münster cf. Paranothosaurus (Cornalia 1854) Nothosaurus ster 1834 Nothosaurus mirabilis Paranothosaurus giganteus (Münster 1834) 1834/ Peyer 1939 sii SpeciesNeusticosaurus Material" Coll.-no. Cont. Table 1:

Unauthenticated Download Date | 2/2/16 9:56 PM Middle Triassic vertebrate assamblage in Europe Ë 765 LAM-626, 638, 1476, 1496 LAM-613, 829 LAM-613, 722-730, 1372-1444,1735, 1829, 1447, 1832-1833, 1951 1535, 1537-1538, 1705, LAM-1508a, 1508b One questionable cervicalbra, verte- one dorsal neural arch, anterclavicle fragment, in- one question- able half ulna, one tibia,gatralia and one One femur shaft, one dorsal neural spine 63 vertebrae centra from Lamerden can be attributedreptile, to 16 this neural marine archare fragments insecure in its determinations, but one is for sure, also onespine dorsal fragment Two sacral vertebra fragmentsform of in lateral processi gaillardoti cf. SpeciesSimosaurus Material Coll.-no. Meyer 1842 PistosaurusMeyer 1839 Blezingeria longaevus ichthyospondyla (Fraas 1896) Tanystropheus conspicuus Meyer 1855 Cont. Table 1:

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Table 2: Abundance, bone presence and fragmentation degree, and primary facies of the amphibians and reptiles from Lamerden, Germany (NISP = number of bone per species).

Species NISP Bone elements Degree of fragmen- Original facies tation Mastodonsaurus giganteus 2 Massive cranial High, strong en- Terrestrial, freshwater bone plate frag- rolled rivers, coastal swamp lakes ments Gerrothorax/Plagiosuchus 2 Massive cranial High, strong en- Terrestrial, freshwater sp. bone plate frag- rolled rivers, coastal swamp lakes ments Tanystrophaeus conspicuus 2 Vertebrae fragments High, strong en- Terrestrial, coastal rolled Blezingeria ichthyospondy- 80 Vertebrae, mainly No to low enrolled Brackish fresh water influ- lus enced estuaries/lagoons Anarosaurus sp. 1 Vertebrae, mainly No to low enrolled Lagoons Neusticosaurus cf. pusillus 475 Vertebrae centra No to low enrolled Lagoons dominantly, all body part elements Neusticosaurus cf. edward- 42 Vertebrae centra No to low enrolled Lagoons sii dominantly, all body part elements Nothosaurus giganteus 21 Vertebrae centra No to low enrolled Shallow marine, lagoons dominantly, all body part elements Simosaurus gaillardoti 4 All body part ele- Medium enrolled Shallow marine, lagoons ments, mainly large bones Nothosaurus mirabilis 7 All body part ele- Medium enrolled Shallow marine ments, mainly large bones Pistosaurus longaevus 2 Only large bones High, strong en- Shallow marine rolled

Table 3: Revised comparison of reptile species in the Germanic Basin (Germany) and Monte San Giorgio or Perledo Alpine sites (Switzer- land/Italy) concerning the time frame of the upper Illyrian to lower Longobardian (based after [3, 4, 7, 9, 10, 38, 41, 42, 46–52, 55, 56, 58, 60, 65, 66, 69, 76]).

Species Germanic Basin shallow Northwestern Tethys Habitat marine intracratonic Basin, intraplatform la- basin lagoons goons High marine adapted Open marine ichthyosaurs Shastasaurus/Besanosaurus sp. Extremely rare Common Open marine Cymbospondylus sp. Extremely rare Common Open marine Omphalosaurus sp. Extremely rare Common Open/shallow ma- rine Mixosaurus cornalianus Extremely rare Very common Open/shallow ma- rine

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Table 3: Cont.

Species Germanic Basin shallow Northwestern Tethys Habitat marine intracratonic Basin, intraplatform la- basin lagoons goons Phalarodon sp. Absent (yet) Rare Open marine Mikadocephalus sp. Absent (yet) Rare Open marine Wimanius sp. Absent (yet) Rare Open marine

Shallow marine nothosaurs Simosaurus gaillardoti Very common Absent (yet) "Shallow marine, la- goons" Nothosaurus giganteus Very common Rare Shallow marine, la- goons Nothosaurus mirabilis Very common Rare Shallow marine Lariosaurus balsami Common Rare Shallow marine

Lagoonary pachypleurosaurs Anarosaurus sp. Rare Rare Very shallow marine lagoons Neusticosaurus pusillus Very common Very common Very shallow marine lagoons Neusticosaurus edwardsii Common Common Very shallow marine lagoons Serpianosaurus mirigiolensis Rare Rare Very shallow marine lagoons

Shallow marine placodonts Paraplacodus broilii Rare Rare Shallow marine Placodus gigas Very common Extremly rare Shallow marine, macroalgae mead- ows Cyamodus kuhnschnyderi Common Rare Shallow marine, macroalgae mead- ows Helveticosaurus zollingeri Absent (yet) Extremly rare Shallow marine Eusaurosphargis sp. Absent (yet) Extremly rare Shallow marine

Lagoonary thalattosaurs Clarizia schinzi Absent (yet) Rare Nearshore shallow lagoons Askeptosaurus italicus Absent (yet) Rare Nearshore shallow lagoons Blezingeria ichthyospondylus Very common Absent (yet) Nearshore shallow lagoons Terrestrial coastal reptiles Batrachotomus kupferzellensis Common Absent (yet) Coastal sand beaches and swamps

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Table 3: Cont.

Species Germanic Basin shallow Northwestern Tethys Habitat marine intracratonic Basin, intraplatform la- basin lagoons goons Ticinosuchus ferox Common (by Isochi- "Rare (by one skele- Coastal carbonate inter- rotherium footprints only) ton only)" tidals and sand beaches

Macrocnemus bassanii Common (by Rhyn- "Rare (by few skele- Coastal carbonate inter- chosauroides footprints tons)" tidals and sand beaches only) Hescheleria ruebeli Common (by Procolo- "Rare (by one skele- Coastal carbonate inter- phonichnium footprints ton only)" tidals and sand beaches only) Tanystrophaeus Rare (and by Synaptichnium Rare (by few skele- Coastal carbonate inter- conspicuus/longibardicus footprints) tons) tidals and sand beaches

Freshwater amphibians Mastodonsaurus giganteus Very common Absent (yet) Freshwater rivers, coastal swamp lakes Gerrothorax pustuloglomeratus Common Absent (yet) Freshwater rivers, coastal swamp lakes Plagiosuchus pustuliferus Common Absent (yet) Freshwater rivers, coastal swamp lakes which was found with several badly-preserved steinkern- val of the nodosus Biozone. At the same time, more north specimens in the bonebed layer (Figure 4) or below and and west (Lüdge, Vahlbruch, Eilversen, Figure 1B) ceratites above. Overlaying the bonebed, a ve centimetre thick reach up further in time into the marine deposits of the dor- marl can be absent. This is found at places where the marl soplanus Biozone. layer was eroded and replaced by a bioclastic tempestite Finally, the top of the Lamerden section represents the that also contains ceratites. In some areas, teeth are at- Warburg Formation and possibly parts of the Erfurt Forma- tached on the basal limestone bed. The channels often tion, but the exact stratigraphic subdivision above the sub- were additionally tectonically pressed onto the bonebed. laevigatus Biozone is not possible to present due to lack of The underlying limestones built in several areas smaller ceratites. diaper-like structures (Figure 4) penetrating the underly- ing marls and the bonebed from below. The ve cm thick marlstone above the bonebed (if present) contains an au- tochthonous fauna with infaunal bivalves of Myophoria vulgaris and epifaunistic Bakevellia and Hoernesia and the 4 Palaeontology ceratites C. enodis/posseckeri, which are sometimes en- riched in narrow-short-elongated subtidal channels [17]. 4.1 The vertebrate biodiversity of Lamerden The top of the Meißner Formation is dominated by claystones. It represents the transition to the terrestrially Sharks are represented mainly with teeth and few n inuenced following Warburg Formation. A test excava- spines by the common species (Table 1), such as Hybo- tion shows, that the nal bonebed (= Albertii Bed) con- dus longiconus Agassiz, 1843, Acrodus gaillardoti Agas- tains only sh scales (and rare shark teeth) in Lamerden. siz, 1837, Acrodus lateralis Agassiz, 1837, Palaeobates an- The last ceratites in Lamerden are not praenodosus forms, gustissimus (Agassiz, 1838), and Polyacrodus polycyphus which support a much earlier uvial inuence and re- (Agassiz, 1837) (Figure 3). Actinopterygian sh teeth and gression (caused by ?uplift Rhenish Massif) in this region. scales are from Gyrolepis sp., Colobodus maximus Quen- This was already brackish/terrestrial during the time inter- stedt, 1835, Colobodus frequens Dames, 1888, Dollopterus

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Figure 6: A. Nothosaurus mirabilis Meyer, 1834 remains from the enodis/posseckeri bone bed of Lamerden, Germany. 1. Lower jaw symphy- ses of a small juvenile individual (LAM-2124; cf. Figure 12.3). 2. Ulna (LAM-641), dorsal. 3. Metapodial (LAM-1635), dorsal. 4. Ilium (LAM- 637), lateral. 5. Anterior dorsal rib (LAM-1530), lateral. 6. Central gastralia (LAM-636), ?dorsal/ventral. 7. Cervical vertebra centrum (LAM- 2116), a. cranial, b. lateral.

Unauthenticated Download Date | 2/2/16 9:56 PM 770 Ë C.J. Diedrich sp., Saurichthys sp. or Birgeria sp. (Figure 3). The sh and been transported too far or too long, and were also not shark species are well known and described from several sorted by grain sizes. Vertebrae centra and neural arches sites all over Europe [3, 7, 9, 18–33]. are the dominant, and anyway most robust bone types The sauropterygian pachypleurosaur marine reptile represented, including ribs and teeth or extremity long- Anarosaurus sp. (? A. heterodontus) (Figure 5A.1a-c) is rare bones. The bone taphonomy (especially Paranothosaurus, (few well to identify vertebra centra). Most common is the Nothosaurus, Simosaurus, and Blezingeria: Figures 6-9) small pachypleurosaur Neusticosaurus cf. pusillus (See- is similar to the pachypleurosaur bones (Figure 5): long ley, 1882) (Figures 5B.2-6), followed by the double-sized bones and appendicle, pelvic and pectoral bones are rare Neusticosaurus cf. edwardsii (Cornalia, 1854) (Figure 5C.7- supporting a mainly sub-autochthonous character. 16). The large nothosaurs are Nothosaurus mirabilis Mün- The majority of reptile remains are vertebrae centra ster, 1834 (Figure 6A), Paranothosaurus giganteus (Mün- of the pachypleurosaurs, about two to four nds each ster, 1834) (Figure 17A), Simosaurus cf. gaillardoti Meyer, square metre (Figure 3A, 5). The main bone remains are 1842 (Figures 8A.1-6). Pistosaurs are represented by Pis- the ve to 15 mm small and most robust vertebrae centra tosaurus longaevus Meyer, 1839 (Figure 8B.1-2). Thalat- of pachypleurosaurs. The high amount of vertebrae cen- tosuchian remains are from Blezingeria ichthyospondyla tra per square metre (Figure 3) stands in anatomically cor- (Fraas, 1896) (Figures 9A.6-7). All those marine reptiles are rect contrast to the few appendicular bones and cranial described in most cases now by skeletons and more and remains, and reects non-sorting, because axial skeleton more with isolated bones in monograph catalogues and bones dominate naturally (Figure 5D). The problem of the were found again all over Europe [4, 7, 9, 34–55]. “missing” small pedal bones or cranial remains is also The fresh water amphibians Mastodonsaurus a. gi- the result of the excavation technique and the problem of ganteus (Jaeger, 1828), and Plagiosaurinae indet. (?Ger- the above mentioned bone preservation (bonebed weath- rothorax/Plagiosuchus) (Figure 10A-B) were described with ering), which does not allow a successful sieving to ob- skulls and skeleton parts from several German Lower tain the very small bone material. Here skeletons of the Keuper aged (Longobardian) sites, and mainly from the isolated larger bones of two Neusticosaurus species were southern Germanic Basin [36, 56–62]. The terrestrial pro- composed (Figures 5A-C). This composing experiment was torosaur reptile Tanystropheus conspicuus Meyer, 1855 important to understand a non-selection/sorting of dis- (Figures 10C.1-2), which is described to be limited on the articulated pachypleurosaur skeletons; these bones are Upper Muschelkalk Germanic Basin, was reviewed [63], mostly autochthonous with short transport within the la- and new discussed with the holotype and original mate- goon habitat (Figure 5D). rial from Bayreuth [7]. The few remains of pistosaurs (Figure 8B), amphib- ians, but also protorosaurs (Figure 10) in contrast, are highly fragmented and are additionally well-rounded. 4.2 Taphonomy They all indicate further transport form a fresh water u- vial or opposite deeper marine environment, or came even The bonebed contains mainly sh remains, whereas scales from eroded underlaying layers with nal deposition into were weathered often too much for determination. The a lagoon facies (allochthonous, Figure 5D). shark teeth are dominated by Hybodus and Acrodus. Their incompleteness are not the results of transport mainly. They are used teeth or have broken and polished tips, and 5 Discussion seem to represent dominantly replacement teeth; there- fore roots are missing. Abundant sh remains are the teeth of Saurichthys, Birgeria and Colobodus (Figure 3B). 5.1 Bonebed layers in the Germanic Basin There is less than one reptile nd of a size larger and their dierent ages than ve mm per square metre (Figures 3-4). Those bones were mapped on a 160 square metre surface (Figure 4) In Germany, several bonebed sites in “Upper Muschel- and were excavated on much larger areas (total about 600 kalk” quarries contain dierent aged bonebed layers of the square metres). Those generally are non-elongated bone Middle Triassic (upper Anisian/lower Ladianian). Those pieces, which have no use for palaeocurrent interpreta- layers are more abundant to the coastal facies, espe- tions. Bones are scattered irregularly across the bonebed cially terebratulid-rich tempestite facies (Figure 11A). The surface and are medium to strongly fragmented. Gener- oldest Upper Muschelkalk bonebeds are from the pul- ally, they are not well-rounded (Figures 5A-C) and have not cher/atavus Biozone at Bad Sulza [10] and Bayreuth [7]. At

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Figure 7: Paranothosaurus giganteus Meyer, 1834 remains from the enodis/posseckeri bone bed of Lamerden, Germany. 1. Anterior or lat- eral fang tooth (LAM-632), lateral. 2. Anterior fang tooth (LAM-1486, No. 12 map Figure 4), lateral. 3. Nasal of a large animal (LAM-833), dorsal. 4. Dentary fragment of a lare animal with well visible alveoloar grooves (LAM-2115), lateral 5. Humerus shaft (LAM-830), dorsal. 6. Sauropterygian carpalia/tarsalia (LAM-1646). 7. Sauropterygian carpalia/tarsalia (LAM-1668). 8. Sauropterygia carpalia/tarsalia (LAM- 643). 9. Sauropterygian metapodial (LAM-1637), dorsal (skeleton reconstruction from [7]. 10. Cervical neural arch (LAM-1482, No. 7 map Figure 4), lateral right. 11. Dorsal neural arch (LAM-839), lateral left. 12. Dorsal neural arch (LAM-627), caudal. 13. Cervical vertebra centrum (LAM-1501, No. 27 map Figure 4), a. cranial, b. dorsal, c. lateral. 14. Dorsal vertebra centrum and articulated neural arch (Lam-2116), cau- dal. 15. Thoracic rib fragment (LAM-1513, No. 39 map Figure 4), lateral. 16. Thoracic rib fragment (LAM-1512, No. 38 map Figure 4), lateral. 17. Thoracic rib fragment (LAM-1478, No. 3 map Figure 4), lateral. 18. Thoracic rib fragment (LAM-635). 19. Sacral rib (LAM-1571), lateral. 20. An- terior caudal rib (LAM-1534, No. 60 map Figure 4), ventral. 21. Distal gastral rib fragment (LAM-1480, No. 5 map Figure 4), lateral. 22. Middle gastral rib (LAM-1520, No. 46 map Figure 4), dorsal.

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Figure 8: A. Simosaurus gaillardoti Meyer, 1842 remains from the enodis/posseckeri bone bed of Lamerden, Germany. 1. Interclavicle frag- ment (LAM-1476). 2. Ulna fragment (LAM-638). 3. Cervical vertebra centrum of (LAM-626). 4. Dorsal neural arch (LAM-without no.), caudal. 5. Tibia (LAM-1496, No. 2 map Figure 4), lateral (skeleton reconstruction from [7], reconstructed after the Simosaurus skeleton casts in the SNSD and SMF and original in the UT) (skeleton reconstruction from [9]). 6. Gastral rib (LAM-without no.). B. Pistosaurus longaevus Meyer, 1839 remains from the enodis/posseckeri bone bed of Lamerden, Germany. 1. Dorsal neural spine (LAM-613), lateral. 2. Femur shaft rounded at the ends by transport (LAM-829) (skeleton reconstruction from [7], after the the Pistosaurus skeleton in the SMF and isolated material).

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Figure 9: Blezingeria ichthyospondyla (Fraas, 1896) remains from dierent sites of southern Germany. 1. Anterior dorsal vertebra spine from the Muschelkalk/Keuper-Grenzbonbeds (Meißner Fm, Fassanian/Longobardian boundary) of Neidenfels I (MHI-1113a), a. cranial, b. lat- eral. 2. Anterior dosal vertebra centrum from the Muschelkalk/Keuper-Grenzbonbeds (Meißner Fm, Fassanian/Longobardian boundary) of Brettenfeld (MHI-1076), a. cranial, b. lateral, c. dorsal. 3. Anterior dorsal vertebra spine from the Muschelkalk/Keuper-Grenzbonbeds (Meißner Fm, Fassanian/Longobardian boundary) of Neidenfels I (MHI-1113b), a. cranial, b. lateral. 4. Posterior dorsal vertebra spine from the Muschelkalk/Keuper-Grenzbonbeds (Meißner Fm, Fassanian/Longobardian boundary) of Neidenfels I (MHI-1113), a. cranial, b. lateral. 5. Sacral vertebra centrum from the Muschelkalk/Keuper-Grenzbonbeds (Meißner Fm, Fassanian/Longobardian boundary) of Neidenfels I (MHI-1113d), a. cranial, b. lateral, c. dorsal. 6. Base of a dorsal vertebra spine (distally incomplete) of a young animal from Lamerden (No. LAM-2034), cranial. 7. Dorsal vertebra of a young animal from Lamerden (No. LAM-722), cranial, b. lateral, c. dorsal, see Figure 5 No. 67. 8. Left scapula from the Muschelkalk/Keuper-Grenzbonbeds (Meißner Fm, Fassanian/Longobardian boundary) of Neidenfels I (MHI-1113d), dorsal. 9. Rigth humerus from the Lower dolomite bonebeds (Erfurt Fm, Longobardian) of Öhringen-Unterohrn (MHI 1345), dorsal. 10. Left pubis from the Muschelkalk/Keuper-Grenzbonbeds (Meißner Fm, Fassanian/Longobardian boundary) of Satteldorf-Neidelfals (MHI 1120), dorsal. 11. Fibula from the Muschelkalk/Keuper-Grenzbonbeds (Meißner Fm, Fassanian/Longobardian boundary) of Neidenfels I (MHI- 1113e) (Hypothetical skeleton reconstruction based according to the skeleton reconstruction of Askeptosaurus in [44] and the herein gured isolated bone remains).

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Figure 10: Lacustrine and brackish Lower Ladinian (Longobardian) temnospondyl amphibians and terrestrial reptiles. A. Mastodonsaurus giganteus (Jaeger, 1828) and B. Gerrothorax pustuloglomeratus (Huene, 1922) (modied redrawn after [58] and reconstructed after [57]). 1. Mastodonsaurus sp. tooth fragment (No. LAM-1540). 2. Mastodonsaurus sp. skull roong element (No. LAM-1562). see Figure 5 No. 100. 3. Gerrothorax or Plagiosuchus (No. LAM-1503) cranial bone fragment, see Figure 5 No. 29. C. Tanystropheus conspicuus Meyer, 1855 remains from the enodis/posseckeri bone bed of Lamerden, Germany. 1. Lumbar vertebra processus lateralis (LAM-1508a), dorsal. 2. Left lumbar vertebra processus lateralis (LAM-1508b), see Figure 4 No. 34, dorsal (skeleton reconstruction from [9]).

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Bayreuth several bonebed layers are found in nearly each is indicated by shell-rich and ceratite accumulation tem- ceratite biozone between the atavus to dorsoplanus Bio- pestites and scour-troughs or shallow marine subtidal zones [7] (Figure 11A). The compressus bonebed from Bis- channels which are also surface documented at Bissendorf sendorf (also present in Bayreuth) falls within the max- [9], the Bayreuther region [7] and in southern Germany imum ooding in the middle Upper Muschelkalk of the in many quarries [59]. The marine reptile fauna from Anisian/Ladinian boundary [9]. In Eilversen, Alverdissen, Lamerden has some dierences to the Upper Muschelkalk Lüdge, Vahlbruch, and Heesten, bones were found mainly bonebeds of Bad Sulza, Bayreuth, Bissendorf (Figure 11A) in the few younger evolutus Bonebed similar as in Bayreuth or Crailsheim. In the enodis/posseckeri bonebed of Lamer- [7, 64] (Figure 11A). In Lamerden, the evolutus and com- den, placodonts are absent. They are in contrast very pressus Bonebeds are also present, but contain few ver- frequent with abundant Placodus and fewer Cyamodus tebrate remains. The younger and rich enodis/posseckeri in the bonebeds of Bad Sulza and Bayreuth (pulcher Bonebed, of which the herein described fauna was former to sublaevigatus Bonebeds [7, 10]). Placodonts are also briey reported [17] is analysed herein with much more present in Bissendorf (compressus Bonebed [9]). Pachy- material (additional excavations after 2003). This bonebed pleurosaur remains are very abundant in the bonebed of is from the lowermost Lower Ladinian, dated by the Lamerden, such as in Bissendorf [9], but possibly less in cephalopod ceratites (see gures in [14]) of Ceratites (Gym- Bayreuth (remains unclear, because of selective collect- noceratites) enodis Quenstedt, for which steinkerns under- ing of larger bones in historic times). At Bissendorf and and overly the bonebed. Last ceratite nds above are from especially at Lamerden large Neusticosaurus populations Ceratites (Ceratites) sublaevigatus Wenger in the nal ma- of juveniles to old individuals were present with two dif- rine inuence in this region (last limestone beds, see Fig- ferent sized species of N. cf. pusillus (smaller form, max ure 2, 11A). The bonebed is little younger as the bonebeds 60 cm) and N. cf. edwardsii (larger form, max 1,2 m). Of of the Muschelkalk/Keuper boundary in southern Ger- both, composite skeleton experiments were made, which many [1–3, 24, 25, 27] (Figure 11A). The enodis/posseckeri demonstrate a short-distance transported of their well bonebed seems to correlate to the Ladinian black shales preserved and non-rounded isolated bone material (Fig- (Kalkschieferzone, upper bonebed layers) of the Monte ure 5D). Similar is the abundance of those small marine San Giorgio (Switzerland/Italy, [65]. There, many kinds of reptiles in the lagoons of the Monte San Giorgio or Perledo articulated sh and reptile skeletons are found and com- (northern Tethys), but there they are preserved mostly parable, but in those northern Tethys lagoons amphibian as full articulated skeletons including all age classes skeletons are completely absent [4, 31, 66]. [4, 42, 43, 45, 65, 66]. At both localities Lamerden and Bissendorf, and other Upper Muschelkalk vertebrate sites of the Germanic Basin, the pachypleurosaur Anarosaurus 6 Palaeoenvironments and related is rare, even in the Lower/Middle Muschelkalk bonebeds [53]. In the Upper Muschelkalk already large nothosaurs reptile record in Lamerden have evolved with the large P. giganteus and N. mirabilis [10]. The nothosaur material of Lamerden is dominated by Upper Middle/Lowermost Upper Muschelkalk vertebrate Paranothosaurus teeth, vertebra centra and rib fragments, tracks of Rhynchosauroides and Procolophonichnium were whereas Nothosaurus is rarer (Figures 6-7). This is simi- found on polygonal cracked carbonate biolaminates in lar to the fauna assemblage of Bissendorf [9]. At Bayreuth several outcrops around and at Lamerden. Those indicate complete skeletons and abundant isolated material of N. tidal at conditions at the end of the Middle Muschelkalk mirabilis is instead more dominant [7]. In the Monte San (Diemel Formation; Figures 2, 11A; [12]. The extended mud Giorgio lagoons Nothosaurus is instead extremely rare, ats are known for the entire Germanic Basin to have been whereas there the only complete P. giganteus skeleton is horse shoe crab reproduction zones [67]. known with stomach contents of pachypleurosaurs [38]. Uppermost Upper Muschelkalk: The en- Comparing all those European Upper Muschelkalk, sites, odis/posseckeri Bonebed was built under very shallow Paranothosaurus is a lagoon facies, and Nothosaurus a full sub-tidal conditions. Generally, the Lamerden bonebed marine facies related large reptile form [7, 9, 10]. represents a lagoon and few fresh water inuenced The Muschelkalk/Keuper boundary bonebeds of bonebed association (Figure 11) of the carbonate ramp southern Germany can-not be compared well, because in the east of the Rhenish Massif (Figure 11A). In Lamer- species-quantitative and taphonomic abundance statistics den the terrestrial/uvial inuence started earlier than the (per square metre) or systematic surface excavations (for Weserbergland in the north [64]. The shallow bathymetry taphonomy analyses) are missing. The studies presented

Unauthenticated Download Date | 2/2/16 9:56 PM 776 Ë C.J. Diedrich there are based on “collected” material [1–3, 24, 25, 27]. as known from pachypleurosaur species of China [71]. As Using those statistics, obviously, in those bonebeds more demonstrated in Germany (Lamerden), at other sites with amphibian remains are present compared to all Upper abundant pachypleurosaur skeletons such as Switzerland Muschelkalk bonebeds, including the Lamerden fauna (Monte San Giorgio), Italy (Perledo) and in China [e.g. assemblage. The Muschelkalk/Keuper bonebeds have al- [43, 45, 71] (Figure 5D), juvenile skeletons and remains are ready a stronger uvial inuence in the Hesse-Bavarian quite abundant as result of populations in their primary Depression within the southern Germanic Basin. The habitats. The highly abundant 50 to 120 cm long parax- full marine palaeoenvironment disappearance during the ial swimming reptiles seem to have been the base of the Erfurt Formation nally appeared also in southern Ger- reptile food chain all over the Triassic Globe, especially many [3]. In the middle Lower Keuper (Longobardian), known in the Germanic Basin, northern Tethys and China the marine fauna disappeared in the Athraconit Bonebed, lagoons. where archosaurs, thalattosaurs and temnospondyls dom- inate within a reduced lagoonary marine pachypleu- rosaur/nothosaur vertebrate fauna [59]. Finally, only ter- 6.3 Paranothosaurus – an alligator-like life restrial and brackish or fresh water reptiles and abun- dantly amphibians appear in the youngest Germanic style Basin bonebeds, the Kupferzell Bonebeds of Vehlheim The here-presented fragmentary humerus and a long dor- and Kupferzell, which represent a swamp landscape with sal spine from the Lamerden bonebed ts more to the partly brackish conditions [56–59, 62, 69, 70]. paraxial and not-highly marine “alligator-convergent like” paraxial swimming (typical low dorsal spines) Paranoth- 6.1 The vertebrate assemblage osaurus giganteus (Figure 11). Other large nothosaur bone material, such as a humerus and a long dorsal spine (typ- ical for N. mirabilis [10].) belongs to the full-marine axial Of 2,036 remains, the better preserved material consists and rowing moving “gavial-convergent like” few smaller of 1,588 determinable isolated bones and teeth, whereas Nothosaurus mirabilis (cf. new anatomic comparison of about 54% (862) are sh remains (mainly teeth). 722 are skeleton characters in [7, 10]. Both are common at many reptile bones and teeth and only four are from amphib- sites in the Germanic Basin in the Upper Muschelkalk ians; about 313 bones are not even identiable as to bone [7–9, 64], but as mentioned, in dierent amounts depend- type or genus level and are not used in the NISP analyses ing on the facies, whereas Paranothosaurus remains ap- (Figure 11B). Whereas teeth were more easily identied in pear most common in lagoons [7, 10, 38]. Stomach con- most cases on genus and species level and also some sh tents of pachypleurosaurs have been found in the P. gigan- scales, the postcranial bones (especially ribs, incomplete teus skeleton of Monte San Giorgio [38] which proves the vertebra centra) from reptiles are mostly problematic, even feeding on Neusticosaurus. This must be expected also in in its genus identications. Pelvic/pectoral or appendi- Lamerden and in the Germanic Basin or NW-Tethys at all, cle bones and even vertebra centra can be well attributed also because of the co-occurrence of Paranothosaurus and more and more to genus level within the sauropterygians Neusticosaurus in the same lagoon palaeoenvironments due to detailed analyses of skeletons and new bone g- (Figure 11). If coprolites with sh remains originate from urations of Pistosaurus, Paranothosaurus, Nothosaurus, those large “nothosaurs” [59, 69], remains unproven, but Neusticosaurus, Serpianosaurus, etc. [e.g. [4, 37–42, 44, their forms let exclude sharks as producers. 48–51, 55, 76].

6.2 Thousands of small pachypleurosaur 6.4 Absence of placodonts, pistosaurs and reptiles on the food chain base ichthyosaurs

The most frequent reptiles in the marine Middle Trias- Placodonts, especially Placodus and Cyamodus, are abun- sic lagoons, such as large young to old animal popula- dant in the brachiopod-tempestite dominated shallow ma- tions in Lamerden (Figure 11), are the small Neusticosaurus rine facies of Bayreuth or Bad Sulza [7, 10, 46]. In Bis- species, which are also found with a few articulated speci- sendorf, Placodus appears similar in Coenothyris brachio- mens in the southern Germanic Basin in the Lower Keuper pod tempestite shell beds. Those indicate not only bra- lagoons [4, 36]. Those gave birth to live young in the sea, chiopod colonies on the hard-and rmgrounds, they also

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Figure 11: A. Correlation of the enodis/posseckeri bonebed at Lamerden and its facies position. B. Faunal compositions (NISP = bone amount per species) in the bonebeds of the enodis/posseckeri bonebed in Lamerden. C. The uppermost Upper Muschelkalk/basal Lower Keuper near-beach coastal swamps and its amphibian dominated fauna, here Mastodonsaurus and Gerrothorax. In a hunting scenario of the the- codont Batrachotomus a Tanystropheus dropped its tail to confuse the predator. B. The lagoon in the uppermost Upper Muschelkalk/Lower Keuper with its typical reptile fauna in the Germanic Basin (Illustration G. Teichmann). Nothosaur Simosaurus gaillardoti, or Paranoth- osaurus giganteus hunted smaller pachypleurosaur Neusticosaurus reptile species, whereas Blezingeria ichthyospondyla predated possibly onto sh in the fresh water influenced Lamerden lagoon (Illustrations G. Teichmann).

Unauthenticated Download Date | 2/2/16 9:56 PM 778 Ë C.J. Diedrich correlate to the main habitat of macroalgae meadows and calities (Figure 9), a postcranial hypothetical reconstruc- placodonts abundance [7, 10, 46] (Figure 11B, C). Placo- tion diers to the thalattosaur Askeptosaurus italicus Nop- dus and Cyamodus or other placodonts were recently re- sca, 1925 [54]. Herein, the Blezingeria skeleton with its typ- vised to be more likely macroalgae feeder that lived in ical high dorsal spines (in Askeptosaurus very low spines) shallow marine sand bar or hardground areas which rep- is more similar in anatomy as the long spines of the Chi- tile and palaeoenvironments of Bissendorf, Bad Sulza, and nese Hupehsuchus or Hanosaurus marine reptiles, which Bayreuth supported this theory [7, 10, 46, 52]. There, shal- were also found in lagoons [74, 75]. With the add of fur- low sub-tidal environments must have been covered by ther postcranial bones from various German sites (Fig- macroalgae meadows, which is furthermore indirectly in- ure 9) the habitus (extremities and pelvic/pectoral girdles) dicated by benthic invertebrate palaeocommunities, es- have closest bone shape characters to the Early Triassic pecially algae adapted snails [7, 10, 59]. Those facies (Olenekian) Chinese marine reptile Nanchangosaurus or types were absent both in contemporaneous brackish la- again, Hupehsuchus [74] which latter resemble an anatom- goons such as in the Lamerden bonebed layer times, ical “cross” between an ichthyosaur and crocodilian. The or in general of the intraplatform lagoons of the Monte humerus presented herein (Figure 9.9) is strongly curved, San Giorgio/Perledo [65, 72, 73]. This facies related occur- massive and very distinctive to all discussed marine rep- rence explains also a nearly absence of Placodus in the tiles, so its attribution to Blezingeria is most possible, northern Tethys lagoon black shale facies (only few teeth whereas closest characters are with the humerus of Askep- nds), which is instead very frequent in the shallow ma- tosaurus italicus [44, 54, 76]. Possibly, this humerus even rine terebratulid tempestite facies of the Germanic Basin belongs indeed to this genus, and not to Blezingeria, which [7, 10, 46, 51]. would be the rst proof within the Germanic Basin of this Open marine ichthyosaurs, which are generally rare “endemic northern Tethys” thalattosaur. The scapula pre- in the Upper Muschelkalk of the Germanic Basin [22], sented herein (Figure 9.8) is also very similar in its outline are missing in the Lamerden enodis/posseckeri Bonebed, shape to the one of Askeptosaurus [44, 76]. Also similar to which are instead very abundant in the northern Tethys Askeptosaurus is the herein gured bula (Figure 9.11) with lagoons [65, 72]. its typical distally widened shape [44]. From the pelvic, Also the few open marine Pistosaurus longaevus re- herein a pubis with its typical open foramen obturatum mains found in the Lamerden bonebed seem to have been seems to be present from Blezingeria as the only pelvic el- transported from more far (or reworked underlaying lay- ement. This is distinct in its polygonal shape to the one ers) into the Lamerden lagoons. The dorsal spine and fe- of Askeptosaurus (Figure 9.10) [44, 76]. Based on the axial mur (Figure 8) are strongly rounded and fragmented. Also skeleton only, which can be reconstructed best at the mo- this reptile, which is recently demonstrated to have been ment only, Blezingeria with its high neural spines and typ- very common in the Germanic basin including a skeleton ical ichthyosaur-like vertebra centra is better to position [50], is missing in the northern Tethys lagoon record [com- within the nanchangosaurs, instead of the thalattosaurs, pare [50] and [65]]. which have very dierent vertebral column morphologies [44, 74]. The rst preliminary postcranial reconstruction of 6.5 Blezingeria – a skeleton reconstruction Blezingeria (Figure 9) is therefore based of the Chinese approach nanchangosaurs. The axial skeleton and the few (clearly non-sauropterygian/ichthyosaur) appendicle bones pre- Blezingeria vertebra centra were attributed rst to a sented herein from German sites might nally belong to nothosaur “Nothosaurus ichthyospondylus” by [36] and dierent reptile groups. Those rare and complete bones were later incorrectly identied as ichthyosaur remains demonstrate the problem for their identication, whereas [39]. After new interpretations, they are thought to be- their origin of all other sauropterygians or amphibians long to marine thalattosaurs. Those typical vertebrae are can be excluded well. Possibly here both, thalattosaur and often found in the Muschelkalk/Keuper Bonebeds and nanchangosaur material is gured (in such a case then Lower Ladinian (Lower Keuper) bonebeds of southern Ger- combined incorrectly), which can not be solved yet. The many [44, 69]. Material of Lamerden is mostly smaller as presentation of this rare material is important for future maximum sized vertebrae of southern Germany, which discussions, cladistic analyses concerning evolution and might indicate younger individuals, or even dierent gen- palaeobiogeography of those rare marine reptiles, which era/species. Using those typical centra and isolated dor- seem to be also distributed much more wide globally, as sal spine material from Lamerden and other German lo- believed before.

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The presence of Blezingeria vertebrae/dorsal spines 6.7 Conclusions in the enodis/posseckeri Bonebed lagoon sediments of Lamerden (Figure 11) and more abundance in the Muschel- Benthic palaeocommunities and bonebed layers of kalk/Keuper boundary Bonebeds of southern Germany is the “Upper Muschelkalk” in the Germanic Basin and obviously a result of an adaptation to the food-chain and their dierent stratigraphic occurrence depend on the brackish or lagoonary conditions. The long dorsal spines intracratonic Germanic Basin development and mor- let think about an axial ichthyosaur-like swimming style, phology. Those layers spanning about three My time whereas in Askeptosaurus those dorsal spines are quite frame in several quarries built a perfect basis for short (centra instead are very long), instead those are sim- the understanding of marine reptile palaeobiogeogra- ilar long in Hupehsuchus or Hanosaurus [44, 75]. The legs phy/phylostratigraphy/biodiversity and evolution. First and pectoral and pelvic girdle are expected similar as in Upper Muschelkalk Illyrian-Fassanian aged bonebeds Askeptosaurus, to be reduced, and therefore, leg propul- are found at small scaled sequence boundaries (pulcher, sion was not the locomotion motor. atavus, evolutus Bonebeds), around the maximum ood- ing (compressus Bonebed), at the transition from marine to brackish conditions (enodis/posseckeri and Muschel- kalk/Keuper boundary Bonebeds) and the nal brack- 6.6 Terrestrial reptiles and amphibians ish/limnic stage (Lower Keuper Bonebeds). The faunal analyses of reptile/amphibian assemblages of those dif- The few material of Tanystrophaeus from the Lamerden ferent bonebed layers support bathymetrical, salinity and bonebed is strongly fragmented and rounded, and there- palaeoenvironmental reconstructions of the Germanic fore has longer/further transport, most probably from the Basin. Those are very dierent at the herein compared Rhenish Massif mainland direction. Also other more ma- bonebeds of Bayreuth, Bad Sulza, Bissendorf and Lamer- rine dominated bonebeds rarely contain few remains of den (all Germany). The facies-related reptile faunas sup- Tanystrophaeus which were found with teeth e.g. in the port the reconstruction of the presence or absence of compressus Bonebed of Bissendorf [9]. Bones are most macroalgae meadows, which placodont marine reptiles common at the Bayreuth sites, where the holotype T. con- (Placodus, Cyamodus, Paraplacodus) depended on as their spicuus Meyer, 1855 was established on isolated bones [7, main food source. Those are absent in Lamerden in the en- 8, 63]. Tanystropheus was a terrestrial coastal adapted pro- odis/posseckeri Bonebed (Meißner Formation, Fassanian). torosaur reptile [63, 68] (Figure 11), which carcasses drifted Placodonts instead were abundant in shallow marine also into the the Monte San Giorgio and Perledo lagoons, terebratulid-rich shallow marine palaeoenvironments, es- together with several other terrestrial reptile skeletons [12, pecially in the Bad Sulza, Bissendorf and Bayreuth areas. 65]. The terrestrial reptile was partly revised [68] and was In Lamerden the small pachypleurosaurs are the domi- corrected anatomically in its limb and neck biomechanics nant reptiles, similar as in the northern Tethys Monte San and positions [77], which was further corrected [12]. The zy- Giorgio (Kalkschieferzone black shales) and Perledo Lon- gapophyses of all vertebrae, the elongated and with long gobardian aged (Meride Limestone black shales) lagoons. neck ribs “xed” neck was constructed for a life on land In both regions, large nothosaurs, especially Paranoth- similar as in Macrocnemus [12]. Tanystropheus must have osaurus giganteus, seem to have been their main predator. hunted on the beach areas similar as gured in rst recon- Dierent to the Monte San Giorgio/Perledo, in Lamerden structions [68], whereas the elongated neck allowed the limnic amphibian remains result form freshwater inu- reptile to hunt deeper into the beach shore. Finally, the ence at the nal marine basin stage on is costs east of the new discoveries of a skeleton from Perledo reproduced an Rhenish Massif. The partly brackish lagoons of Lamerden unrecognized “cartilage” regrowing tail [77], which is only contain few and far transported temnospondyl remains known in modern lizards today. Those might have been at- (most probably Mastodonsaurus, Plagiosuchus, and Ger- tacked most probably by the large thecodont Ticinosuchus rothorax), but also terrestrial protorosaur Tanystropheus (Figure 11C), or other thecodonts. and marine pistosaur Pistosaurus reptile remains which Mastodonsaurus and the other temnospondyls such were accumulated. The most bone material is from small as Gerrothorax and Plagiosuchus had their habitats in the to medium-sized Neusticosaurus species of dierent indi- limnic swamp and river environments of the Rhenish and vidual ages. Blezingeria remains from the Lamerden and Bohemian Massif costal areas (Figure 11C), but those large southern German lagoon deposits allow a hypothetical amphibians might have been adapted also to brackish con- reconstruction of the postcranial skeleton. Those thalat- ditions [57, 58]. tosaurs are herein revised to belong to nanchangosaurs

Unauthenticated Download Date | 2/2/16 9:56 PM 780 Ë C.J. Diedrich which obviously adapted to limnic inuenced lagoons. [8] Meyer von H., Zur Fauna der Vorwelt. 2 Abt. Die Saurier des Nanchangosaurs were found with dierent forms in Ger- Muschelkalkes mit Rücksicht auf die Saurier aus Buntem Sand- many and China, but are absent in the normal saline la- stein und Keuper. Frankfurt a. Main, 1847-1855, 8-175. [9] Diedrich C., The vertebrates of the Anisian/Ladinian boundary goons of the Monte San Giorgio/Perledo. (Middle Triassic) from Bissendorf (NW Germany) and their con- tribution to the anatomy, palaeoecology, and palaeobiogeogra- Acknowledgement: For the excavation permissions, and phy of the Germanic Basin reptiles. Palaeogeogr. Palaeoclima- transfer of the owner rights of the bone material, I thank tol. Palaeoecol., 2009, 273, 1-16. the Fischer GmbH und CO KG, especially A. Fischer. They [10] Diedrich C., Shallow marine sauropterygian reptile biodiver- supported the excavations each year with large machines. sity and change in the Bad Sulza Fm (Illyrian, Middle Trias- sic) of Central Germany - and a contribution to the evolution of Dr. J. Fichter supported the rst campaigns. I thank for Nothosaurus in the Germanic Basin. New Mex. Mus. Nat. Hist. comparative bone material Dr. R. Schoch of the Staatliche Sci., 2013, 61, 132-158. Sammlung für Naturkunde Stuttgart. Also Dr. R. Werneb- [11] Kozur H.W., Bachmann G.H., Updated correlation of the Ger- urg (head Naturhistorische Museum Schleusingen) al- manic Triassic with the Tethyan scale and assigned numeric lowed the comparison to Thuringian material. Prof. Dr. R. ages. Ber. Geol. Bundesanst., 2008, 76, 53-58. [12] Diedrich C., Palaeogeographic evolution of the marine Middle Springhorn of the Lippische Landesmuseum Detmold gave Triassic marine Germanic Basin changements - with emphasis access to the Weserbergland-Muschelkalk collection. Dr. on the carbonate tidal flat and shallow marine habitats of rep- H. Hagdorn allowed the study of the Muschelkalkmuseum tiles in Central Pangaea. Glob. Planet. Change, 2008, 65(2009), Ingelngen material. I thank S. Rein for some ceratite de- 27-55. terminations. The excavations, ocially permitted by the [13] Fredd D., Unser Traum - ein Massenvorkommen von Encrinus province Hesse (Denkmalbehörde), were undertaken by liliiformis in Lamerden. Arbeitskreis Paläontologie Hannover, 1994, 3, 61-80. the support of the company and private research institute [14] Diedrich C., Tsunami killed and backwashed accumulated PaleoLogic (www.paleologic.eu). Prof. Dr. M. Benton, Dr. F. crinoids in Middle Triassic intracratonic Germanic Basin carbon- Witzmann, and Dr. J. Liu made helpful critical reviews. G. ates of Central Europe. Carbonate. Evaporate., 2015, (in press). Teichmann illustrated nally the Lamerden lagoon fauna [15] Diedrich C., Ein Ceratiten-Pflaster aus der penndor-Zone situation. des Oberen Muschelkalkes (Mitteltrias) Nordhessens (Nord- Deutschland). Philippia, 2003, 11(2), 93-102. [16] Fürsich F.T., Die Entalis-Faunengemeinschaft in Weichböden des Oberen Muschelkalkes. In: Mc Kerrow W.S., (Ed.), Palökolo- References gie. Schweizerbart, Stuttgart, 1981, 134-135. [17] Diedrich C., Die Wirbeltier-Fauna aus dem Bonebed der [1] Reif W.E., Zur Genese des Muschelkalk/Keuper-Grenzbonebeds enodis/possseckeri-Zone des Oberen Muschelkalkes (Mittel- in Südwestdeutschland. Jahrb. Geol. Palaeont., 1971, 139, 369- trias) von Lamerden (NW-Deutschland). Philippia, 2003, 11(2), 404. 133-150. [2] Reif W.E., Muschelkalk/Keuper bone-beds (Middle Triassic, SW- [18] Agassiz L. Recherches sur les Poissons fossiles. 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