18x6 m fossilized floating wood (Agathoxylon), with Crinoids attached (Pentacrinites & Seirocrinus). It is one of the most emblematic fossils of the formation, where the anoxic seas of the lower toarcian lead to an exquisite preservation.
The Sachrang Formation or "Posidonienschiefer" Formation (Whose Vulgar name is "Posidonia Shale") is a geological formation of southwestern Germany, northern Switzerland, northwestern Austria, southeast Luxembourg and the Netherlands, that spans about 3 million years during the Early Jurassic period (early Toarcian stage). It is known for its detailed fossils, especially sea fauna, listed below.[1] Composed mostly by black shale, the formation is a Lagerstätte, where fossils show exceptional preservation (Including exquisite soft tissues), with a thickness that varies from about 1 m to about 40 m on the Rhine level, being on the main quarry at Holzmaden between 5 and 14 m.[1] Some of the preserved material has been transformed into fossil hydrocarbon Jet, specially wood remains, used for jewelry.[2] The exceptional preservation seen on the Posidonia Shale has been studied since the late 1800s, finding that a cocktail of chemical and environmental factors let to such an impressive conservation of the marine fauna.[2] The most common theory is the changes in the oxygen level, where the different anoxic events of the Toarcian left oxygen-depleted bottom waters, with the biota dying and falling to the bottom without any predator able to eat the dead bodies.[3]
Biological interaction frozen in time
Several animal behaviours have been recovered on the Posidonia Shale. The Monotis-Dactylioceras bed is one of them, as it shows an accumulation of the bivalves Meleagrinella substriata and the ammonite Dactylioceras, that were the most abundant representatives of its group on the Altdorf region, and were probably washed to near epicontinental waters by a rapid event, or as result of a large succession of events.[4] This assemblage has been compared with modern Brazilian coastal Mangroves and also linked to Tsunami events.[5]
Related to the Ammonite fauna, on Holzmaden there have been found several empty shells of this cephalopods, with associated crustaceans inside.[6] The original specimen was reported in 1995 and consisted on a possible member of the genus Paleastacus inside a chamber of a Harpoceras.[6] Other epizoans are found related to the decayed Ammonite shells, such as Serpulid Annelids and Bivalves, creating what was denominated as "benthic islands" as reference at be isolated benthic units that attracted fauna.[6] The Decapod is related to the family Erymidae, that are considered as possible bottom-dweller carnivorous or carrion feeders.[6] The associated fossil has several spherical structures that had been interpreted as decapod coprolites, implying that the animal lived for a long period on the shell, and maybe changes on the bottom oxygen stopped the process.[6] More recent studies had recovered new data about the inquilinism of decapods inside ammonites, this time, however, recovering three Eryonoidea lobsters together within a body chamber.[7] The lobsters most likely used the ammonoid as some kind of shelter, where was excluded the possibility of transportation into the body chamber by bottom currents.[7] There are several theories about the gregarious inquilinism showed by this specimens, such as that the shell was and ideal location to molt, as has not been proved to be corpses or molts; that the shell provided protection against predators; the decomposing soft body of the ammonoid was a source of food that attracted the decapods or that was used as a long-term shelter.[7] One key aspect found was that the muddy bottom was not suitable for burrowing, implying that the decapods look for a different shelter due to being unable to make their own.[7]
Beyond trace fossils, several vertebrate specimens show associations with crustacean exoskeletal remains such as GPIT-PV-31586 and SMNS 58389 (Pachycormus macropterus) with necrophagous interaction as well SMNS 55934 (Stenopterygius quadriscissus) or SMNS 95401 (Metopacanthus sp.).[8]
Fossil of Clarkeiteuthis preying on Leptolepis
The genus Clarkeiteuthis and its predatory behaviour, found associated with fishes of the genus Leptolepis.[9] Based on the position of prey and predator, was suggested that the coeloid cephalopods caught and killed the fishes while the schools still in well-oxygenated waters and then descended into oxygen-depleted water layers where the cephalopod suffocated and died attached to its prey.[9] The fish measured 12 centimetres (4.7 in), while the coeloid 21 and it was measured by the fossilized arms of 14 specimens of coeloids that the hunting specimens arms where contracted over the fish, probably quickly killing it by cutting its spine.[9]
Several Geotheutis have been reported with eumelanin preserved along with its ink sacs.[10]
A specimen of Jeletzkyteuthis found on Ohmden has appeared predating a Parabelopeltis. The association of this 2 genera shows the predatory behaviour of this group when lived on Epicontinental seas, being rather different than extant Vampyromorphs.[11]
A Pabulite (fossilized meal when it never entered the digestive tract) was recovered on Holzmaden, being composed by an associated Passaloteuthis laevigata with its arms embracing an exuvia of a crustacean.[12] The own Belemnnite can be the remnant of a failed prey of a Hybodus, corroborating a possible tropic chain.[12]
Seirocrinus subsingularis stems over a branch
One of the most complex organism interactions on the Posidonia Shale where the crinoid megarafts, that group a wide variety of animals, creating large floating ecosystems, being the longest surviving communities to exist in the fossil record.[13] The largest megaraft found measured 18 metres (59 ft), and is based on an Agathoxylon trunk, where different animals were attached.[13] The first attached animals would have been the growing community of oysters, bivalves and crinoids, that would suppose and small weight to the raft about 800 kilograms (1,800 lb).[13] The presence of this megaraft was in part possible due to the absence of marine wood worms, that destroy tree logs on less than 3 years along without the presence of modern raft wood predators (that appeared on the Bathonian) those rafts can last up to 5 years, being that the main reason the crinoids attached were able to reach huge sizes.[13] Probably where also essential to distribute animals along the sea basins.[13]Seirocrinus & Pentacrinites where various of the main crinoid colonizers of the floating rafts.[14]Seirocrinus is the main representative of the pelagic crinoids, being among the tallest animals know, with a size of 26 m the largest documented specimen.[14] The ecology of the genus is widely known, where is known that the smallest stems were among the first animals to colonize the rafts, with at least 2 generations of crinoids found per raft, where the hydrodynamic changes of the log influenced the settlement of the crinoids.[14] It is believed that Seridocrinus had a seasonal reproduction, linked to the monsoonal conditions that sent new logs to the sea.[14] The large crinoids would have feed on pelagic micronutrients, and afer fall on the bottom, all the colony would have died.[14]
Thoracic cirripedes of the genus Toarcolepas became the oldest epiplanktonic cirripede known on the fossil record, probably motivated by the appearance of the giant crinoid rafts. It has been found in situ associated with fossil wood.[15]
Hybodus hauffianus with skin and Belemnnite traces
The shark Hybodus includes specimens with the gastric contents, being full of belemnnite fragments.[16] That implied active predatory behaviour by the genus of several kinds of belemnnites, such as Youngibelus.[16]
A Spienballen, a regurgitated mass composed of indigestible stomach contents had been found on the Holzmaden quarry.[17] The Speinballen measures 285 mm length with a diameter of 160 mm, and consists of 4 members of the genus Dapedium (Dapediidae) and a jaw identified as Lepidotes (Semionotidae).[17] The animals capable of it had been suggested as sharks like Hybodus, actinopterygians and several marine reptiles.[17]Hybodus, being was able to reach nearly 3 meters long and with a dentition suitable to hunt fish, although its stomach contents suggest it is a mostly invertebrate hunter.[17] Actinopterygians like Saurostomus grew up to 2 m long, and have been found with fishes, coeloids and ammonites in its stomach contents, however, not the fishes present on this Spienballen.[17] Marine reptiles included marine crocodiles, such as Platysuchus or Pelagosaurus, associated to the fishes of the Speinballen, although are proven to have eaten gastroliths to improve buoyancy and digestion.[17] Ichthyosaurs, whose diet is among the best studied of the Posidonia Shale, with Dapedium specimens in juvenile stomachs, along with coeloids. Temnodontosaurus, measuring between 8 and 13 metres (26 and 43 ft), would have been able to do such a large Speinballen.[17] Plesiosaurs were disaccredited due to the study of its teeth, that proves a diet based on soft-body prey, such as fishes of the genus Leptolepis and coeloids. Dapedium and Lepidotes, with a heavy and solid squamation can be excluded.[17]
A specimen of Pachycormus has been found with stomach contents that include hooks similar to the ones found on genera like Clarkeiteuthis.[18]
Another specimen of Pachycormus macropterus preserves an ammonite inside its gut, likely swallowed by accident and directly responsible for the fish’s death.[19]
SMNS 51144 (Saurostomus esocinus) was found with Chondrites isp. burrows in the abdominal cavity, what indicates a possible opportunistic scavenger. Other Chondrites isp. includes SMNS 17500 and MHH 1981/25 (Stenopterygius uniter) that can either suggest ichthyosaurs were preserved immediately below one such bioturbation horizon or scavenger association.[8]
Stenopterygius quadriscissus, mother with embryo
One of the most emblematic finds of the formation its that of a mother Stenopterygius giving birth living young, like the modern dolphins and marine mammals, being born with the tails first. Other specimens have been found with Embryos inside, but with the bones of them scattered, partly beyond the body limits of the mother.[20] There have been various theories about this scenario: the bones of embryos had been deposited before the body of the adult went to the sea floor, covering the embryo bones and implying that the adult would not be the mother of the embryos.[20] Another option is that a pregnant ichthyosaur on its last moments sank to the bottom and may have struggled for life, given untimely birth to some of the foetuses.[20] Other option follows the presence of foetus bones outside the mother body, where a dead female sank to the bottom, with the water warm enough, helping the putrefaction gases to start to develop while the hydrostatic pressure was too high to be prevented by the body. Scavengers must have started eating from the dead body, until the chamber retaining the pressure was to thin and exploded.[20] These theories where however contested after the study, where it was criticised the absence of the presence of the bottom-current activity in the epicontinental sea covering Central Europe during the Toarcian, pointing that the mother carcass should have been translated after it sank to the bottom floor, probably exploding or expelling its embryos first, that would be transported along.[21]
Specimen SMNS 53363 (Eurhinosaurus?) from Aichelberg was found with two encrusted large oysters (Liostrea) on the right pterygoid, considered to be part of a reef stage over bones.[8]
SMNS 80234 (Stenopterygius quadriscissus) represents another female with embryos, yet also shows ribs broken perimortem that can be either of intraspecific aggression or a predation attempt. This specimen has several taxa associated: ammonite aptychi and two ophiuroids (Sinosura brodiei) and a articulated echinoid (Diademopsis crinifera), indicating a short-lived deadfall community.[8]
SMNS 81841 (Stenopterygius quadriscissus) represents one of the most clear examples of deadfall communities described in the formation: the skeleton is associated with serpulids surrounded by a mass of disarticulated ophiuroid remains, indeterminate echinoid tests, an isolated crinoid ossicle, the byssate bivalve Oxytoma inaequivalvis, the pectinid Propeamussium pumilus, Eopecten strionatis, Plagiostoma sp. , Meleagrinella sp., "Cucullaea" muensteri, with the genera Parainoceramya dubia and Liostrea associated with the carcass.[22] As many of this bivalves shown overgrowth likely the community persisted for some time.[22] Fossil traces of Gastrochaenolites isp. attributed to mechanical bivalve borers are abundant implicating prolonged exposure of the skeleton on the seafloor.[22]
SMNS 81719 (Stenopterygius uniter) includes Liostrea, Propeamussium pumilus, Plagiostoma sp. and Parainoceramya dubia, with other invertebrates found (?) not being part of the deadfall community, such as several ammonites and Parainoceramya valves stratigraphically below the specimen.[8] This specimen includes also traces of scavenging activity, possibly by crustaceans.[8]
SMNS 80113, (Stenopterygius triscissus) was found populated by Parainoceramya, a specimen of Eopecten strionatis and an unexpected specimen of the small infaunal lucinid Mesomiltha pumila, equivocal evidence for the sulfophilic stage.[8]
Local ichthyosaur soft tissues include skin enough well preserved to infer coloration and appearance on the living animal, as well evidence for homeothermy and crypsis.[23]
Microbial activity
Non-fenestrate Stromatolite crusts formed in Aphotic deep-water environments during intervals of very low sedimentation are recovered in places such as Teufelsgraben, Hetzles.[24] The Stromatolites of this region have evidence of live on a deeper shelf environment with a quietwater deposit which suffered repeated phases of stagnant bottom waters, where a depth water habitat developed, probably at more than 100 meters depth.[24] There is a thin, southern widespread Stromatolite crust on the Top of the Sachrang Formation, called "Wittelshofener Bank", that has made rethink the depth of the major southern basin of the formation, where with the absence of phototrophic calcareous benthic organisms (probably due to the lack of light), shows the depth character of the basin.[24] On the "Wittelshofener Bank" there is also the only occurrence of Ooids, presumably formed in the same deep-water environment.[24]
Probably related with Archaea activity.[24] Although Frutexites is a cryptic microfossil and an important element of many deep water stromatolites, with an Inorganic origin proposed, where are interpreted as dendritic shrubs to purely inorganic growth of Aragonitic crystals, but also resemble shrubs of the cyanobacteria Angulocellularia.[24] On the Posidonia a cryptoendopelitic mode of life is assumed, being only possible for Heterotrophic bacteria or Fungi.[24] As seen on the Stromatolites of the Posidonia, Frutexites acted mainly as a dweller or secondary binder of the deep-water stromatolites, not as their major constructor.[24]
A Cyanobacteria, member of the family Oscillatoriales. The only difference compared to the Paleozoic forms is the much thinner cellular wall of the specimens described, which otherwise seems to be always thicker, and the more irregular shape of the individual tubes. Girvanella is almost rock-forming in the Lower and Upper levels, and is very common, but can only rarely be detected in the bituminous clay marl slate due to conservation reasons.[25] The Girvanella specimens recovered on the Posidonienschiefer can come from Diatomeas rather than Cyanobacteria, but its assignation is rather controversial.[25]
A benthonic Foraminiferan, member of Vaginulinidae inside the family Vaginulinida (Lagenina). An extant genus. Its shell resemble a mixture between an ammonite conch, due to having a lower spiral, and a mussel.
A benthonic Foraminiferan, member of Nodosariidae inside the family Nodosariacea (Lagenina). Dentalina is an extant genus, with an elongated shell, that resemble a small worm.
A benthonic Foraminiferan, type member of Vaginulinidae inside the family Vaginulinida (Lagenina).
Dinoflagellata
Dinoflagellate cysts
The evolutionary burst of the Toarcian Dinoflajellates led the first appearance and rapid radiation of the Phallocystaceae (Susainium, Parvocysta, Phallocysta, Moesiodinium and related forms).[28] This occurred at the time of a widespread Lower Toarcian bituminous anoxia-derived shale of the Posidonienschiefer Formation. Is recovered on the Posidonienschiefer, Pozzale, Italy, Asturias, Spain, Bornholm, Denmark, the Lusitanian Basin of Portugal, the Jet Rock Formation in Yorkshire and to the "Schistes Carton" in northern France. Whether there is a causal connection in this co-occurrence of Phallocystaceae and bituminous facies is a problem still to be resolved. This family has its acme in diversity and quantity in the latest Toarcian and became less important in the Aalenian.[28]
A Dinoflagellate cyst, type member of Luehndeoideae. Luehndea spinosa is common on the medium layers of the lower Sachrang Formation, while restricted to some areas on the Lias delta.[31]
A Dinoflagellate cyst, member of Dinophyceae of the family Nannoceratopsiaceae. On the Lias Epsylon Interval (Lowermost Toarcian), most of the assemblages are dominated by Nannoceratopsis gracilis. Nannoceratopsis senex becomes highly abundant until the uppermost Tenuicostatum.[31]
A Dinoflagellate cyst from the family Comparodiniaceae.
Algae
Includes abundant variety of algae, such as the genus of colonial Green algae Botryococcus, or the unicellular algal bodies Tasmanites, and other small examples. Algae are a good reference for changes on the oxygen conditions along the Toarcian.[34]
An Acritarch probably from Algal origin. Its fossils indicate nearshore or estuarine to shallow lagoon and/or slightly brackish-water environments. It is the dominant on the nearshore sections.
An Acritarch probably from Algal origin. Veryhachium fossils represent open marine and transgressive conditions. It has high presence on most of the samples studied from the Sachrang Formation, being nearly the 50% of the Acritarch fraction on some locations.
A member of the family Parhabdolithaceae inside Stephanolithiales. Shore deposits genus. The abundance drop of M. jansae further characterise the T-OAE perturbation, where becomes the dominant Genus on most of the Saxony Basin.
Type member of the family Schizosphaerellaceae inside Parhabdolithaceae. Towards the Pliensbachian-Toarcian extincion this genus gets a decrease in abundance and size that shows the change and biotic crisis.
Fungal Spores, hypae and undeterminated remains are a rare element of the otherwise openmarine deposits of the Posidonienschiefer formation, but where recovered at Dormettingen.[43] This fungal remains are composed mostly by indeterminate spores and indicate oxygenated environments and suitable transportation by rivers.[43]
Fungal patches in ammonite shells and belemnite rostra
A Marine parasitoid Fungus of uncertain relationships, linked with shells of marine invertebrates. Extant Ostracoblabe implexa is usually found associted with Bivalve shells as an external parasiotid. Beyond this genus, other fungal remains include indeterminate endolithic fungi linked with microbial mats
Ichnofossils
The major ichnological analyses of the Posidonian Shale come from Dotternhausen/Dormettingen, where the ichnogenus Phymatoderma formed the so-called Tafelfleins and Seegrasschiefer.[45] The Tafelflein bed was deposited under anoxic bottom and pore water, where a recover of oxygen allow the Phymatoderma-producers return.[45] The two organic-rich layers (Tafelfleins and Seegrasschiefer) are characterized by the dense occurrence of trace fossils such as Chondrites and Phymatoderma, done episodically due to the fall of the oxygen levels.[45] The Coeval more nearshore Swiss deposits referred Posidonian Shale (Rietheim Member) hosted similar trace fossils asthose recovered on SW Germany.[45] Tougth this setting apparently evolved faster to more oxic-to-dysoxic bottom waters.[45]
At Unken, laminated deposits of red limestone suggest well oxygenated active waters (as lack shale), where high amounts of Chondrites are found.[37]
Burrow-like ichnofossils. Interpreted as the feeding burrow of a sediment-ingesting animal.[48] A more recent study has find that Scoloplos armiger and Heteromastus filiformis, occurring in the German Wadden Sea in the lower parts of tidal flats, make burrows that are homonymous with numerous trace fossils of the ichnogenus.[49]Chondrites burrows from Holzmaden are mostly filled with granular Calcite crystals, clay minerals, and rare framboidal Pyrite. The local dysoxic seawater is reflected on the palaeoredox conditions, relating framboidal pyrite and biogenic processes. Chondrites appear to be capable of colonizing environments characterized by oxygen levels well below levels needed for survival of other animals, being considered an "extremotolerant" ichnotaxon.[50]
Burrow-like ichnofossils. It consists commonly on a subhorizontal branching burrow system consisting of radiating tunnels filled with fecal pellets.[53] It has been interpreted as a product made by an Endobenthic deposit-feeding animal, specially a Fodinichnia, burrows produced by benthonic subsurface food-mining activity, as is proved by the tunnels and pelletal infill.[53] The study of the Fecal Pellets has revelated that the maker of this ichnogenus was an epicontinental shelf setting non-selective deposit feeder, ingesting particles on the sediment surface without selection. A mode of feeding common on aquatic Benthos, reported on modern animals such as Spionid Polychaete worms, tropical Holothurians and Spatangoid urchins.[54]
U-shaped tubes with sinuous, bifurcating or planispiral spreite, that can be related to Crustaceans, Annelids and Fishes, being both Domichnia and/or fodinichnia.
Burrow-like ichnofossils, that can be related to Crustaceans, Annelids and Fishes.[55] The presence of this burrows changed along the different depositional layers, interpreted as result of relative magnitudes and durations of a series oxygenation events.[55] Increased Oxygen conditions eventually led to a level that permitted both the survival of larger Chondrites and Thalassinoides producing organisms, as well the depth of the Burrow-like structures.[55] The changes on the layers are detailed enough to know that oxygenation-change events duration was sufficient to allowe the migration and establishment of trace-producing organisms, establishing an "equilibrium" with bottom-water oxygen conditions.[55]
Burrow-like ichnofossils. It has been related to Echiuran annelids, but also from moving and feeding polychaete worms.[57]
Example of Zoophycos fossil
Invertebrata
Porifera
In the non-bituminous facies located on Obereggenen im Breisgau (Shore of the Black Forest High), especially the lower semicelatum subzone, pyritized individual needles of silica sponges (Demospongiae and Hexactinellida) are found, rarely on pelagic layers to very often on the low depth marine deposits.[25] They are usually associated with radiolarian stone cores. In Dusslingen and Reutlingen, these sponge needles could be barytized in phosphorites of the Haskerense subzone and are much more common here than in any other zone of the Lower Toarcian. These needles are absent in the bituminous horizons of the entire Lower Toarcian.[25]
Increased amounts of Sponge needles (dominated by Hexactinellida) are also found on the arenaceous facies of the nearshore unit that is the Unken member, being the only section if its region hosts them, probably due to be an active and well oxygenated bottom.[37] The location of this member as a possible bay on the south of the vindelician land probably allow to the development of more pre-Toarcian AOE conditions, hence the presence of biota otherwise rare on bituminous layers.[37]
A sessile, marineannelidtube worm of the family Serpulidae. Its affinities with the genus Serpula are controversial, since the genus is known mostly since Cretaceous strata. Although there are other fossils assigned to the genus on same age deposits of France. Presumably these specimens have fallen from their growth areas.[25]
A polychaete of the family Dorvilleidae inside Eunicida. It was considered as part of the genus Halla. Eunicidan species with prionognath jaws, absent on Bituminous layers
A OncousoeciidaeStenolaematan. Colonies consists of bands that are the same width throughout their entire extent and can branch, without fan-shaped spread
A Lingulidaerhynchonellatan. Associations of bioturbation infauna are dominated on certain sections by Palaeonucula/Lingula agrupations, developed under longer-term oxygenated conditions within the substrate and bottom waters.[59]
A Discinidaerhynchonellatan. This genus was found had a planktotrophic larval stage, that adapted while growing to the local redox boundary, when this fluctuated near the sediment–water interface and oxygen availability prevailed, allowing benthic colonization. Is found on associations with Grammatodon and Pseudomytiloides.[59]
A posidoniid ostreoidan. It is the type fossil of the Sachrang Formation. Originally it was named "Posidonia bronni", thought to be a new genus, and the strata was denominated the Posidonia layers after it. Years later it turned out to be a junior synonym of Bositra, and thus, it was reassigned. However, the name of the layers was retained. The habitat and mode of life of Bositra has been debated for more than a century. There have been different interpretations, such as a pseudoplanktonic organism,[61] a benthic organism related to open marine floor, where it was the main inhabitant of the basinal settings, and a hybrid mode, where it has a life cycle with holopelagic reproduction controlled by the change on Oxygen levels, and even a chemosymbiotic lifestile, related to the large crinoid rafts, being the main "Safe conduct" to evade anoxic events. All the opinions along the years led to a large study in 1998, where the size/frequency distribution, the density of growth thanks to the lines related to the shell size and the position of the redox boundary by total organic carbon diagrams has revealed that Bositra probably had a benthic mode of life.[62]
A pectinoid scallop. The presence of this genus along endo- and epibenthic bivalves, which are absent farther up the section, suggest a delayed overstepping of anoxic bottom waters on the Altdorf High.[63]
An inoceramidclam. Being the second most common genera of Bivalve on the Formation, it had been object to several studies to find its ecological niche, like Bositra. Several opinions include a pseudoplanktonic-only organism, able to live in open sea, or a benthonic-only organism. On the 1998 evaluation with Bositra, was found that probably has a benthic early life that translated to a faculatively pseudoplanktonic mode of adult life.[62]
A Coelodiscidae sea Snail. Is the oldest known holoplanktonic gastropod, thanks to a bilateral symmetrical shells as an adaption to active swimming. Also the most common of the sea snails of the Formation, it is also one of the most varied in size terms, with some of the biggest specimens of snail from the Lower Toarcian know.[69] It has been related to large floating driftwood as one of the primary settlers.[69]
A Pterotracheidae sea Slug. Among the oldest pelagic floating Slugs, Pterotrachea liassica had a more extended larval period than modern extant Pterotrachea coronata, because one additional whorl is present.[70]
Ammonite internal moulds of uncertainf affinity. Represent the most diverse Ammonite remains found on all the formation and are usually not identificable to family level
A Megateuthididae Belemnite. Includes some of the Biggest Know Belemnites, with an estimated maximum up to 2 m long in life, although, most specimens would have been rather smaller.
A Belemnotheutidae Belemnite. Chitinobelus rostrum was composed of aragonite with organic material, while normal Belemnites had calcite. Has been suggested this rostrum was calcitic.
Complete Specimens with tissue, complete Arm Crown, ink Sacs and organ Traces
Isolated Gladius
A Diplobelidae Coleoidean. It has been found adult individuals of Clarkeiteuthis which caught small teleost fish of the species Leptolepis bronni. Further indirect evidence for the hunting behaviour comes from their body orientation in the water during life.[9]
Ammonite internal moulds of uncertainf affinity. Represent the most diverse Ammonite remains found on all the formation and are usually not identificable to family level
A Hildoceratidaeammonite. The co-occurrence on Altdorf of boreal (Pseudolioceras) and Tethyan faunal elements (Frechiella) is striking, suggesting clear connection with both regions.[88]
Complete Specimens with tissue, ink Sacs and organ Traces
Isolated Gladius
A possible primigenial Cuttlefish. Is one of the most important fossils of Cephalopods on the Sachrang Formation, due to be one of the Earliest examples of Pigments found on any species, also one of the first historically.[92] The pigments are preserved on various specimens with Eumelanin related to its ink sacs and include even phosphatized musculature.[10]
Partial Specimens with tissue Impressions and ink Sacs
Isolated Gladius
A Loligosepiidae Loligosepiidan (Vampyromorpha). Related to the modern Vampyroteuthis infernalis. Gladii of Loligosepia can be distinguished from Jeletzkyteuthis by the transition lateral field/hyperbolar zone. A specimen of this genus found on Ohmden has appeared predating a Parabelopeltis.[11] The association of this 2 genera shows the predatory behaviour of this group when lived on Epicontinental seas, being rather different than extant Vampyromorphs.[11]
Type member of the Lioteuthididae inside Vampyromorphida. The taxonomic position of Lioteuthis is uncertain, although the wings reaching the proximal gladius section and the smooth median field suggest affinity to the Prototeuthididae[94]
Complete Specimens with tissue, complete Arm Crown, ink Sacs and organ Traces
Isolated Gladius
A Loligosepiidae Loligosepiidan (Vampyromorpha).[96] The Loligosepiidae is believed to be ancestral to the Recent vampire squid, Vampyroteuthis infernalis.[89] Hooklets in the food residues in the posterior mantle indicate that Loligosepia preyed upon belemnites.[95]
Complete Specimens with tissue, complete Arm Crown, ink Sacs and organ Traces
Isolated Gladius
A Geopeltidae Loligosepiidan (Vampyromorpha). Related to the modern Vampyroteuthis infernalis. It is distinguished from Geoteuthis and Loligosepia by its median rib: this rib forms a narrow ridge between two narrow grooves. Probably hosted finds similar to modern Vampyroteuthis.[11]
A PhylloceratidaeAmmonite. The largest ammonite found in the Posidonienschiefer comes from the Ohmden quarry,and belongs to a Phylloceras heterophyllum with a diameter of 87 cm.[73]
The First Cycloid Arthropod from the Jurassic, from the family Halicynidae inside Cycloidea.[104] Cycloids are a group of maxillopod arthropods that span between the Paleozoic until the latest Cretaceous, probably related to the crustaceans and probably detritivores.[104]
A marine Ostracodan, member of the family Healdiidae inside Podocopida. Rather abundant on the Toarcian profiles on Europe, this genus has a Mussel-like shape, with a very clean and round morphology.
The specimen reported represents the oldest fossil record of an achelatan lobster larva, and the first representative of achelatan lobsters in the Posidonia Shale. Shares similarities with the late Jurassic genus Cancrinos. It is also the first example among crustaceans, which possibly could have lived as part of the plankton.[111]
California spiny lobsters are relatives of the Larval specimen from the Sachrang Formation
Type genus of the Erymidae Decapodan family. Originally, was named Glyphea amalthei,informally used by Quenstedt and housed on the Museum Naturkunde in Württemberg. A series of posterior revisions probe it was a different genus.[115]
A ColeiidaeDecapodan. Was confussed with Proeryon hartmanni specimens. Although these finds point to some potential for exceptional preservation, the Pfronten locality was never systematically sampled.[116] It was assigned to the genus Gabaleryon the basis of its ovoid carapace, a transverse groove cutting the axial carina on pleonal terga and an uropodal exopod with curved diaeresis.[116] It shares similarities with Gabaleryon garassinoi and Gabaleryon moorei.[116] A species from Gomaringen is the first know with preserved ommatidia.[117]
A Coleiidae Decapodan. The second largest Decapodan from the formation, P. giganteus is a species reaching a larger size than most other polychelidans, with up to 15 cm. On the Sachrang Formation there is the most abundant variety of species from the genus, ranging from different sizes and morphologies, that indicate different habitat & feeding adaptations on the genus. Some like P. hartmani show less adaptations to hunt for small nectobenthic preys than other relatives, being abundant on Oyster-filled waters. There is a relatively abundance of the genus in deep-water settings from the Toarcian onward.[123]
An Erymidae Decapodan. It was erroneously reported from the Late Toarcian, as was found on bituminous limestone on the Unterer Stein it is clear its provenance from the Sachrang Formation.
A gregarious PolychelidanLobster. Specimens of Tonneleryon schweigerti where recovered generally in cluster of several individuals, a characteristic unique to this species on the whole Polychelida group.[113] The specimens in these accumulations are of similar size, lacking characters of exuviae such as a split median line or disjunction of carapace and first pleonite. Due to that and the disposition of the specimens probably represent a mass-mortality assemblage and suggest this species was gregarious.[113]
An Astacidea Decapodan of the family Uncinidae. Reaching large sizes of almost half a meter (39–47 cm), Uncina Posidoniae is among the largest known Jurassic Crustaceans. Uncina posidoniae is also the largest representative of the genus Uncina.[125] This large crustacean has been found associated with Ammonite and Bivalve filled Bentos, where probably hunted different kinds of prey.[125] Its large claws would have been perfect to hunt small invertebrates and vertebrates.[125]
The Largest complete Uncina posidoniae specimen, with 44 cm long.
Numerous disarticulated individuals, associated with fossil wood.[15]
A phosphatic-shelled Cirripede of the family Eolepadidae.[15] Toarcolepas is provisionally interpreted as the oldest epiplanktonic cirripede known, and is thought to have lived attached to floating driftwood.[15]
Modern genus Lepas is among the most related taxa to Toarcolepas
The type genus of the family Liassoscorpionididae, probably related to Mesophonoidea. Being the only Jurassic scorpion known, there is no evidence that L. schmidti was aquatic (which was suggested in the past) and in the absence of further, better preserved material it should be excluded from future considerations of broad patterns of scorpion evolution.[127]
Insecta
Insects are a common terrestrial animals that were probably washed into the sea due to monsoon conditions present on the Sachrang Formation.[128]
Uncertain Affinity Insects, whose remains are scarce to allow an assignation to a concrete family. Also Includes insects with characters never seen in other families and genera.
A dragonfly of the family Heterophlebiidae. Heterophlebia is a relative abundant Genus, present on most of the pits of the Sachrang Formation, even on Holzmaden. Heterothemis is the second most distributed insect on the formation.
A dragonfly of the family Sphenophlebiidae. The Kerkhofen specimens of Sphenophlebia pommerana shows an almost complete dragonfly with head and abdomen as well the two pairs of wings that are partially on top of each other, consisting of fore and hind wings.[138]
A stick insect of the family Aerophasmidae. Schesslitziella is one of the described insects found more near the Bohemian Massif, where probably belong most of the terrestrial invertebrate fauna.
A Giant Lacewing (Kalligrammatidae) of the subfamily Liassopsychopinae and Kallihemerobiinae. It is one of the oldest known representatives of the Giant pollinator lacewings. The genus Liassopsychops was previously referred to Psychopsidae. Another specimen related, Specimen Hi 21 is regarded here as Kalligrammatidae incertae sedis. This along the occurrence of two distantly-related genera of Kalligrammatidae in the lower Toarcian is unexpected.[153] The Toarcian Kalligrammatidae lived in warm and dry conditions, as proven by its recovery from the zone of Hondelange but absence on Grimmen and other northern locations. Both genera where among the largest insects found on all the formation.[153]
Lance Lacewings of the family Osmylidae. Tetanoptilon is the largest non-Kalligrammatidae lacewing of the Jurassic, with a forewings length up to 470 mm and a wingspan estimated at 11 cm. Like the Kalligrammatids, this genus is limited to the southern zones, asbsent in northern deposits.[153]
Extant member of Osmylidae, Posidonia genera where probably similar
Beetles of uncertain placement inside Coleoptera. Diverse beetle specimens that due to its preservation (incomplete) or its morphological traits are not assigned to any concrete family. The abundance of beetle Elytrons indicates the proximity of terrestrial habitats.
An Amphiesmenopteran of the family Necrotauliidae. The ovipositor, like terminalia of female N. parvulus, indicate that these insects laid their eggs rather in soil than in water
Lepidopterans probably related with the family Micropterygidae. Compared with their record on Grimmen, on Lower Saxony Lepidopterans are rather scarce and bad preserved.
Extant member of Micropterygidae, Posidonia genera where probably similar
A primitive Crane fly of the family Tanyderidae. Extant members of the family are nectar feeder while extinct members cannot be determined precisely.[161]
Extant member of Tanyderidae, Posidonia genera where probably similar
Echinoderm debris is pretty abundant on the shale-free Unken and Salzburg members, including Crinoid skeleton elements, also that of the Ophiurida; the Echinoids take their place, where really blossomed at that time. That's why Pedicellaria are observed very often.[37]
A sea urchin of the family Cidaridae. Common on several layers. Cidaris is genus that still alive today. A bottom dweller, is commonly found associated with Belemnnite fossils, probably due to eating its carcasses.
A sea urchin of the family Pedinidae. It is the most common sea urchin in the formation, present on all the levels with specimens of various sizes. Vinculated to sea bottom sediments, before Gasteropods and Bivalves, Diademospsis was the third major colonizer of the bottom, in between anoxic changes.
A sea cucumber of the family Chiridotidae. It is the only major genus of Sea Cucumbers reported locally on the Posidonienschiefer. It was identified originally as Chirodonta mesoliasicus and Chirodonta heptalmorpha. Represents a possible cosmopolitan holothurian that occupied all three types of deep-sea ecosystems as an organic-enrichment opportunist.
Megaraft Colonies with +thousand associated specimens
Isolated Stems
Type genus of Crinoidean from the family Pentacrinitidae. Like Seirocrinus, Pentacrinites formed colonies on rafting wood, getting a different role than bigger crinoid and appearing on the first stages of the decomposition of the rafting wood. Was a smaller genus, with specimens of no more than 1 meter long, usually measuring 40–70 cm.
Close view of one specimen from the Posidonienschiefer
Exceptionally well preserved individual with the arms,pinnules and cirri largely Intact
A Crinoidean of the family Isocrinida. This Benthic Crinoid clearly represents an exotic elementement of the typical Posidonia fauna, likely moved from the coastal settings
Megaraft Colonies with +thousand associated specimens
Isolated Stemns
The largest known Crinoidean, from the family Pentacrinitidae. Among the tallest animals of its period, Seirocrinus is also one of the most famous fossils from the Sachrang Formation. It consists of fossils of colonies along large wood trunks, with specimens up to 14 m long, with the largest specimen reaching 26 m long, what makes it among the tallest know Mesozoic organisms, one of the largest invertebrates know on the fossil record and one of the tallest know animals.[171] It was an open ocean organism that lived in rafting woods, probably filtering food and serving as a refuge for other animals, such as ammonites. The crinoids had a large colonization process, based on the status of the fossil wood found.[172] The large rafts were the home for a high variety of marine organisms, such as Balanoideans, Ammonites and other. It has been estimated that without the presence of modern raft wood predators (that appeared on the Bathonian) those rafts can last up to 5 years, being that the main reason the crinoids were able to reach such huge sizes. The large rafts were also probably essential to distribute animals along the Early Jurassic Seas.[13]
Close view of one specimen from the Posidonienschiefer
A member of Callorhynchidae inside Chimaeriformes. Similar to Callorhinchus, among the oldest known of its type. It the first Modern lineage Chimaeras from the Toarcian.
Meckelian Cartilages, Jaws, teeth, Palatoquadrates, placoid scales and dearticualted parts of the labial, hyoid and branchial skeleton.
A shark of the family Hybodontidae. The Type specimen belongs to a large hybodontid, with an estimated total length of up to 3 m.[176] It has a Meckelian Cartilage more robust than Hybodus hauffianus and an acrodontine dentition, as seen on Bdellodus.[176] Probably is related with "Hybodus" delabechei, very likely a junior synonym of Crassodus.[176]
Articulated cranial fragment preserving the dentition, lower jaw, cephalic spines, palatoquadrate
SMNS 10062 - a complete skeleton with belemnites in the gastric mass
SMNS 15150 - a complete skeleton with the dentition
SMNS 80594 - (in three parts) a small cranial fragment with at least 12 associated teeth
SMNS 54048 - a disarticulated anterior part of a skeleton with well preserved teeth
SMNS 51949 - a virtually complete specimen
SMNS 52460 - a virtually complete skeleton with dentition
SMNS 51136 - a partial anterior skeleton with a single tooth exposed in labial view
Unnumbered Partial skull and articulated dentition
BMNH P.5880 - a disarticulated anterior part of a skeleton
SMNS 10062 - isolated parts of a skull
SMNS 8663 - a partial skeleton
SMNS 80595 - dorsal fin spine and scapulocoracoids plus associated skin and other cartilages
SMNS 51 135 - the partial skeleton of a juvenile
SMNS 58749 - virtually complete skeleton
SMNS 58748 - a fragment of a hyomandibula
SMNS 52222 - a partial skeleton with both dorsal fin spines
SMNS 50804 - a partial skeleton with both dorsal fin spines
Unnumbered Complete wall mounted specimen in left lateral view
Isolated Teeth
Isolated Fin spines
Type genus of the family Hybodontidae. It is the most abundant shark on the layers of the Sachrang Formation, with some of the best preserved specimens of the genus known. It was probably an open ocean hunter, with small horns over the eyes. With a size around 2 m, it was also one of the largest representatives of the Chondrichthyes on the formation.[179]
Hybodus Holzmaden specimen, among the best preserved of the genus, with Belemmnites inside.
Anterior part of body with basicranium, palatoquadrates, Meckel's cartilage, ceratohyals, epihyals, teeth, traces of the branchial arches and the anterior finspine
A possible member of Batoidea. It was originally identified as a member of Galeiformes. This Genus was found to be sister taxa to the extant genus Raja, yet maybe represents an stem Rajiform. It is probably the oldest described pelagic Rajiform. It wears an enlogated rostrum.
A deep-bodied neopterygian, the type genus of the family Dapediidae. Unpublished material indicates the presence of one or even two more still undescribed species of Dapedium in the Lower Toarcian.[189] A specimen of Lytoceras has a Dapedium sp. in the body chamber probably trapped when feeding on the carcass of the cephalopod, and may therefore indicate an opportunistic carrion feeder.[190]
First ganoin-scaled Ophiopsiformes (Halecomorphi) from the Posidonienschiefer. The type specimen measures 51 cm, and has elongated and serrated body scales before the dorsal fin and tiny ganoid scales after it.[194]
A member of the family Leptolepididae. The most common member of its family, Leptolepis is commonly associated with crustaceans and small marine invertebrates, probably the main constituents of its diet. One on the most predated vertebrates on the formation, with abundance of larger fishes and reptiles with specimens associated.
A member of the family Leptolepididae. Was identified as Paraleptolepis, but this name is currently occupied by a Japanese fish genus of Early Cretaceous age.[200] It differs from Leptolepis coryphaenoides in the presence of a few autapomorphies and also in the retention of several primitive features not present on the last one.[199] Small genus, of about 14 cm length.[199]
First ganoin-scaled Ophiopsiformes (Halecomorphi) from the Posidonienschiefer. Elongated morphology, with a length of ~39 cm, covered by smooth, massive ganoin scales.[194]
Type member of the family Pachycormidae. Large representative of its family, with a size up to 1.5 m. One specimen preserved the alimentary canal, with the stomach filled by numerous hooklets that can be referred to the coleoid cephalopod Phragmoteuthis, implying a diet of cephalopods for this genus.[18]
A member of the family Pholidophoridae. Is among the most abundant fishes on the late liassic of Europe, present on the sub-Mediterranean boreal, with specimens of several sizes.
A large member of the Chondrosteidae and the largest non-reptilian marine vertebrate of the Sachrang Formation, with a size between 3 and 4.5 m, and an estimated weight over 800 kg to 1 tonne.[208]
A large coelacanth of the family Mawsoniidae, related to the genera Axelrodichthys, Chinlea, Diplurus and the type, Mawsonia.[211] The largest specimen known from the Sachrang Formation is GPIT.OS.770 (Holotype), with a length over 1.6 m.[210] The specimen presents an ossified lung inside the abdominal cavity, and most of the body, being also one of the most complete coelacanths of the Jurassic found.[210][211]Trachymetopon precedes the presence of the family Mawsoniidae in Europe by about 120 Ma and the northernmost occurrence of a member of the group, implying an extensive geographical range during the Early Jurassic.[211] Due to the specimens being found on pelagic deposits suggest that probably was an open ocean swimmer.[210][211]
A large ichthyosaur of the family Leptonectidae with convergent evolution with modern swordfish. Like these fishes, Eurhinosaurus is believed to be a fast swimming predator, able to hunt fish schools on same way. Large specimens of up to 6 m are known.
Small sized ichthyosaur, probably a member of Parvipelvia, sister group to Stenopterygius + Ophthalmosauridae. A small- to mid-sized ichthyosaur, 2–3 m in length, with a relatively short and slender antorbital rostrum.[215]
A possible member of the family Leptonectidae. Mostly of the specimens of this genus have been referred to Leptonectes or Temnodontosaurus, although some remains on the Posidonienschiefer are too complex to being clearly referred.
An Ichthyosaur of the family Stenopterygiidae. Magnipterygius may not have grown to a total length of much more than 120 cm. It is therefore potentially only the second post-Triassic ichthyosaur known with such a small body size
Type genus of the family Stenopterygiidae. A common Toarcian ichthyosaur, present in multiple layers. The rather exquisite level of preservation has led to know even the coloration, that exposes a clear countershading, with an upper part being more obscure than the lower, similar to modern killer whales, the Heaviside's dolphin or the Dall's porpoise. There is also evidence of changes in color with ontogenic changes, going from dark juveniles to countershaded adults. The skin was flexible & scaleless, as in dolphins.[219] The study of several specimens has revelated that Stenopterygius quadriscissus underwent a size-related trophic niche shift through ontogeny, shifting from a piscivorous diet to a teuthophagous diet, known thanks to exquisitely preserved stomach contents.[220]
Type genus of the family Temnodontosauridae. A large macroraptorial ichthyosaur, apex predator of its environment. It ranges between 9 and the 12 m, being one of the largest known ichthyosaurs, characterized by skulls and jaws over 1 m in length, with the largest being over 1.9 m long. It has been found with fragments of young icthyosaur in his stomach.[17]
A basal member of Pliosauridae. A moderately sized (3.4 m) Pliosaurid, ecologically adapted to fish hunt, as has been observed due to comparing the long snouted skull with that of Peloneustes, Gharial Crocodiles or Dolphins.[226]
Type member of the Plesiosaur family Microcleididae. Small Plesiosaur, with a length of less than three metres. It is characterised by a really elongated neck, and was probably an ichthyophagous form that occurred rarely in the Posidonienschiefer fauna.
SMNS 51945, an almost complete articulated skeleton lacking most of the skull
Skin Impressions
Phosphatised collagen fibres
An indeterminate plesiosauroid plesiosaur. An impressively preserved immature specimen, different from Hydrorion brachypterygius and Seeleyosaurus guilelmiimperatoris, the most abundant plesiosaurs found locally, and its anatomical characters suggest it represents a new genus.[228] It fossilized buff-coloured material, identified as mainly composed of calcium phosphate and interpreted as phosphatised muscle tissues.[228] It also contains eumelanin and hence possibly corresponds to areas dark-coloured in life.[228] The stomach contains quartz grains that were most likely ingested during the animal's life, maybe used for food trituration. This gastroliths have importance for its exotic provenance compared with the surrounding lithology of the deposits (mostly shale), and as sandy turbidites have never been reported from the SW German Basin, the individual may have acquired the sand-sized grains many kilometres away from the burial site.[228] Strata containing fine sand ('Glaukonit und viel Feinsand') at Obereggenen im Breisgau (western side of the Black Forest between Freiburg and Basel), 200 km from Holzmaden suggest a nearshore deposit was allocated here, and that the black forest emerged at this time. Probably this young specimen reached that location in search for gastroliths.[228]
A basal Plesiosaur that has been linked with Cryptoclididae. It is one of the smallest from the Posidonia, with a complete skeleton measuring less than 2.5 m. It is considered a possible junior synonym of Seeleyosaurus.
A Plesiosaur of the family Microcleididae. It was named originally "Plesiosaurus guilelmiimperatoris". It was a moderate-sized plesiosauroid, measuring up to 3,5 m in length with a skull length of 170 mm.
An aquatic sphenodont of the family Pleurosauridae. Palaeopleurosaurus evidences that there was a slightly skeletal specialization for an aquatic lifestyle, achieved through the Jurassic gradually on pleurosaurs.[234] Recent studies suggest a shorter lifespan than modern Tuatara, based on irregular spacing of growth marks.[235]
A marine turtle of the superfamily Eurysternidae inside Thalassochelydia. Is the main formally identified fossil as Turtle from the Sachrang Formation, representing a rather basal genus. The pleurals resemble those of the genus Plesiochelys.[236] Being found in the zone of Franconia that on the Toarcian was at -80 km from the shore can suggest that early marine turtles lived in the epicontinental waters of the European shallow seas before reach richer ecosystem diversity on the Late Jurassic.[236] This would explain the serious lack of Turtle fossils on the formation, as mostly of the deposits are located far from the coast.[236]
Possible unclassified testudine remains. Münster (1834) cited: "there were also rare things at the quarries of Altdorf, among other remains there were ones of a turtle on lias limestone"". The remains are not catalogued and some specimens are in Private Collections.[239]
A marine turtle of the clade Thalassochelydia inside Angolachelonia. The only testudine identified from the northern realm of the Sachrang Formation.[240]
A longirostrine thalattosuchian of the family Machimosauridae. Was considered synonymous with Steneosaurus until in 2020 this last was recovered as invalid. It reached large sizes, with specimens exceeding 5 m, being a generalist predator.[241]
A mesorostrine thalattosuchian of the family Teleosauridae. A marine crocodylomorph with a diet probably based on fish. Was considered synonymous with Steneosaurus until recently.[244] Due to this unusual placement of the external nares, Mystriosaurus was more terrestrial, or spent a greater amount of time on land, than other teleosauroids. This would explain its greater presence in zones of the formation more proximal to the emerged landmasses. Its morphology suggest it was a mesorostrine generalist.[244]
A thalattosuchian with a complex assignation, probably the basalmost metriorhynchoid. Pelagosaurus typus was a small-bodied thalattosuchian (~1 m in length) considered to be an adept aquatic pursuit predator, with a long streamlined snout ideal for snapping at fast moving prey (one specimen was found with Leptolepis fishes inside) and large, anterolaterally placed orbits for increased visual acuity.[245]
A Longirostrine Thalattosuchian, the most basal know. Was considered synonymous with Steneosaurus. Longirostrine specialist, probably active fish hunter.
A longirostrine thalattosuchian of the family Teleosauridae. Platysuchus was slightly more robust than its contemporaneous relatives, being probably adapted to hunt more voluminous fish. A heavily armoured, semi-terrestrial longirostrine generalist form, indicated by the extensive and tightly packed rows of dorsal osteoderms.[241]
A Novialoidean Pterosaur, type genus of the family Campylognathoidea. Mark Witton suggests the construction of Campylognathoides' extremely robust forelimbs, with proportionally long wing fingers, could be a specialization for a fast aerial lifestyle comparable to those of Falcons and mastiff bats, being more probably an insect & vertebrate hunter and living on nearshore environments.
A Rhamphorhynchinae Pterosaur. It is one of the best known Early Jurassic Pterosaurs.[248] Unlike Campylognthoides, Dorygnathus was an oceanic hunter, with teeth disposed to catch marine prey, such as Belemnittes and several species of fishes. Dorygnathus mistelgauensis is considered a junior synonym until more data can be recovered from the specimen, held on a private collection.[248]
A Novialoidean Pterosaur, probably a member of the family Campylognathoidea. Has been assigned to the genus Campylognathoides, although it is clearly different than any other pterosaur from the Sachrang Formation.[91] The name "Schandelopterus" is invalid and lacks any study, assigned without species to refer to the specimen on private German Fossil Groups. The pelvis indicates a laterally, slightly upwardly directed orientation of the acetabula which does not support a bird-like bipedal locomotion of this pterosaur as has been suggested on the past.[91]
A Rhamphorhynchinae Pterosaur. Has been assigned to the genus Dorygnathus. It has a really complete skull that can help to explain the status of the genus Parapsicephalus.[253]
An indeterminate possible Sauropodomorph dinosaur, possibly a member of Sauropodiformes inside Anchisauria (Resemble Yunnanosaurus teeth).[255] The Fossil was reported from the Lias Epsilon level, that on Oedhof is occupied by the Sachrang Formation.[255] It was found with abundant Plant debris and Belemnite remains.[255] Hasn't been revised since 1956
The Oedhof jaw can belong to a creature similar to Yunnanosaurus
Plantae
The Flora is dominated by horsetails, what can be due to a similar ecological niche to modern genus Phragmites, able to resist saline conditions. Storms or floods maybe where the major events that transported this flora to the sea.
The macroflora of the Posidonia slate can be described as extremely poor in species.[256] Apart from the remains of Horsetails, it is without exception the remains of coarse branches and fronds from gymnosperms, in which one has a certain can assume transport resistance. Remains of Ferns are completely missing, except for tall arboreal ferns (Peltaspermales).[257] Mostly of the flora was reported from the area of Braunschweig.[256] The major explanation for the flora could be that the plants in question are mono-or oligotypic stands on the edge of the waters that flow into the Posidonienschiefer sea, probably tear away in the course of flood events, easily fragmented during transport and wave waves, possibly especially in the occasional storm events postulated.[258] In terms of taphonomy, this would result in a comparison with today's reed Phragmites, which can form extensive stocks on the edge of shallower and slowly flowing waters ("Reed belts").[256] The Wood remnants clearly indicate one higher diversity of Coniferous flora in the delivery area than the remains of leafy branches.[256] This fact is likely to be proportionate, similar to that frequent occurrence of charcoalized or gagged trunks, mostly of them are believed to be "driftwoods" that only take a long time drifting also suggests a frequent settlement with mussels and full-grown Sea Lilies.[256][258] The deposition settings are at large distance from the nearest coastline (for southern Germany about 100 kilometers), making only plants strong to transportation able to resist enough to get deposited.[259][142] At Irlbach and Kheleim, NE of Regensburg, where the Posidonienschiefer has its near mainland deposit with abundant sand, a rich deposit filled with plant remains of different kind (Seds, Reproductive organs, Leafs, Stems, Cuticles and wood) with traces of coal was recovered, however, it was never studied in depth.[80] Of all the plant material expected only a few Bennetites leafs and two conifer branches with leaves where cited and none studied.[80]
At the Austrian realm The sachrang Member was developed in the basinal area, while the Unken Member, sandwiched between red, often condensed limestones, represents the marginal facies.[37] Due to be more marginal and connected with the southern Vindelician land, the most diverse palynological assemblages of the formation are found, transported from zonas with moldanuvian granites as proven by the feldspar accumulations.[37]
Phytoclasts
Phytoclasts have been recovered from several sections on the formation, but only studied in depth from the Dotternhausen and specially Dormettingen.[43] Here two kinds of Phytoclasts where recovered, opaque phytoclasts (charcoal, indicator of wildfire activity on nearby landmasses, indicator of seasonal alterations of the water column) and translucent phytoclasts (indicator of proximal landmasses with high availability of wood and other plant material, as well transport conditions).[43] On the lowermost part of the section opaque phytoclasts are low (15% of the total organic matter) while translucent are incredibly abundant (40%), lowering its abundance to a 20-10% on the next section.[43] The Exaratum Subzone is the only one with an inverse trend and more abundance of opaque phytoclasts. On the Bifrons level, both types reach between a 15% and a 30%, showing a rapid increase, to decrease on the end of the section to values of less than 10%.[43] Opaque Phytoclasts, for a supposed marine deposit are relatively abundant on some sections, whose decreasing on others suggest (along with increasing levels of Kaolinite) an increased delivery of land plant material by rivers, from areas with wetter climate and less frequent fires, while its rise suggest the opposite, nearby continental setting with dry climate and continuous wildfire activity.[43]
Affinities with the families Peltaspermaceae, Corystospermaceae or Umkomasiaceae inside Peltaspermales. Pollen of Uncertain provenance, that can be derived from any of the members of the Peltaspermales. The lack o distinctive characters and bad conservation are among the main factors to make this Palynological residues difficult to classify. Arboreal to arbustive seed ferns.
Affinities with both Sciadopityaceae and Miroviaceae inside Pinopsida. This Pollen resemblance with extant Sciadopitys suggest that Miroviaceae can be an extinct lineage of sciadopityaceaous-like plants.[263]
Extant Sciadopitys. Cerebropollenites likely come from a related plant
Affinities with Cycadaceae and probably Cycadales. Alternatively can be Pollen from Bennettitales. It is the most abundant non conifer Pollen recovered on the formation, recovered on all the major sampled areas. Probably derived from arbustive cycads, this genus is related with dry settings, even from desertic regions.
Closer Look of Macrozamia cones, a common Cycad. Chasmatosporites maybe come from a related plant
Affinities with Cheirolepidiaceae inside Coniferae. Abundant on the Lower Jurassic of North and Southern Europe, represents pollen of medium to large arboreal plants, specially coniferales. The abundance of pollen of Classopollis and other thermophile plants was observed in this region in the lower Toarcian from the end of the antiquum (= tenuicostatum) zone to the middle of commune zone.[265]Classopollis is correlated with evaporites and are therefore associated with desert basins, but the shrubs may have also lived in xeric upland areas with seasonal fires. Evidence of fires is absent on the marine Posidonienschiefer, but has been recovered on the coeval nearshore calcareous sandstones.[265] It increases with the appearance of charcoal phytoclasts, as derived from dry settings with increased wilfires.[43]
Affinities with Gnetopsida and probably Gnetophyta. Has Been considered Pollen of Chloranthaceae. However, it is to old for belonging to advanced Angiosperms. It probably comes from cones related to the Genera Piroconites kuesperti from the Lowermost Jurassic of Germany, resembling pollen of extant Ephedra and Welwitschia.
Closer Look of Ephedra cones, a common Gnetal. Clavatipollenites maybe come from a related plant
Affinities with Gleicheniaceae inside Gleicheniales. Suggest relative increase of humidity on the rivers flowing towards the Austrian realm. Most abundant Fern spore in this region.
Dicranopteris, Concavisporites come probably from similar genera
Affinities with the Cycadopsida inside Cycadales. Pollen related with modern Cycas, arbustive to lower floor plants, relatively abundant, present on various of the measured samples. The mos common found on the Austrian realm, indicator of dry settings.
Encephalartos, Cycadopites come probably from similar genera
Affinities with Selaginellaceae and probably Lycopsida. A rare element on the palynological records of the German Basin, although more abundant than any other Spore recovered locally.
Affinities with the Pinidae inside Coniferae. Abundant on the Lower Jurassic of NW Europe. Its identification on the Posidonienschiefer is rather complex due to the bad preservation of the Pollen Grains.
Extant Pinus cembra Cone, example of the Pinidae. Inaperturopollenites is similar to the pollen found on this genus
Affinities with the Ophioglossaceae inside Filicopsida. Spores related with modern floor Ferns, that appear on abundant water locations. The Unken Member is considered a more basinal deposit, where Wood and Sporomorph remains are more common.
Ophioglossum, Lycopodiacidites come probably from similar genera
Affinities with Cheirolepidiaceae inside Pinaceae. Abundant on the Lower Jurassic of NW Europe. Spheripollenites co-occurs on the coeval Sorthat Formation with cuticles of Dactyletrophyllum ramonensis, and after a test of relationships it was found a highly significant correlation that may suggest that the species S. psilatus was produced by the conifer genus Dactyletrophyllum.[267]
Affinities with Equisetaceae inside Equisetopsida. Number of mostly very fragmented and not particularly well preserved, but clear horsetail remains described. So far recognizable, leaf sheaths where developed in most cases, but the state of preservation does not allow a more precise determination.[268][269]
Affinities with Equisetaceae inside Equisetopsida. Neocalamites is the most common more distributed of all the Posidonia Shale, being even found on Luxembourg Posidonia Strata.[270] Mostly of the Stems reported come from Aeolian-Dunar related deposits, or from nearshore-basinal deposition. Probably was related to the seashore.[270] Some stems are big, resembling the rates of growth seen on modern Bamboo specimens, suggesting +6–7 m tall Equisetopsids.[270]
Affinities with Umkomasiaceae inside Corystospermaceae. Is based on bipinnate leaves, rachis longitudinally striated, with a long petiole and secondary rachises. It belongs to large tree ferns. Sachrang Formation Specimen is characterized for its large size and probably where attached to trunks similar in built to the Cretaceous genus Tempskya.[271]
The only Pachypteris specimen know from the Sachrang Formation
Affinities with Cycadeoidaceae inside Bennettitales. It is the most abundant medium-sized plant on the environment. Found specially on seashore depositional settings, but also on deltaic and lagoonar environments. It was a low arbustive-arboreal Bennetite related to arid environments, with a leave similar of that of the modern genus Encephalartos, specially Encephalartos munchii, but also Dioon mejiae. Otozamites has been considered synonym with Otopteris, but since the 1990s everybody used the name Otozamites, and Otopteris was forgotten.[274]
Otozamites gracilis specimen from the Sachrang Formation
Affinities with Cycadeoidaceae inside Bennettitales. This Bennettitalean is related to Shrub built. Some specimens were assigned to Dioonites acutifolium (Junior synonym). Consists on leaves with pinnate, leaflets perpendicular or oblique to the rhachis, on the top of the leaf axis. It comes from fragments of fairly large fronds. In its external form it closely follows the modern genus Dioon, only the leaflets are of our kind wider and shorter, they are further apart, but the tendency to rhachis is the same.
Affinities with Williamsoniaceae inside Bennettitales. Arboreal Cycadaceans, some with the presence of flower-like structures. It resembles the leaf of the modern Microcycas calocoma, and probably had a similar arboreal built, being the leave of tall Bennetite trees such as Bucklandia, found on the middle jurassic of England.
Ptilophyllum sp. specimen from the Sachrang Formation
A member of Williamsoniaceae inside Bennettitales. It has been interpreted as a cycad in the family Cycadaceae or a Bennettitalean plant, and also a late surviving member of Noeggerathiales. Leaflets somewhat removed, oval-oblong, a little narrower near the base, rounded at the tip, nerves partially diverging from the base towards the edge. It was assigned to Pterophyllum oblongifolium and on the genus Glossozamites. This genus was the leaf of arboreal Bennetites, similar in appearance to the modern Encephalartos woodii.
Affinities with Ginkgoaceae inside Ginkgoales. Various leaves are known from Ohmden, coming from marine deposits and where identified as Ginko digitata, identified originally as the only Ginko specimen reported on the Sachrang Formation. In the southern and northern Germany, there are regular remains of coal, which are initially reminiscent of small Ginkgo leaves. The leaves are hard to identify, more or less regularly concentric structures, as they sometimes appear like the coarse fruiting bodies of wood-dwelling fungi, such as the genus Trametes.
Affinities with Araucariaceae or Cheirolepidiaceae inside Pinales. Specimens whose spiral foliage of the branch, in which the individual leaves open about 2/3 of their length (without the tip) are fused with the branch they hold in their free part fits tightly, suggests belonging to the genus Brachyphyllum.
Affinities with Cheirolepidiaceae. Is the type genus of this family, and is related with Arid Settings. Coming from The land-plant-taphocoenose from posidonia slate 3, is ready for processing.
Fragmentary axis compressions with preserved leaves
Cuticles
Isolated Leave Fragments
Affinities with Araucariaceae or Cheirolepidiaceae inside Pinales. Pagiophyllum araucarinum predominates among the two types of leafy coniferous branches that have become known from the Posidonia. However, there is no indication whether this fact reflects their respective share in the vegetation of the delivery area. Cheirolepidiaceae Pollen is the most abundant and diverse found on the formation, what is correlated with the abundance of this genus. Other factor that puts local Pagiophyllum on Cheirolepidiaceae is the dominance of an arid climate, the preferred for this type of conifers.
Fragmentary axis compressions with preserved leaves
A possible ancestral member of the Callitroideae inside Cupressaceae, or a member of Cheirolepidiaceae. Named also "Cupressites" liasinus, represents probably Arbustive to arboreal-derived axis.
Widdringtonites liasinus specimen from the Sachrang Formation
Fossil wood
Fossil Wood increases on the marginal Unken Member, with great amounts of logs and fragments of more than 1 m. Surface studies suggest relationships with the wood genera identified on the coeval Úrkút Manganese Ore Formation.[276]
Affinities with Araucariaceae inside Pinales. Has araucarioid structures, but lacks proper characters of extant genera such as Araucaria & Agathis The largest known rafting wood on the fossil record is assigned to this genus, with a length of 18 m. The rafts were populated with Crinoid colonies, and a wide variety of organisms.[14]
Affinities with Abietinae inside Pinales. This genus was found to be related with Protocedroxylon found on the coeval Whitby Mudstone, crucial to the interpretation of the fossil record of Early Abietinae.
Abies specimen, whose role was probably covered by the makers of the Araucariopytis woods.
Affinities with Podocarpaceae inside Pinales. Acmopyle pancheri is the only living conifer that has real resemblance with the wood assigned to Circoporoxylon. Xenoxylon phyllocladoides and C. grandiporosum are commonly found associated, for example in Braunschweig area.
Affinities with Cupressaceae inside Pinales. Wood with a morphology similar to the modern genus Metasequoia. Probably related to Widdringtonites liasinus.
Metasequoia specimen, whose role was probably covered by the makers of the Cupressinoxylon woods.
"Georg-Friedrich" mine not far from Liebenberg near Grauhof in the Harz foreland
Wenzen between Einbeck and Alfeld
Mistelgau
Altdorf
Fossil Wood
Affinities with Podocarpaceae inside Pinales. resembling modern genera such as Dacrycarpus, with other specimens resembling Juniperus. Includes wood more related to nearshore arbustive Conifers (columnar or low-spreading Shrubs with long, trailing branches), being the most abundant, but also medium to large arboreal conifers from nearshore forests.
Affinities with Cheirolepidiaceae inside Pinales. Includes large sized trunks up to 1.7 m tall and 115 cm wide. Wood of other species of this genus, such as Protocupressinoxylon purbeckensis grown on flood-influenced settings of vegetated soil with brackish-water, like modern Swamp Cypress.
Wood of the Protopinaceae, a possible "morpho-group" of the family Cheirolepidiaceae. Belongs to an invalid group of mostly Paleozoic Woods, and all the woods assigned to the family should be treated as Incertae Sedis.[281]
Affinities with the Cupressaceae inside Pinales. Probably related to the coastal settings. It maybe covered a similar role that modern Swamp Cypress, Taxodium distichum.
Taxodium distichum specimen, whose role was probably covered by the makers of the Taxodioxylon woods.
Dörnten not far from Liebenburg, district of Goslar
Fossil wood
Affinities with Coniferales, concretely is closer to the Podocarpaceae, Cupressaceae and in a lesser extend to the Cheirolepidiaceae. Finally can be a member of the extinct family Miroviaceae. It is the more abundant genus of wood present on the Bohemian Realm of the Sachrang Formation.
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