Cranial Anatomy of the Aetosaur Paratypothorax andressorum

Schoch, R. R., and J. B. Desojo. 2016. Cranial anatomy of the aetosaur Paratypothorax andressorum Long & Ballew, 1985, from the Upper Triassic of Germany and its bearing on aetosaur phylogeny. Neues Jahrbuch für Geologie und Paläontologie - Abhandlungen 279(1):73-95. DOI: http://dx.doi.org/10.1127/njgpa/2016/0542
http://www.ingentaconnect.com/content/schweiz/njbgeol/2016/00000279/00000001/art00008


Abstract - The large aetosaur Paratypothorax andressorum has so far been known only by its osteoderms. Here we describe for the first time the skull of a complete, articulated specimen of this taxon that was found in the type horizon at Murrhardt, southwestern Germany. Paratypothorax
andressorum has the following cranial autapomorphies: (1) upper jaw margin with deep notch between premaxilla and maxilla, (2) maxilla-lacrimal suture with finger-like projection, (3) upper
temporal fenestra triangular, and (4) first paramedian cervical osteoderms narrow and oval, much smaller than second row. Apart from these features, the skull of P. andressorum closely resembles that of the small aetosaur Aetosaurus ferratus known from the same horizons, despite major differences in the morphology of osteoderms. Both taxa share (1) the pointed, beak-shaped premaxilla which expands only gently anterior to the nasal, (2) maxilla and lacrimal excluding jugal
from margin of antorbital fenestra, (3) exclusion of squamosal from margin of infratemporal fenestra, and (4) posterior part of jugal not downturned. Phylogenetic analysis reveals poorly resolved relationships within Aetosauria, but exclusion of a problematic taxon Coahomasuchus results in a much better resolution, with Paratypothorax to nest with Rioarribasuchus, Tecovasuchus, Typothorax, and Redondasuchus within a monophyletic Typothoracinae. Interestingly, Aetosaurus and Stenomyti form successive sister taxa of this clade rather than fall within an aetosaurine grade of basal aetosaurs, as suggested by previous authors. The resemblance of Paratypothorax and Aetosaurus in many cranial features, their close relationship as suggested by the present analysis, and the immature state of all available Aetosaurus specimens suggest two new alternative hypotheses: (1) Aetosaurus is the juvenile of a close relative of Paratypothorax or (2) it is itself the juvenile of Paratypothorax.

Reanalysis of the Diapsid Reptile Elachistosuchus huenei from the Late Triassic of Germany

Sobral, G., Sues, H.-D., and J. Müller. 2015. Anatomy of the enigmatic reptile Elachistosuchus huenei Janensch, 1949 (Reptilia: Diapsida) from the Upper Triassic of Germany and its relevance for the origin of Sauria. PLoS ONE 10(9): e0135114. doi:10.1371/journal.pone.0135114

Abstract - The holotype and only known specimen of the enigmatic small reptile Elachistosuchus huenei Janensch, 1949 from the Upper Triassic (Norian) Arnstadt Formation of Saxony-Anhalt (Germany) is redescribed using μCT scans of the material. This re-examination revealed new information on the morphology of this taxon, including previously unknown parts of the skeleton such as the palate, braincase, and shoulder girdle. Elachistosuchus is diagnosed especially by the presence of the posterolateral process of the frontal, the extension of the maxillary tooth row to the posterior margin of the orbit, the free posterior process of the jugal, and the notched anterior margin of the interclavicle. Phylogenetic analyses using two recently published character-taxon matrices recovered conflicting results for the phylogenetic position of Elachistosuchus–either as an archosauromorph, as a lepidosauromorph or as a more basal, non-saurian diapsid. These different placements highlight the need of a thorough revision of critical taxa and new character sets used for inferring neodiapsid relationships.

Redescription of "Paleorhinus" (Phytosauria) Specimens from Germany

Butler, R. J., Rauhut, O. W. M., Stocker, M. R., and R. Bronowicz. 2013. Redescription of the phytosaurs Paleorhinus (‘Francosuchus’) angustifrons and Ebrachosuchus neukami from Germany, with implications for Late Triassic biochronology. Zoological Journal of the Linnean Society Early View. DOI: 10.1111/zoj.12094

 
Abstract - Phytosaurs are a diverse and morphologically distinctive clade of superficially crocodile-like archosauriforms that had a near global distribution during the Late Triassic. Because their remains are among the most abundant vertebrate remains recovered in many Upper Triassic terrestrial formations, phytosaurs are used extensively in long-range biochronological and biostratigraphic correlations. The biochronologically oldest and earliest branching known phytosaurs include an array of nominal species from the early Late Triassic of the United States, Germany, Poland, Morocco, and India that have been synonymized within the genus Paleorhinus, and subsequently used to define a global ‘Paleorhinus biochron’. However, recent phylogenetic work suggested that the North American species previously referred to Paleorhinus are paraphyletic. Here, we reassess the systematics and anatomy of putative specimens of Paleorhinus from southern Germany. Two well-preserved basal phytosaur skulls from the Blasensandstein (Carnian) of Bavaria form the holotypes of Francosuchus angustifrons and Ebrachosuchus neukami, both of which were synonymized with Paleorhinus by previous workers. We demonstrate that Francosuchus angustifrons shares unique synapomorphies with specimens referred to Paleorhinus bransoni from the Late Triassic of Texas, and thus refer the species to Paleorhinus. By contrast, the longirostrine Ebrachosuchus is highly distinctive in morphology, and our new cladistic analysis of Phytosauria demonstrates that it represents a valid taxon that is more closely related to Phytosauridae than to Paleorhinus. We provide the first autapomorphy-based support for a monophyletic but restricted Paleorhinus (supported by a nodal row on the jugal, and low paired ridges on the squamosal) and confirm that previous broader conceptions of Paleorhinus are likely to be paraphyletic.

Paleoenvironmental Interpretation of a Fossil Locality from the Late Triassic of Germany

Havlik, P., Aiglstorfer, M., Atfy, Haytham, E. and D. Uhl, Dieter. 2013. A peculiar bonebed from the Norian Stubensandstein (Löwenstein Formation, Late Triassic) of southern Germany and its palaeoenvironmental interpretation. Neues Jahrbuch für Geologie und Paläontologie, Abhandlungen 269 : 321-337
DOI: http://dx.doi.org/10.1127/0077-7749/2013/0354

Abstract - Despite the abundance of Triassic outcrops in S Germany knowledge about continental ecosystems from the Norian is rather scarce so far for this region. A new fossil-bearing site from the Lower Löwenstein Formation (Unterer Stubensandstein; Norian) of north-eastern Baden-Württemberg (SW Germany), discovered in 2011, yielded disarticulated vertebrate remains together with a moderately diverse palynoflora, ichnofossils and gymnospermous charcoal. These remains are accumulated in a bonebed layer. Palynomorphs support a Late Triassic age of the source sediment, showing a high diversity of conifers. The new site provides the second conclusive evidence for wildfires in the Norian of Europe and the first record of phytosaurs from Schwäbisch Gmünd. Based on sedimentological evidence and taphonomical interpretations of the charcoal remains, it seems possible that the bonebed can be regarded as the result of increased erosion following catastrophic wildfire.

First record of Colognathus (?Amniota) from the Middle Triassic of Europe

This is a short note redescribing material from the Middle Triassic of Germany that is referable to the enigmatic vertebrate taxon Colognathus, which has been considered everything from a 'fish' to and archosauromorph reptile. Unfortunately the new material does not clarify the systematic position of Colognathus, but does represent a temporal and geographic extension.

Sues, H. -D.,  and R. R. Schoch. 2013. First record of Colognathus (?Amniota) from the Middle Triassic of Europe, Journal of Vertebrate Paleontology, 33:4, 998-1002. http://dx.doi.org/10.1080/02724634.2013.732977

A new interpretation of material of an armored archosauriform from the Middle Triassic of Germany that solves the mystery of the purported Middle Triassic aetosaur.  Instead this specimen is from a taxon more closely to Doswellia kaltenbachi. This new taxon supports the growing realization that doswelliids had a broader range geographically and temporally.

Schoch, R. R., and H.-D. Sues. 2013. A new archosauriform reptile from the Middle Triassic (Ladinian) of Germany. Journal of Systematic Palaeontology (advance online publication) DOI:10.1080/14772019.2013.781066

Abstract- Numerous well-preserved skeletal remains of a distinctive armoured archosauriform reptile from the Lower Keuper (Erfurt Formation; Middle Triassic: Ladinian: Longobardian) of Baden-Württemberg (Germany) represent a new taxon, Jaxtasuchus salomoni gen. et sp. nov. The dermal armour of Jaxtasuchus comprises transverse rows of four rectangular osteoderms each in the cervical, dorsal and caudal regions, with the individual plates closely resembling those of Doswellia kaltenbachi from Carnian strata in North America. The long and low maxilla of Jaxtasuchus held at least 15 teeth. The labial and lingual surfaces of the tall, only slightly recurved crowns of the maxillary teeth bear distinct vertical ridges and smooth mesial and distal carinae. The cervical region of the vertebral column is long. Phylogenetic analysis places Jaxtasuchus as the sister taxon to
Doswellia in Doswelliidae sensu Desojo et al. (2011). Doswelliidae is diagnosed by the coarsely reticulate, incised ornamentation of osteoderms composed of central regular pits of subequal size and contour, and a mostly smooth anterior articular lamina on each osteoderm. The discovery of Jaxtasuchus confirms that Doswelliidae had a wide palaeogeographical distribution during the latter half of the Triassic.

Reevaluation of the Enigmatic Archosaur Dyoplax arenaceus from the Upper Triassic of Germany

Maisch, M. W., Matzke, A. T., and T. Rathgeber. 2013. Re-evaluation of the enigmatic archosaur Dyoplax arenaceus O. Fraas, 1867 from the Schilfsandstein (Stuttgart Formation, lower Carnian, Upper Triassic) of Stuttgart, Germany. Neues Jahrbuch für Geologie und Paläontologie - Abhandlungen 267(3): 353-362
DOI: http://dx.doi.org/10.1127/0077-7749/2013/0317


Abstract - The holotype and only specimen of the small suchian archosaur Dyoplax arenaceus O. Fraas, 1867 from the Stuttgart-Formation (Schilfsandstein) of southwestern Germany is partially redescribed and re-evaluated. The type locality can be identified as Stuttgart, not Stuttgart-Feuerbach as erroneously suggested by previous authors. A re-description of the skull and the dorsal armour provides several new characters and a restoration of the skull is attempted for the first time. The phylogenetic placement of Dyoplax is discussed. Although it is agreed with previous authors that the taxon is not an aetosaur, its placement in Crocodylomorpha is questioned. Instead we demonstrate that Dyoplax has several important cranial and postcranial features in common with Erpetosuchus from the Late Triassic of Scotland and North America, and it is tentatively re-assigned to ?Erpetosuchidae.

First Evidence of Late Triassic Dicynodonts from Germany

Schoch, R. R. 2012. A dicynodont mandible from the Triassic of Germany forms the first evidence of large herbivores in the Central European Carnian. Neues Jahrbuch für Geologie und Paläontologie, Abhandlungen 26:119-123. DOI: http://dx.doi.org/10.1127/0077-7749/2012/0216


Abstract - A new partial mandible from the Schilfsandstein (Stuttgart Formation, Middle Carnian) of southern Germany forms the first unambiguous evidence of dicynodonts in the German Triassic. The preserved anterior part of the mandible is most consistent with kannemeyeriiform dicynodonts known from the Middle and Late Triassic of South America, southern Africa, North America, and the Eastern European Platform. Extrapolation of body size from the mandible indicates that the Schilfsandstein dicynodont was moderately large (∼2m estimated body length). This find is significant as it forms the first evidence of large herbivores in the Carnian pre-dinosaur faunas of Central Europe.         

Two New Triassic Temnospondyl Papers in the Journal Palaeontology

Witzmann, F., Schoch, R. R., Hilger, A., and N. Kardjilov. 2012. Braincase, palatoquadrate and ear region of the plagiosaurid Gerrothorax pulcherrimus from the Middle Triassic of Germany. Palaeontology 55:31-50. DOI: 10.1111/j.1475-4983.2011.01116.x

Abstract - The complete neurocranium plus palatoquadrate of the plagiosaurid temnospondyl Gerrothorax pulcherrimus from the Middle Triassic of Germany is described for the first time, based on outer morphological observations and micro-CT scanning. The exoccipitals are strong elements with paroccipital processes and well-separated occipital condyles. Anterolaterally, the exoccipitals contact the otics, which are mediolaterally elongated and have massive lateral walls. The otics contact the basisphenoid, which shows well-developed sellar processes. Anteriorly, the basisphenoid is continuous with the sphenethmoid region. In its posterior portion, the sphenethmoid gives rise to robust, laterally directed laterosphenoid walls, a unique morphology among basal tetrapods. The palatoquadrate is extensively ossified. The quadrate portion overlaps the descending lamina of squamosal and ascending lamina of pterygoid anteriorly, almost contacting the epipterygoid laterally. The epipterygoid is a complex element and may be co-ossified with otics and laterosphenoid walls. It has a broad, sheet-like footplate and a horizontally aligned ascending process that contacts the laterosphenoid walls. The degree of ossification of the epipterygoid, however, is subject to individual variation obviously independent from ontogenetic changes. The stapes of Gerrothorax is a large, blade-like element that differs conspicuously from the plesiomorphic temnospondyl condition. It has a prominent anterolateral projection which has not been observed in other basal tetrapods. Morphology of neurocranium and palatoquadratum of Gerrothorax most closely resembles that of the Russian plagiosaurid Plagiosternum danilovi, although the elements are less ossified in the latter. The extensive endocranial ossification of Gerrothorax is consistent with the general high degree of ossification in the exo- and endoskeleton of this temnospondyl and supports the view that a strong endocranial ossification cannot be evaluated as a plesiomorphic character in basal tetrapods.

Dias-da-Silva, S., Sengupta, D. P., Cabriera, S. F., and L. R. Da Silva. 2012. The presence of Compsocerops (Brachyopoidea: Chigutisauridae) (Late Triassic) in southern Brazil with comments on chigutisaurid palaeobiogeography. Palaeontology 55:163-172. DOI: 10.1111/j.1475-4983.2011.01120.x

Abstract - Chigutisauridae is the longest-lived trematosaurian clade (from early Triassic to early Cretaceous). They were reported in Argentina, Australia, India and South Africa. This contribution reports a putative chigutisaurid specimen in the Carnian of southern Brazil (Santa Maria Formation, Paraná Basin). The material comprises two skull fragments, a mandibular fragment, a clavicular blade and a humerus. Ontogenetic features point to an early development stage of the specimen. The presence of a long, straight and pointed tabular horn, which runs parallel to the skull midline towards its tip, and a distinctive projection in the posterior border of the postparietal indicates a close relationship of the Brazilian chigutisaurid with the Indian Compsocerops cosgriffi. Three distinctive and combined characters suggest that the Brazilian chigutisaurid is a distinctive specimen: the presence of an alar process of the jugal in the ventral margin of the orbit; jugal does not extend well beyond the anterior margin of the orbit; and tabular does not contact the parietal. These characters could justify the erection of a new taxon; however, they might reflect its immature ontogenetic stage as well. Accordingly, we attribute this new specimen to Compsocerops sp. Argentinean and Indian occurrences are dated as Norian, so the presence of a Carnian chigutisaurid in southern Brazil indicates that western Gondwana chigutisaurids have first occupied the Paraná Basin and later migrated towards west (to Argentina) and east (India). However, the presence of ghost chigutisaurid taxa cannot be dismissed, because their long temporal range contrasts with their still short (in comparison with other temnospondyl groups) geographic distribution. Hence, they might have been more geographically widespread than their fossil record suggests.

Braincase of the Temnospondyl Gerrothorax from the Middle Triassic of Germany

Witzmann, F., Schoch, R. R., Hilger, A. and N. Kardjilov. 2011. Braincase, palatoquadrate and ear region of the plagiosaurid Gerrothorax pulcherrimus from the Middle Triassic of Germany. Palaeontology (early online). doi: 10.1111/j.1475-4983.2011.01116.x

Abstract - The complete neurocranium plus palatoquadrate of the plagiosaurid temnospondyl Gerrothorax pulcherrimus from the Middle Triassic of Germany is described for the first time, based on outer morphological observations and micro-CT scanning. The exoccipitals are strong elements with paroccipital processes and well-separated occipital condyles. Anterolaterally, the exoccipitals contact the otics, which are mediolaterally elongated and have massive lateral walls. The otics contact the basisphenoid, which shows well-developed sellar processes. Anteriorly, the basisphenoid is continuous with the sphenethmoid region. In its posterior portion, the sphenethmoid gives rise to robust, laterally directed laterosphenoid walls, a unique morphology among basal tetrapods. The palatoquadrate is extensively ossified. The quadrate portion overlaps the descending lamina of squamosal and ascending lamina of pterygoid anteriorly, almost contacting the epipterygoid laterally. The epipterygoid is a complex element and may be co-ossified with otics and laterosphenoid walls. It has a broad, sheet-like footplate and a horizontally aligned ascending process that contacts the laterosphenoid walls. The degree of ossification of the epipterygoid, however, is subject to individual variation obviously independent from ontogenetic changes. The stapes of Gerrothorax is a large, blade-like element that differs conspicuously from the plesiomorphic temnospondyl condition. It has a prominent anterolateral projection which has not been observed in other basal tetrapods. Morphology of neurocranium and palatoquadratum of Gerrothorax most closely resembles that of the Russian plagiosaurid Plagiosternum danilovi, although the elements are less ossified in the latter. The extensive endocranial ossification of Gerrothorax is consistent with the general high degree of ossification in the exo- and endoskeleton of this temnospondyl and supports the view that a strong endocranial ossification cannot be evaluated as a plesiomorphic character in basal tetrapods.

Redescription of a Nearly Complete Skull of Plateosaurus from the Late Triassic of Germany

Prieto-Márquez, A., and M. A. Norell. 2011. Redescription of a nearly complete skull of Plateosaurus (Dinosauria: Sauropodomorpha) from the Late Triassic of Trossingen (Germany). American Museum Novitates 3727 :1-58. 
doi: http://www.bioone.org/doi/abs/10.1206/3727.2

Abstract - The nearly complete, disarticulated skull of AMNH FARB 6810, a specimen of the basal sauropodomorph Plateosaurus collected in 1925 from the Norian (Late Triassic) strata of the Knollenmergel beds of Trossingen (Germany), is redescribed. This study supports referral of AMNH FARB 6810 to P. erlenbergiensis on the basis of the following characters: occipital condyle above level of parasphenoid; basisphenoid with transverse, subvertical lamina extending between basipterygoid processes, with ventrally projecting median process; and peglike process projecting medially from the middle of the palatine. Furthermore, P. longiceps is regarded a junior synonym of P. erlenbergiensis because the type specimen of the latter is diagnostic (displaying the above-noted apomorphies of the braincase and palatine) and, chronologically, P. erlenbergiensis has priority over P. longiceps.

Redescription of the Sail-backed Poposauroid Ctenosauriscus from the Early Triassic of Germany

Available now at PLoSONE is a redescription of this important historic Triassic taxon which is the namesake of a clade of sail-backed poposauroid archosaurs that have come into recent prominence given the discovery of a well-preserved specimen of Arizonasaurus babbitti in 2002. 

  

Butler, R. J., Brusatte, S. L., Reich, M., Nesbitt, S. J., Schoch, R. R., and J. J. Hornung. 2011. The sail-backed reptile Ctenosauriscus from the latest Early Triassic of Germany and the timing and biogeography of the early archosaur radiation. PLoS ONE 6(10): e25693. doi:10.1371/journal.pone.0025693

 

Background

Archosaurs (birds, crocodilians and their extinct relatives including dinosaurs) dominated Mesozoic continental ecosystems from the Late Triassic onwards, and still form a major component of modern ecosystems (>10,000 species). The earliest diverse archosaur faunal assemblages are known from the Middle Triassic (c. 244 Ma), implying that the archosaur radiation began in the Early Triassic (252.3–247.2 Ma). Understanding of this radiation is currently limited by the poor early fossil record of the group in terms of skeletal remains.

Methodology/Principal Findings

We redescribe the anatomy and stratigraphic position of the type specimen of Ctenosauriscus koeneni (Huene), a sail-backed reptile from the Early Triassic (late Olenekian) Solling Formation of northern Germany that potentially represents the oldest known archosaur. We critically discuss previous biomechanical work on the ‘sail’ of Ctenosauriscus, which is formed by a series of elongated neural spines. In addition, we describe Ctenosauriscus-like postcranial material from the earliest Middle Triassic (early Anisian) Röt Formation of Waldhaus, southwestern Germany. Finally, we review the spatial and temporal distribution of the earliest archosaur fossils and their implications for understanding the dynamics of the archosaur radiation.

Conclusions/Significance

Comprehensive numerical phylogenetic analyses demonstrate that both Ctenosauriscus and the Waldhaus taxon are members of a monophyletic grouping of poposauroid archosaurs, Ctenosauriscidae, characterised by greatly elongated neural spines in the posterior cervical to anterior caudal vertebrae. The earliest archosaurs, including Ctenosauriscus, appear in the body fossil record just prior to the Olenekian/Anisian boundary (c. 248 Ma), less than 5 million years after the Permian–Triassic mass extinction. These earliest archosaur assemblages are dominated by ctenosauriscids, which were broadly distributed across northern Pangea and which appear to have been the first global radiation of archosaurs.

First Evidence of Aetosaurs from the Plateosaurus Quarry of Germany

Matzke, A. T., and M. W. Maisch. 2011. The first aetosaurid archosaur from the Trossingen Plateosaurus Quarry (Upper Triassic, Germany). Neues Jahrbuch für Geologie und Paläontologie – Abhandlungen (advance online publication) DOI: 10.1127/0077-7749/2011/0203http://www.ingentaconnect.com/content/schweiz/njbgeol/pre-prints/0203

Abstract -Two associated cervical paramedian osteoderms and one isolated paramedian osteoderm of an aetosaur from the famous Trossingen Plateosaurus Quarry are described. They represent the first evidence of aetosaurs from Trossingen as well as the stratigraphically youngest remains of this group from Germany. Therefore, the Trossingen assemblage consists now of four species level taxa. Hitherto only three species level taxa were known from this quarry. Plateosaurus longiceps and Proganochelys quenstedti from complete specimens as well as one tooth of cf. Liliensternus. The finds indicate that more small- to medium-sized taxa may be present in the Trossingen Quarry.

Review of the Lower Triassic Temnospondyl Fauna of Germany

Schoch, R. R. 2011. How diverse is the temnospondyl fauna in the Lower Triassic of southern Germany? Neues Jahrbuch für Geologie und Paläontologie - Abhandlungen 261:49-60. DOI: 10.1127/0077-7749/2011/0147

Abstract - Among the few fossiliferous Lower Triassic deposits in Europe, the Lower to Middle Buntsandstein of the German southwest has yielded a range of interesting finds. A review of the published and new material gives the following results: (1) The oldest identified remain is a mandible of a small stereospondyl from the Bernburg Formation (Induan); (2) A new palate from the early Olenekian (Badischer Bausandstein) stems from a basal capitosaur similar to Wetlugasaurus; (3) In the late Olenekian (Hardegsen Formation), two well-defined temnospondyl genera are present: the basal capitosauroids Meyerosuchus fuerstenbergianus and Odenwaldia heidelbergensis. Large remains formerly referred to Mastodonsaurus either pertain to a different family or are undiagnostic, pushing the first occurrence of mastodonsaurids into the Anisian. These finds highlight the presence of large aquatic predators by early Triassic time, revealing an increasing diversity of aquatic predators from the Induan to the Anisian. The documented faunal changes involved patterns of immigration and extinction/emigration rather than endemic evolution within the basin. In contrast to the northern parts of the basin, the genera Trematosaurus, Parotosuchus, and Sclerothorax are absent in the southern part, where Meyerosuchus and Odenwaldia existed.

The Aetosaurus Block

After a crazy spring things are pretty slow in the Triassic right now, with hardly any new papers coming out in recent weeks.  Even our fieldwork has pretty much ground to a halt given the recent extreme heat in the western U.S. Plus, I'm taking a week off from work, so things will be even slower; however, I'm expecting things to take off again pretty soon. 

In the meantime here is a shot showing the famous block of Aetosaurus ferratus specimens from the Lower Stubensanstein of Germany.

a close-up of one of the best preserved skulls

and an older reconstruction (from Wikipedia) showing several Aetosaurus perishing in a sandstorm.

These specimens (collected and first described in the 1800s) were the subject of an excellent detailed description by Rainer Schoch in 2007.

REFERENCE

Schoch, R. R. 2007. Osteology of the small archosaur Aetosaurus from the Upper Triassic of Germany. N. Jb. Geol. Paläont. Abh. 246:1– 35

Procolophonid-like Tetrapod from the Middle Triassic of Germany

Schoch, R. R. 2011. A procolophonid-like tetrapod from the German Middle Triassic. Neues Jahrbuch für Geologie und Paläontologie, Abhandlungen 259: 251–255. DOI: 10.1127/0077-7749/2011/0124
Abstract - A new find of a tetrapod mandible with unique teeth is reported from the Middle Triassic Lower Keuper. The outline of the dentary, morphology of the teeth and their emplacement suggest that the remain represents a new procolophonid parareptile. The abbreviated dentary, the high coronoid process, and the shape of the teeth resemble those of leptopleuronines. Unique features are the crown morphology, marked parallel striations, the possession of two similar, very large durophagous teeth, the edentulous anterior portion, and the lack of a coronoid suture. Thus, although the jaw resembles procolophonids in some features, it is argued that referral to this group needs recognition of more clearly established character-states, prompting the search for more complete material in the same deposits.

New Seed Ferns and Cycadophytes from the Middle Triassic of Germany

Kustatscher, E.,  and J. H. A. van Konijnenburg-van Cittert. 2010. Seed ferns and Cycadophytes from the Triassic Flora of Thale (Germany). Neues Jahrbuch für Geologie und Paläontologie - Abhandlungen 258: 195–217.

Abstract: This is the second paper on the Middle Triassic flora from Thale, dealing with the seed ferns and cycadophytes. Scytophyllum bergeri Bornemann is one of the common elements in the flora, in contrast to Sagenopteris sp. which is rare in the assemblage. However, it is the first evidence of Sagenopteris for the German Basin. The specimen described as ?Peltaspermum sp. is the only peltasperm ovuliferous organ known to date with attached ovule-bearing discs from the Middle Triassic of Europe. The cycad Apoldia tener (Compter 1883) Zijlstra et al. 2009 (formerly Sphenozamites tener Compter) is by far the most common fossil plant in the assemblage. Nilssonia cf. neuberi Stur ex Pott et al. is relatively rare. The genus Bjuvia is emended to accommodate more species of large entire leaves characterised by amphistomy and cycadalean stomatal morphology. This includes Bjuvia thalensis n. sp., which is a common element in the Thale flora but is unknown from elsewhere. Narrow, long entire leaves from the Middle and Upper Triassic of Europe have been described under the name Taeniopteris angustifolia Schenk, but have now been transferred to Taeniopteris kelberi n. sp. for nomenclatorial reasons. The macrosporophyll Dioonitocarpidium pennaeformis (Schenk) Ruhle von Lilienstern also occurs in the Thale flora. Two cycadalean leaf species (Apoldia tener and Bjuvia thalensis) from Thale flora sometimes have stomata with only two subsidiary cells instead of the usual 4-6, a feature that in gymnosperms occurs mainly in the Bennettitales.

On the Procompsognathus postcranium

F KNOLL (2008). On the Procompsognathus postcranium (Late Triassic, Germany)☆ Geobios, 41 (6), 779-786 DOI: 10.1016/j.geobios.2008.02.002

ABSTRACT - A review of the historical background of the material housed in the Staatliches Museum für Naturkunde (Stuttgart) and ascribed to Procompsognathus triassicus (Upper Triassic, Germany) is provided. The systematic position of the postcranial remains is discussed. The combined results of cladistic analyses suggest that the type material, an incomplete postcranial skeleton in two pieces (SMNS 12591), is from a theropod close to Segisaurus and Coelophysis. An isolated manus (SMNS 12352a) is definitely not theropodan, but could be from any small basal archosaur. The remarkable diversity of the carnivorous guild that dwelled in southern Germany before the end-Triassic events is underlined.

This is the latest paper in the Procompsognathus saga, regarding the taxonomic status of four specimens refered to this taxon, a purported theropod dinosaur, from the Late Triassic of Germany. To recap, Fraas (1913) named Procompsognathus triassicus based on a partial skeleton (SMNS 12591a) and skull (SMNS 12591), and refered it to the Dinosauria. Huene (1921) discussed the material further and assigned another partial skull (SMNS 12352) and manus (SMNS 12352a) from the same quarry to the taxon. Since that time this material has been reviewed by Ostrom (1981), Sereno and Wild (1992), Chatterjee (1993, 1998), Rauhut and Hungerbuhler (2000), Rauhut (2003), Allen (2004), and most recently by Knoll and Schoch (2006) which was an abstract previewing the current study by Knoll. All of these authors came to differing conclusions regarding the taxonomic affinities of the material as listed below:

Ostrom (1981)
SMNS 12591 - Procompsognathus triassicus
SMNS 12591a - Procompsognathus triassicus
SMNS 12352 - non Procompsognathus triassicus
SMNS 12352a - non Procompsognathus triassicus

Sereno and Wild (1992)
SMNS 12591 - Theropod similar to Coelophysis and Segisaurus
SMNS 12591a - Saltoposuchus connectens (Crocodylomorpha)
SMNS 12352 - Saltoposuchus connectens
SMNS 12352a - Saltoposuchus connectens

Chatterjee (1993, 1998)
SMNS 12591a - Theropod (contra Sereno and Wild [1992])

Rauhut and Hungerbuhler (2000)
SMNS 12591 - Theropod similar to Coelophysis and Segisaurus

Rauhut (2003)
Procompsognathus is a theropod but a metataxon

Allen (2004)
Procompsognathus is a non-dinosaurian ornithodiran

Knoll and Schoch (2006)
SMNS 12591 - Theropod similar to Coelophysis and Segisaurus
SMNS 12591a - Theropod, possibly tetanuran
SMNS 12352 - indeterminate crocodylomorph
SMNS 12352a - indeterminate crocodylomorph

The current paper (Knoll, 2008) follows the finding of Knoll and Schoch (2006) except that the isolated manus (SMNS 12352a) is considered to represent an unknown basal archosaur. Thus Knoll (2008) argues for a previously unrecognized diversity of carnivorous archosaurs in the Upper Triassic Stubensandstein. Hopefully this is the final word, but given the numerous differing hypotheses put forth this taxonomic argument may never be fully resolved.

The image is from here.

REFERENCES

Allen, D. 2004. The phylogenetic status of Procompsognathus revisited. Journal of Vertebrate Paleontology 24:34A.

Chatterjee, S. 1993. Procompsognathus from the Triassic of Germany is not a crocodylomorph. Journal of Vertebrate Paleontology 13:29A.

Chatterjee, S. 1998. Reassessment of the Procompsognathus skull, p. 6 in Wolberg, D.L., Gittis, K., Miller, S., Carey, L., and A. Raynor (eds.), Dinofest International. The Academy of Natural Sciences, Philadelphia.

Fraas, E. 1913. Die neuesten Dinosaurierfunde in der schwabischen Trias. Die Naturwissenschaften 1:1097-1100.

Huene, F.v. 1921. Neue Pseudosuchier und Coelurosaurier aus dem wurttembergischen Keuper. Acta Zoologica 2:329-403.

Knoll, F., and R. Schoch. 2006. Does Procompsognathus have a head? Systematics of an egnimatic Triassic taxon. Journal of Vertebrate Paleontology 26:86A.

Knoll, F. 2008. On the Procompsognathus postcranium (Late Triassic, Germany). Geobios 41:779-786.

Ostrom, J.H. 1981. Procompsognathus - theropod or thecodont? Palaeontographica A 175:175-195.

Rauhut, O.W.M. 2003. The interrelationships and evolution of basal theropod dinosaurs. Special Papers in Palaeontology 69:1-213.

Rauhut, O.W.M., and A. Hungerbeuhler. 2000. A review of European Triassic theropods. GAIA 15:75-88.

Sereno, P.C., and R. Wild. 1992. Procompsognathus: theropod, "thecodont", or both? Journal of Vertebrate Paleontology 12:435-458.