Reassessment of cf. Halticosaurus orbitoangulatus
from the Upper Triassic (Norian) of Germany –
a pseudosuchian, not a dinosaur
HANS-DIETER SUES FLS
1
* and RAINER R. SCHOCH
2
1
Department of Paleobiology, National Museum of Natural History, Smithsonian Institution, MRC
121, P.O. Box 37012, Washington, DC 20013-7012, USA
2
Staatliches Museum für Naturkunde Stuttgart, Rosenstein 1, D-70191 Stuttgart, Germany
Received 4 October 2012; revised 31 January 2013; accepted for publication 19 March 2013
The holotype of cf. Halticosaurus orbitoangulatus Huene, 1932, comprises an incomplete and macerated but
associated skull of an archosaurian reptile from the middle (second) Stubensandstein (middle Löwenstein
Formation; Upper Triassic: Norian) of Baden-Württemberg, Germany. It was originally interpreted as a theropod
dinosaur but more recently it has been suggested that this taxon has crocodylomorph affinities. Detailed
preparation of the holotype of cf. H. orbitoangulatus has revealed much new anatomical information and permitted
reassessment of its affinities. The maxilla lacks both a distinct antorbital fossa and a medial bony lamina bordering
the antorbital fenestra. The lateral surface of the dentary bears a pronounced horizontal ridge. The squamosal
differs from that of basal crocodylomorphs in being L-shaped rather than arcuate in dorsal view, lacking a
dorsolateral overhang, and lacking an interlocking contact with the paroccipital process as, for example, in the
basal crocodylomorph Saltoposuchus connectens from the same horizon and locality. Phylogenetic analysis placed
cf. H. orbitoangulatus amongst loricatan pseudosuchians (but not amongst Crocodylomorpha) rather than amongst
theropod dinosaurs. The holotype of cf. H. orbitoangulatus represents a previously unrecognized taxon of loricatan
pseudosuchian, which is here named Apatosuchus orbitoangulatus and set apart from other known Norian-age
non-crocodylomorph loricatans by its apparently much smaller size.
© 2013 The Linnean Society of London, Zoological Journal of the Linnean Society, 2013, 168, 859–872.
doi: 10.1111/zoj.12038
ADDITIONAL KEYWORDS: Archosauria – Baden-Württemberg – Loricata – Löwenstein Formation –
Paracrocodylomorpha.
INTRODUCTION
Non-avian theropod dinosaurs were the dominant
predators in terrestrial ecosystems for much of the
Mesozoic Era. The oldest known undisputed members
of this group date from the Norian stage of the Late
Triassic (Nesbitt, Irmis & Parker, 2007). However, the
existence of late Carnian representatives of Sauro-
podomorpha (Martínez & Alcober, 2009; Cabreira
et al., 2011; Martinez et al., 2011), the sister taxon of
Theropoda, implies that the latter clade must have
the same age.
Late Triassic continental strata of the Middle
Keuper in Baden-Württemberg, Germany, have
yielded numerous, well-preserved skeletal remains
of the basal sauropodomorph dinosaur Efraasia
minor (see Yates, 2003) but surprisingly few fossils
of theropods (Huene, 1907–1908, 1932; Rauhut &
Hungerbühler, 2000). Amongst the latter are several
specimens from the middle (second) Stubensandstein
(Löwenstein Formation; Norian) of the Stromberg
region in northern Baden-Württemberg (Fig. 1).
Almost all the specimens of dinosaurs and crocody-
lomorph archosaurs known from the Löwenstein For-
mation were recovered from a disused sandstone
quarry known as the ‘Weisser Steinbruch’ and located *Corresponding author. E-mail: suesh@si.edu
Zoological Journal of the Linnean Society, 2013, 168, 859–872. With 7 figures
© 2013 The Linnean Society of London, Zoological Journal of the Linnean Society, 2013, 168, 859–872 859
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