Journal of Vertebrate Paleontology

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Osteology and phylogenetic relationships of a new

archosauriform reptile from the Middle Triassic
(Anisian) of Germany

Hans-Dieter Sues, Stephan N. F. Spiekman & Rainer R. Schoch

To cite this article: Hans-Dieter Sues, Stephan N. F. Spiekman & Rainer R. Schoch (12
Jun 2024): Osteology and phylogenetic relationships of a new archosauriform reptile
from the Middle Triassic (Anisian) of Germany, Journal of Vertebrate Paleontology, DOI:
10.1080/02724634.2024.2357326

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Journal of Vertebrate Paleontology e2357326 (19 pages)
Published with license by the Society of Vertebrate Paleontology
DOI: 10.1080/02724634.2024.2357326

ARTICLE

OSTEOLOGY AND PHYLOGENETIC RELATIONSHIPS OF A NEW ARCHOSAURIFORM

REPTILE FROM THE MIDDLE TRIASSIC (ANISIAN) OF GERMANY

HANS-DIETER SUES, *,1 STEPHAN N. F. SPIEKMAN, 2 and RAINER R. SCHOCH 2,3

1
Department of Paleobiology, National Museum of Natural History, Smithsonian Institution, MRC 121, P.O. Box 37012, Washington,
DC 20013-7012, U.S.A., [email protected];
2
Staatliches Museum für Naturkunde Stuttgart, Rosenstein 1, D-70191 Stuttgart, Germany, [email protected],
[email protected];
3
Institut für Biologie, Fachgebiet Paläontologie, Universität Hohenheim, D-70599 Stuttgart, Germany

ABSTRACT—Skeletal remains of archosauriform reptiles from the Lower and lower Middle Triassic Buntsandstein Group in
the Central European Basin are rare. This paper reports on a partial, almost completely disarticulated skeleton of a previously
unknown archosauriform reptile from the lower Middle Triassic (Anisian) Röt Formation of Rotfelden in Baden-Württemberg
(Germany). Marcianosuchus angustifrons gen. et sp. nov. is distinguished by the following combination of features:
posterodorsal process of premaxilla slender, slightly inclined posterodorsally, and with rounded apex; posterodorsal surface
of frontal covered by slightly diverging, fine longitudinal grooves; squamosal with distinct lateral ridge extending for entire
length of element; teeth with proportionately small, weakly recurved crowns; humerus considerably shorter than femur;
distal shaft of ischium rod-like, not plate-like; pubis with flat distal portion forming ‘pubic apron’: neural spines of posterior
cervical and anterior dorsal vertebrae with transversely greatly expanded apices that are Y-shaped in anterior or posterior
view; dorsal vertebrae with centra taller dorsoventrally than long anteroposteriorly; dorsal osteoderms more or less
rectangular in outline, longer anteroposteriorly than wide mediolaterally, with slightly rounded anterior and concave
posterior margins and bearing dorsal ridge or eminence; and dorsal surfaces of osteoderms with unsculptured anterior
region and posterior region bearing keel or eminence and covered by sculpturing of radially arranged grooves and pits.
Marcianosuchus angustifrons represents the first definitive record of a non-archosaurian archosauriform from the
Buntsandstein Group of Germany. Its body plan most closely resembles that of Euparkeria capensis among well-known non-
archosaurian archosauriform reptiles, but the two taxa are clearly distinct.

http://zoobank.org/urn:lsid:zoobank.org:pub:582821E9-F8576-4CD5-96D2-E0A1BAC2C87C

SUPPLEMENTARY FILES—Supplementary files are available for this article for free at www.tandfonline.com/UJVP.

Citation for this article: Sues, H-D., Spiekman, S. N. F., & Schoch, R. R. (2024) Osteology and phylogenetic relationships of
a new archosauriform reptile from the Middle Triassic (Anisian) of Germany. Journal of Vertebrate Paleontology. https://doi.
org/10.1080/02724634.2024.2357326

Submitted: January 16, 2024

Revisions received: May 11, 2024
Accepted: May 14, 2024

INTRODUCTION
Present-day birds and crocodylians together with their extinct with Archosauriformes as Archosauromorpha (Benton, 1985;
relatives, including non-avian dinosaurs and pterosaurs, consti­ Dilkes, 1998; Ezcurra, 2016; Sues, 2019). Following the end-
tute the clade Archosauria. Archosaurs and their closest relatives Permian extinction event, archosauromorph reptiles rapidly
such as Proterosuchidae and Euparkeriidae are grouped diversified and dispersed across Pangea during the Early and
together as Archosauriformes. In addition, various other, early Middle Triassic (Ezcurra, 2016; Ezcurra & Butler, 2018;
mainly Triassic diapsid reptiles that are more closely related to Foth et al., 2016). Commencing in the Middle Triassic, the evol­
archosauriforms than to lepidosauromorphs have been united utionary radiation of archosaurs profoundly reshaped terrestrial
ecosystems. However, there are still relatively few occurrences of
Early and early Middle Triassic fossils that document the early
*Corresponding author. evolutionary history of archosauriforms.
This work was authored as part of the Hans-Dieter Sues' official duties In the Central European Basin, predominantly continental red
as an Employee of the United States Government and is therefore a work beds of Early to early Middle Triassic (Indian–Anisian) age, col­
of the United States Government. In accordance with 17 U.S.C. 105, no lectively known as the Buntsandstein Group (Deutsche Stratigra­
copyright protection is available for such works under U.S. Law.
Stephan N. F. Spiekman and Rainer R. Schoch hereby waive their right
phische Kommission, 2013; Fig. 1), have yielded many trackways
to assert copyright, but not their right to be named as co-authors in the attributable to a variety of tetrapods (Haubold, 1971; Klein &
article. Lucas, 2021) but few skeletal remains (Krebs, 1969; Schoch,
Color versions of one or more of the figures in the article can be found 2021; Sues et al., 2022). The abundance and diversity of archosaur­
online at www.tandfonline.com/ujvp. omorph footprints in the Buntsandstein Group, especially in the

Published online 12 Jun 2024

 Sues et al.—Middle Triassic archosauriform (e2357326-2)

FIGURE 1. Map of the state of Baden-Württemberg (Germany) with location of the town of Rotfelden (left) and stratigraphic column for the Bunt­
sandstein Group in the Central European Basin (right). Arrow points to the level in the Röt Formation from which the holotype of Marcianosuchus
angustifrons was recovered. The distance between Stuttgart and Rotfelden is approximately 41 km (25.2 miles). Data for the stratigraphic column
based on Deutsche Stratigraphische Kommission (2013).

Hardegsen and Solling formations of the Middle Buntsandstein During a field trip to the Kössig quarry in 1972, Rupert Wild
Subgroup (Haubold, 1971; Klein & Lucas, 2021), indicate that (former curator at the Staatliches Museum für Naturkunde Stutt­
the diversity of these reptiles was much greater than the known gart) collected a sandstone slab with a largely disarticulated skel­
record of body fossils would suggest. eton of a reptile, which is now housed in the museum’s collections
The Röt Formation, which forms the top of the Upper Bunt­ (SMNS 91318). It was recovered from talus that piled up after the
sandstein Subgroup, is early Anisian (Aegean) in age quarry was no longer in use. Thus, its exact stratigraphic origin
(Menning & Käding, 2013). It encompasses variegated sand­ within the originally exposed sequence of the Röt 4 Subformation
stones and mudstones, which typically are purple to red-brown can no longer be determined. However, the bone-bearing rock, a
in color and contain a succession of paleosols that Ortlam fine- to medium-grained, reddish-brown sandstone, best matches
(1967) referred to as ‘Violet Horizons’ (VH 1–5). The strata of the lithological description of unit 9 in Ortlam’s (1968) strati­
this formation have yielded skeletal remains of temnospondyls graphic section for the Kössig quarry.
and reptiles (Krebs, 1969; Ortlam, 1967, 1970; Schoch, 2021; The skeletal remains of SMNS 91318 represent a previously
Schoch et al., 2023). unknown non-archosaurian archosauriform from the lower
The Kössig quarry is a small, long abandoned quarry some Middle Triassic and the first from Central Europe, rendering
100 m south of the village of Rotfelden in the Calw district of this find particularly important. The study presents a description
the northern Black Forest region of Baden-Württemberg of this specimen and assesses its phylogenetic relationships.
(Germany) (Fig. 1). It produced sandstones for local buildings
during the 1960s and exposed a 6- to 8-m-thick section within
the top of the Röt 4 Subformation (Deutsche Stratigraphische
Kommission, 2013). Tetrapod remains have repeatedly been MATERIALS AND METHODS
recovered from different horizons within this section (Ortlam,
Phylogenetic Analysis
1967, 1968, 1970), including an excellently preserved cranium
of the capitosauroid temnospondyl Eocyclotosaurus woschmidti We added scores for the new archosauriform to the character-
(holotype: SMNS 51562; Kamphausen, 1989; Rinehart et al., taxon matrix used by Sookias (2016), which is based on Nesbitt
2015) and skeletal remains of the tanystropheid archosauro­ (2011) and Sookias et al. (2014b). We selected this matrix
morph Amotosaurus rotfeldensis (holotype: SMNS 50830; because it focuses specifically on the interrelationships of Eupar­
Fraser & Rieppel, 2006; Spiekman et al., 2021a). These fossil- keriidae and closely related taxa to which the new taxon exhibits
bearing strata form part of a sequence of fluvial to nearshore- close morphological similarities. We also scored Tarjadia ruthae
marine mudstones and sandstones. The tetrapod remains into the matrix based on its recent descriptions by Ezcurra
occurred together with current marks, invertebrate traces, tetra­ et al. (2017) and Desojo et al. (2024) to add the pseudosuchian
pod tracks, and fossil plant remains (Ortlam, 1967, 1968). clade Erpetosuchidae. We analyzed the character-taxon matrix
 Sues et al.—Middle Triassic archosauriform (e2357326-3)

using a maximum parsimony analysis in TNT 1.6 (Goloboff & no duplication of particular bones (e.g., left humerus). Thus, we
Morales, 2023). confidently interpret SMNS 91318 as the cranial and postcranial
We followed Sookias (2016) in excluding Dongusuchus efremovi remains of a single individual.
and character 93 a priori. Consequently, the final analysis includes Etymology—The genus name Marcianosuchus is derived from
65 taxa and 404 active characters. Following Sookias (2016), Marciana silva (‘border forest’), the Latin designation for the
Youngina capensis was designated as the outgroup and characters Black Forest used by the Roman historian Ammianus Marcelli­
58, 130, 132, 160, 224, 246, 267, 268, 270, 290, 294, 309, 347, 362, and nus in the fourth century CE, and from the Greek σοῦχος for
389 were treated as ordered. The TNT analysis was performed the ancient Egyptian crocodile-headed deity Sobek, a commonly
using an equal-weights analysis through a Traditional Search used suffix for genus names of non-mammalian tetrapods. The
with 1000 Wagner trees replications and TBR branch swapping species name angustifrons is derived from the Latin words angu­
holding 10 trees per replicate. Bremer and bootstrap GC stus, narrow, and frons, forehead, and refers to the transversely
support values were calculated, the latter through a Traditional narrow frontal region of the cranium.
Search with 1000 iterations. The STATSb.run script (see sup­ Diagnosis—Marcianosuchus angustifrons is diagnosed by the
plementary material in Spiekman et al. [2021b]) was used to calcu­ following combination of features (hypothesized autapomorphy
late homoplasy indices. The NEXUS and TNT files used to indicated by *): posterodorsal process of premaxilla slender,
conduct the analyses are included in the Supplementary Material. slightly inclined posterodorsally, and with rounded apex; poster­
odorsal surface of frontal covered by slightly diverging, fine, and
anteroposteriorly extending grooves; *squamosal with distinct
Photogrammetry lateral ridge extending for entire length of element; teeth with
The three-dimensional (3D) photogrammetric model was gen­ proportionately small, weakly recurved crowns; humerus con­
erated following the methodology of Mujal et al. (2020) and siderably shorter than femur; distal shaft of ischium rod-like,
using the freeware Meshroom to generate the mesh and not plate-like; pubis with flat distal portion forming ‘pubic
texture from photographs of the specimen taken under various apron’: neural spines of posterior cervical and anterior dorsal
angles. The mesh was subsequently processed for cleaning, vertebrae with transversely greatly expanded apices that are Y-
scaling, and orientation in MeshLab, and ParaView was used to shaped in anterior or posterior view; dorsal vertebrae with
create images with false color renderings to better visualize centra taller dorsoventrally than long anteroposteriorly; *dorsal
and enhance certain features. osteoderms more or less rectangular in outline, being longer
anteroposteriorly than wide mediolaterally, with slightly
Institutional Abbreviations—SMNS, Staatliches Museum für rounded anterior and concave posterior margins and bearing
Naturkunde Stuttgart, Stuttgart, Germany. dorsal ridge or eminence; and dorsal surfaces of osteoderms
Anatomical Nomenclature—The authors use the terminology with unsculptured region anteriorly and sculpturing consisting
proposed by Gower (2003) for the orientation of limb bones. of radially arranged grooves and pits more posteriorly.
Locality, Horizon, and Age—Kössig quarry, about 100 m south
ZooBank Registration—The nomenclatural acts in this publi­ of Rotfelden, Ebhausen municipality, Calw district, Baden-Würt­
cation are registered at ZooBank. The new genus Marcianosu­ temberg, Germany. Geographic coordinates: latitude 48°
chus is registered under LSID urn:lsid:zoobank.org: 36’9.07"N, longitude 8°41’51.79"E. Röt Subformation 4, Röt For­
act:944F8B65-C04F-4DAF-B6CC-1F71C581785A. The new mation, Upper Buntsandstein Subgroup, Buntsandstein Group
species Marcianosuchus angustifrons is registered under LSID (Deutsche Stratigraphische Kommission, 2013). Age: early
urn:lsid:zoobank.org:act:EDB1E4D4-98AF-4F6E-807A- Middle Triassic (Anisian: Aegean; Menning & Käding, 2013).
E7BA04B71A1A. Comments—The partial skeleton SMNS 91318 is preserved on
a slab of reddish-brown sandstone, which roughly measures 60 ×
30 cm and has an average thickness of about 2–3 cm (Fig. 2).
SYSTEMATIC PALEONTOLOGY
There are two associated small pieces of bone-bearing rock,
DIAPSIDA Osborn, 1903 one of which preserves an incomplete right humerus. The
ARCHOSAUROMORPHA Huene, 1946 sensu Gauthier, bones are yellowish to gray in color and, in places, stained slightly
2020a blue by oxidized vivianite. Many bones underwent diagenetic
ARCHOSAURIFORMES Gauthier, 1994 sensu Gauthier, compaction, distortion, and flattening. Many bony surfaces
2020b show networks of fine fractures. A few larger skeletal elements
MARCIANOSUCHUS ANGUSTIFRONS, gen. et sp. nov. (e.g., scapulae) appear to have been aligned by current activity;
the latter is probably also responsible for the loss of many
Holotype—SMNS 91318, mostly disarticulated but associated smaller, lighter elements such as most osteoderms and gastralia.
elements of a skeleton including a right premaxilla, right The specimen preserves an accumulation of mostly disarticu­
maxilla, frontals, right postorbital, right quadratojugal, right squa­ lated cranial, axial, and appendicular bones. As discussed
mosal, right quadrate, articulated left articular, surangular, and above, there is no evidence to suggest that these skeletal
angular, left prearticular, teeth, both scapulae, left coracoid, remains represent either more than a single individual or more
both humeri, partial right ilium, right ischium, both pubes, both than one taxon. The only still articulated elements are a
femora, partial tibia, fragment of ?fibula, vertebrae from various portion of the skull roof, the bones forming the posterior
regions of the vertebral column, neural arches, ribs, gastralia, portion of the left mandibular ramus, and two articulated
and osteoderms. There are additional bones that could not yet dorsal vertebrae. Most vertebrae present open neurocentral
be identified due to the lack of detailed information for possibly sutures, and several neural arches were completely separated
homologous elements in other known archosauriform reptiles. from their respective centra and turned over during postmortem
Most bones appear to be of congruent size to belong to a single transport. This condition has traditionally been associated with
individual, using the skeletal proportions of the well-known somatic immaturity, but Irmis (2007) cautioned that the ontogen­
early-diverging archosauriform Euparkeria capensis (Ewer, etic sequence and timing of closure of the neurocentral sutures
1965; Sookias et al., 2020) as a guide. Many of them closely can show considerable variation.
resemble the homologous elements in certain non-archosaurian The body size of SMNS 91318 is relatively small for a Middle
archosauriforms, particularly Euparkeria capensis, Dorosuchus Triassic archosauriform, with a reconstructed femoral length of
neoetus, and Halazhaisuchus qiaoensis. Furthermore, there is 115 mm. This compares well with the length of the femora in
 Sues et al.—Middle Triassic archosauriform (e2357326-4)

FIGURE 2. A, three-dimensional photogrammetric model of the holotype of Marcianosuchus angustifrons (SMNS 91318). B, photogrammetric model
with labeling of key skeletal elements. Abbreviations: atl, atlantal neural arches; co, coracoid; f, frontal; fe, femur; fi, fibula; h, humerus; il, ilium; is,
ischium; l, left; m, maxilla; o, osteoderm; pa, prearticular; pd, postdentary bone complex; pm, premaxilla; po, postorbital; pu, pubis; q, quadrate; qj,
quadratojugal; r, right; sc, scapula; sq, squamosal; ti, tibia. Scale bars equal 10 cm.
 Sues et al.—Middle Triassic archosauriform (e2357326-5)

FIGURE 3. Marcianosuchus angustifrons (SMNS 91318, holotype). A, right premaxilla in medial view. B, anterior portion of right maxilla in medial
view. Abbreviations: aof, antorbital fenestra; apm, anterodorsal process of premaxilla; d, depression; en, external narial fenestra; idp, interdental plate;
mf, maxillary foramen; mpm, palatal (= medial) process of maxilla; rt, replacement tooth; ppm, posterodorsal process of premaxilla; 1–4, premaxillary
alveoli 1–4. Scale bars each equal 1 cm.

the holotype of the euparkeriid “Turfanosuchus shageduensis” Osmolskina czatkoviensis in not being distinctly downturned
(127 mm long; Sookias et al., 2014b) and the holotype of Dorosu­ (Borsuk-Białynicka & Evans, 2009; Sookias, 2016). Euparkeria
chus neoetus (98.3 mm; Sookias et al., 2014a), respectively, but capensis (Ewer, 1965; Sookias, 2016) and O. czatkowicensis
the largest known femur of Euparkeria capensis cited by (Borsuk-Białynicka and Evans, 2009) share with SMNS 91318
Sookias et al. (2020) has a length of only 65.5 mm as preserved. the presence of four premaxillary teeth (Sookias et al., 2020);
Given the persistence of open neurocentral sutures, however, elsewhere, this character-state is present in non-crocodylomorph
SMNS 9318 had probably not yet attained somatic maturity, (‘rauisuchian’) loricatans (Nesbitt, 2011; Weinbaum, 2011).
and it is likely that the femur of an adult would have been Maxilla—Only the anterior part of the right maxilla is exposed
longer than those of the aforementioned taxa. in lingual view at the lateral margin of the right femur, which
overlies and conceals the remainder of the element (Fig. 3B).
Furthermore, the dorsal portion of the element is partially
Skull
obscured by attached fragments of unidentifiable bone, possibly
Premaxilla—The right premaxilla is exposed in medial (= of palatal elements. Although only partially exposed, the maxilla
lingual) view (Fig. 3A). It is a gracile, in medial view nearly U- is clearly longer than the premaxilla. Its deep, lappet-shaped
shaped element. It has a horizontal, tooth-bearing bony bar palatal process curves medially and ventrally, similar to the
(length: 15 mm), a gently posterodorsally and somewhat medi­ more gracile process in Euparkeria capensis (Sookias et al.,
ally curving anterodorsal (= nasal) process forming the anterior 2020). Given the degree of its ventral curvature, the premaxilla
margin of the external narial fenestra, and a slender, nearly ver­ would have been deflected to some degree. The ventral margin
tical, and distally tapering posterodorsal (=postnarial) process of the palatal process is continuous with the alveolar margin of
that makes up the posterior margin of the fenestra and which the maxilla posterior to it, whereas its dorsal margin curves med­
has a flat medial surface and a rounded apex. The posterodorsal ioventrally. Based on its orientation, it is unlikely that it con­
process of the premaxilla is shorter than the anterodorsal tacted its antimere medially. A medial contact between the
process. It differs from that of Euparkeria capensis, which is rela­ maxillae is also absent in certain non-archosaurian archosauri­
tively broader and extends almost perpendicular to the alveolar forms, including Euparkeria capensis (Sookias et al., 2020), but
margin of the bone (Sookias et al., 2020). The external narial it is typically present in archosaurs (Nesbitt, 2011). The medial
fenestra is positioned immediately posterior to the anterodorsal surface at the base of the (largely concealed) dorsal process of
process and faced laterally. It is bounded by the anterior, pos­ the maxilla forms a distinct depression, which is delimited by a
terior, and ventral margins of the premaxilla. The bar-like hori­ ridge ventrally. A similar depression is present in E. capensis
zontal portion of the premaxilla is narrower mediolaterally (Gow, 1970; Sookias et al., 2020). The tooth-bearing portion of
than deep dorsoventrally. Its anterior portion arcs anterome­ the maxilla ventral to the antorbital fenestra is dorsoventrally
dially. The medial surface of the tooth-bearing bar has four open­ low and slightly expands dorsoventrally towards its posterior
ings, each of which opens into an alveolus. The third alveolus end, as in several archosauriforms, including Euparkeria capensis
contains the crown of a small replacement tooth deep within it. (Nesbitt, 2011). No teeth are preserved in situ. Four interdental
In overall shape, the premaxilla most closely resembles that of plates are slightly inset from the alveolar margin. They are low,
Euparkeria capensis (Sookias et al., 2020) and differs from not fused to each other or to the body of the maxilla, and have
those of other known archosauriform reptiles (Ezcurra, 2016). somewhat rounded apical margins. The presence of distinct inter­
It differs from that of Euparkeria capensis in the absence of a dental plates is diagnostic for Archosauriformes (Ezcurra et al.,
low ventral peak projecting into the external narial fenestra at 2023). A large foramen, which is continued by a short groove
the anteroposterior midpoint of that opening and from that of posteriorly, is present at the level of the fourth interdental
 Sues et al.—Middle Triassic archosauriform (e2357326-6)

FIGURE 4. Marcianosuchus angustifrons

(SMNS 91318, holotype). A, frontals and rem­
nants of nasals in dorsal view. B, right postorbi­
tal in medial view and isolated tooth.
Abbreviations: app, anteromedial process of
postorbital; f, frontal; fj, recessed area for
contact with jugal; gr, grooves on frontal; n,
nasal; or, orbit; ppp, posterior process of post­
orbital; t, tooth; vpp, ventral process of postor­
bital. Scale bars each equal 1 cm.

plate on the medial surface of the maxilla. It closely resembles Squamosal—The right squamosal is exposed in lateral view
similar openings on the maxillae of other archosauriforms (e.g., (Fig. 5A). It has an anterior process for contact with the posterior
Batrachotomus kupferzellensis), which served as a passage for process of the postorbital, a short, free-standing posterior
the maxillary artery and associated neurovascular structures process, and a ventral process. In lateral view, the lateral
(Gower, 1999). As in other non-archosaurian archosauriforms, surface of the squamosal forms a prominent anteroposteriorly
the maxilla did not contribute to the posterior margin of the extending ridge. This lateral ridge extends from the (slightly
external naris, as can be deduced from the tall posterodorsal damaged) anterior end of the squamosal back to the posterior
process of the premaxilla (Nesbitt, 2011). end of the posterior process, overhanging the ventrolateral
Frontal—The right and part of the left frontal are exposed in surface of the element. Such a distinct laterally facing, overhan­
dorsal view, but no suture is evident between them (Fig. 4A). ging ridge is not known among non-archosaurian archosauri­
Posteriorly, the frontals expand transversely and would have forms except the early-diverging phytosaur Parasuchus spp.
broadly contacted the (not preserved) parietals along a trans­ (Kammerer et al., 2016), but, among Triassic archosaurs, occurs
verse suture. They are narrow transversely between the orbits to a similar extent in loricatan pseudosuchians including crocody­
and form smooth orbital margins. Toward their anterior ends, lomorphs (Nesbitt, 2011). The anterior process of the squamosal
the frontals expand again transversely. There is no visible is rather narrow transversely, bordering the supratemporal fenes­
contact with the poorly preserved nasals. Anterolaterally, the tra laterally. Its slightly mediolaterally concave dorsal surface
right frontal possibly contacts the prefrontal, although bears weak sculpturing. There is no distinct slot for the reception
the identification of this bone can only be made tentatively. of the posterior process of the postorbital, which presumably
The dorsal surfaces of the frontals are, for the most part, flat. contacted the anterior process ventral to the lateral ridge. The
Much of the bone in the anterior portion of both elements medial region of the squamosal is still concealed in the matrix.
was lost but, near the posterior contact with the parietals, the The gently curved ventral process made up part of the posterior
dorsal surface of the right frontal bears fine grooves that slightly margin of the infratemporal fenestra. Its posterolateral surface is
diverge from the posterior end. Similar grooving is present on deeply concave for contact with the quadratojugal, similar to the
the frontals of Euparkeria capensis (Sookias et al., 2020); the condition in Euparkeria capensis (Sookias et al., 2020).
frontals of the proterochampsian Chanaresuchus bonapartei Quadratojugal—The right quadratojugal is exposed in lateral
also bear fine ridges but these ridges form a radiating pattern view (Fig. 5B). It is roughly ‘L’-shaped, with a posteriorly
from a central point on each bone (Trotteyn & Ezcurra, rounded main body (sensu Sookias et al., 2020) at the junction
2020). Posterolaterally, the frontal forms a short, posterolater­ between the slightly anterodorsally extending dorsal process and
ally extending and tapering process that presumably contacted the slender, anteriorly tapering horizontal (= infratemporal)
the postfrontal and parietal. process. The two processes formed the rounded posteroventral
Postorbital—Most of the right postorbital is exposed in medial corner of the infratemporal fenestra. The dorsal process is incom­
view adjacent to the shaft of the left femur (Fig. 4B). It is a trir­ plete and its basal portion is slightly recessed along its anterior
adiate bone with a thick central portion from which a short, ante­ margin. The quadratojugal resembles the homologous elements in
romedially curving anterior process, a long, slender, and Euparkeria capensis (Sookias et al., 2020) and Osmolskina czatko­
anteroventrally curving ventral (= jugal) process, and a robust wicensis (Borsuk-Białynicka & Evans, 2009), but the angle enclosed
posterior process (much of which is concealed by the femur) by its horizontal and dorsal processes is smaller and the main body
extend. The ventral process tapers to a point distally and has of the element is relatively more robust than in the latter taxa.
an anteroposteriorly concave posteromedial surface for contact Quadrate—The left quadrate is partially exposed in medial
with the dorsal (= ascending) process of the jugal, with which it view (Fig. 5C). Its dorsal portion with the small head curves
formed the postorbital bar. somewhat posteriorly. A distinct posterior ridge bounds a
 Sues et al.—Middle Triassic archosauriform (e2357326-7)

FIGURE 5. Marcianosuchus angustifrons (SMNS 91318, holotype). A, right squamosal in lateral view. B, right quadratojugal in lateral view. C, left
quadrate in medial view. Abbreviations: aqj, recessed area for contact with quadratojugal; apq, anterior process of quadratojugal; aps, anterior process
of squamosal; dpq, dorsal process of quadratojugal; itf, infratemporal fenestra; lr, lateral ridge on squamosal; qh, head of quadrate; pps, posterior
process of squamosal; ptf, area for contact with quadrate flange of pterygoid; vps, ventral process of squamosal. Scale bars equal 1 cm (A) and 5
mm (B, C), respectively.

depression on the posteromedial surface of the pterygoid flange et al., 2014a), and is pierced by a foramen (not discernible in
of the quadrate. The ventral articular end of the element is con­ dorsal view) for the passage of the chorda tympani. The antero­
cealed by overlying bone. posteriorly rather short retroarticular process forms the pos­
Palatine—A small piece of bone of what appears to be the terior margin of the postcotylar fossa. Medially, it bears a
anterior part of the left palatine is exposed adjacent to the first distinct dorsomedial projection (of which only the base is pre­
of the two articulated dorsal vertebrae. It has a notch that poss­ served). Much of the dorsal surface of the retroarticular
ibly represents the posterior margin of the choana. process is concave transversely.
Vomer—A slender, slightly spindle-shaped element preserved Prearticular—A complete left prearticular (anteroposterior
near the distal end of the left humerus is possibly a vomer. It pre­ length: 63 mm) is exposed in lateral view (Fig. 6C). It is markedly
sents no noteworthy features. bowed anteroposteriorly, with a concave dorsal margin and a
Surangular—The anteroposteriorly long surangular is rep­ nearly straight ventral margin. The mediolaterally flat anterior
resented by the complete left element, which is ‘L’-shaped as and posterior ends of the slender bone are more than twice as
exposed in dorsal view (Fig. 6A, B). It forms the nearly flat deep dorsoventrally than the mid-section. The anterior end is dor­
dorsal margin of the postdentary region of the left mandibular soventrally less deep than the posterior one, tapering anteriorly
ramus and arcs over the adductor fossa. In dorsal view, the and bearing a short longitudinal ridge, which terminates at the
anterior portion of the surangular curves anteromedially and anterior tip of the prearticular. The posterior end contacted the
tapers toward its anterior end. Medial to it, parts of the articular posteromedially. The mid-section of the prearticular is
angular and of an unidentified flat bone are exposed. Posteriorly, distinctly concave dorsoventrally, forming the medial wall of a
the surangular sharply turns medially and forms a preglenoid lateral groove, which housed Meckel’s cartilage and is bounded
process, which becomes anteroposteriorly wider toward its by ridges dorsally and ventrally. The dorsal ridge forms the
medial end. This process forms the anterior margin of the med­ ventral margin of the adductor fossa whereas the ventral margin
iolaterally expanded glenoid surface of the jaw joint, which has presents a narrow shelf that contacted the angular.
somewhat raised anterior and posterior margins. The glenoid Teeth—There are only a few isolated teeth. One preserves a
surface is formed by the surangular and articular but the suture long, straight, and flattened root; the poorly preserved crown
between these bones is not evident; presumably its course was of this tooth is continuous with the root without a constriction
similar to that in Dorosuchus neoetus (Sookias et al., 2014a). between them (Fig. 4B). The labiolingually narrow tooth
The two cotyles for the articular condyles of the quadrate are crowns are proportionately small and only weakly recurved.
not clearly separated by a low ridge, unlike in Euparkeria capen­ None of the crowns is sufficiently well preserved to determine
sis (Sookias et al., 2020). The lateral and medial margins of the the presence or absence of serrations on the mesial and distal
glenoid surface are rounded. cutting edges.
Angular—Part of the dorsal surface of the angular is exposed
medial to the anterior process of the surangular (Fig. 6A, B) but
Postcranial Skeleton
it presents no noteworthy features.
Articular—The left articular is exposed in dorsal view (Fig. 6A, Vertebral Column—The postcranial axial skeleton is rep­
B). It forms much of the glenoid surface. Its posteromedial resented only by mostly dissociated vertebrae, ribs, and gastralia.
portion forms a ventromedially directed process, as in Dorosu­ Often obliquely embedded in the matrix and affected by crushing
chus neoetus and pseudosuchians (Sookias et al., 2014a). The and distortion during fossilization, it is difficult to determine the
posterior portion of this process is directed posteromedially relative positions and document the structure of some vertebrae.
and somewhat dorsally. Behind the lateral portion of the No intercentra are preserved, but this might reflect the disarticu­
glenoid surface, is a deeper, mediolaterally expanded fossa, lated nature of the skeletal remains. None of the vertebral centra
which Sookias et al. (2014a) named the postcotylar fossa. The presents distinct ventral beveling of the rims of the articular sur­
articular has a ventromedial process, as in D. neoetus (Sookias faces that is frequently associated with the presence of
 Sues et al.—Middle Triassic archosauriform (e2357326-8)

FIGURE 6. Marcianosuchus angustifrons

(SMNS 91318, holotype). A, B, photograph
and explanatory diagram of left postdentary
bones in dorsal view. C, left prearticular in
lateral view. Abbreviations: afm, articular
facet for craniomandibular joint; an, angular;
ar, articular; dp, dorsal process (base only);
mc, medial wall of Meckelian canal; mps,
medial process of surangular; pcf, postcondylar
fossa; rp, retroarticular process; sa, surangular;
vmp, ventromedial process. Scale bars equal
1 cm.

intercentra. However, Halazhaisuchus qiaoensis has well-devel­ over the neural canal and a short, posteriorly directed and taper­
oped intercentra yet lacks distinct ventral beveling of the ing process that met the prezygapophysis of the axis. The anterior
anterior and posterior articular ends of its presacral vertebral part of the dorsal region has a well-defined facet that is ventrally
centra (Sookias et al., 2014b). bordered by a narrow ledge and contacted the proatlas. The
The presacral vertebrae have gently amphicoelous, spool- ventral pedicle of the neural arch has a slightly thickened end
shaped, and non-notochordal centra, which are constricted at for contact with the atlas intercentrum and an anterior part for
mid-length with concave lateral and ventral surfaces between contact with the occipital condyle.
the articular surfaces. None of the centra is anteroposteriorly One cervical vertebra, exposed in obliquely anterolateral view
elongated. The articular surfaces of the centra are moderately (Fig. 7B), has a small, knob-like diapophysis just below the neu­
concave and form distinct rims. In lateral view, the neural rocentral suture and a large parapophysis in an anteroventral
arches are slightly embayed at the bases anteriorly but more position on the rim of the anterior centrum surface. Its neural
deeply posteriorly. Where clearly exposed, the neurocentral spine has a transversely wide distal apex (‘spine table’) with a dis­
sutures are unfused. tinctly concave dorsal surface. The lateral edges of the ‘spine
Both neural arches of the atlas are preserved (Fig. 7A). The table’ form lobes in lateral view. We interpret this vertebra as a
right element is exposed in lateral view. Its dorsal region is more posterior cervical. There is a shallow spinodiapophyseal
divided into a portion that, together with its antimere, ‘roofed’ fossa on the lateral surface of the neural arch. The apex of the
 Sues et al.—Middle Triassic archosauriform (e2357326-9)

FIGURE 7. Marcianosuchus angustifrons

(SMNS 91318, holotype). A, atlantal neural
arches, the right one in lateral view. B, mid-cer­
vical vertebra in oblique anterolateral view.
Abbreviations: di, diapophysis; foc, facet for
occipital condyle; fpa, area for contact with
proatlas; ped, pedicle of neural arch; pp, para­
pophysis; ppr, posterior process of atlantal
neural arch; prz, prezygapophysis; pz, postzy­
gapophysis; sd, Spinodiapophyseal fossa; spt,
‘spine table.’ Scale bars equal 5 mm (A) and
1 cm (B).

neural spine resembles the Y-shaped ‘spine tables’ on the presa­ processes and a low lateral flange extending distally from the
cral and anterior caudal vertebrae of Tarjadia ruthae (Ezcurra tuberculum along the anterior surface of the rib shaft. A more
et al., 2017), the mid-cervical and anterior dorsal vertebrae of posteriorly positioned dorsal rib has a well-developed capitular
Aenigmaspina pantyffynnonensis (Patrick et al., 2019), and the process but the tuberculum lacks a distinct process. A partial
mid-cervical vertebrae of Scolotosuchus basileus (Sennikov, shaft of a dorsal rib preserves its slightly expanded distal end.
2022). The transversely expanded apices on the neural spines Gastralia—The gastralia are mostly dissociated. Four median
resemble those on cervical vertebrae in birds, which are associ­ elements are preserved still largely in association near the
ated with the ligamentum supraspinosum and ligamentum elasti­ distal end of the right femur. Each has straight, slender shafts
cum interspinale and median epaxial muscles (Tsuihiji, 2004). that are fused to each other along the midline of the body and
The prezygapophysis on the right side of the neural arch has an form an open ‘V’ with its apex pointing forward. The anterior
articular facet that faces anteromedially and dorsally and the gastralium of the set has shafts diverging from each other at a
postzygapophysis on the left side has a slightly elongated, poster­ right angle and bears an anterior process. A large V-shaped
oventrally and laterally facing articular facet. On all identifiable gastral element is exposed adjacent to the left surangular.
cervical vertebrae, the anterior and posterior articular surfaces of Various scattered fragments of rod-like bones possibly represent
the centrum are at the same level, rather than offset from each more lateral segments of gastralia.
other, in lateral view. Osteoderms—Specimen SMNS 91318 includes some scattered
The dorsal vertebrae (Fig. 8) are distinguished by the place­ osteoderms (Fig. 10), none of which are preserved in their articu­
ment of diapophyseal facets on slender transverse processes. lar context. We interpret them as dorsal paramedian elements.
Anterior (= parapophyseal) and posterior centrodiapophyseal Each osteoderm is more or less rectangular in outline, with a
laminae extend from the anterior and posterior ends of the rounded anterior and a slightly concave posterior margin. The
centrum, respectively, along the ventral surface of the transverse plates lack anterior or anterolateral processes, unlike in Eupar­
processes to the tips of the processes. Just below the neural keria capensis (Ewer, 1965) and its close relatives (Sookias
arches, two still articulated dorsal centra have round lateral et al., 2014b) as well as in various pseudosuchians (Nesbitt,
fossae (Fig. 8). The prezygapophyseal facets face dorsomedially 2011). Complete osteoderms are approximately twice as long
and those of the postzygapophyses ventrolaterally. The facets anteroposteriorly as wide mediolaterally. Two thin plates
are separated from each other medially by narrow clefts. Some (Fig. 10A) each bear a distinct, somewhat asymmetrically
of the zygapophyseal facets are flat mediolaterally whereas the placed, and anteroposteriorly extending ridge or keel on their
prezygapophyses on one dorsal are slightly convex in anterior dorsal surfaces, which we interpret as laterally offset based on
view. The centra of the mid-dorsal and posterior dorsal vertebrae comparisons with similar features on the osteoderms of other
are anteroposteriorly short. The ventral surfaces of the centra of archosauriforms (although we cannot rule out the possibility
the uncrushed dorsals are rounded transversely. The two articu­ that the ridge was medially offset as in Halazhaisuchus qiaoensis;
lated centra are crushed so that the ventral margins appear sharp. Sookias et al., 2014b). Two thicker, strongly sculptured plates
The neural spines either have slightly transversely expanded lack such a ridge (Fig. 10B), but their sculptured posterior
apices or are somewhat elongated and not transversely expanded portion forms a low, mound-like dorsal eminence. They show
distally. some resemblance to the osteoderms on the dorsal surface of
A vertebra with a tall, anteroposteriorly narrow neural spine, a the tail in Ticinosuchus ferox (Krebs, 1965). On all osteoderms,
dorsoventrally lower neural arch, and the base of a small caudal much of the dorsal surface is ornamented with ridges and distinct
rib is probably an anterior caudal (Fig. 9A). A more posteriorly pits, which radiate from the dorsal ridge or eminence. The pits
situated caudal vertebra lacks any trace of a caudal rib (Fig. 9B). become more elongate toward and open along the periphery of
Its pre- and postzygapophysis are connected by a lateral ridge. the plate. The anterior portion of the dorsal surface lacks sculp­
The prezygapophysis extends anterior to the anterior end of turing and was presumably overlapped by the preceding osteo­
the centrum whereas the postzygapophysis barely does so rela­ derm in life. This contrasts with the smoother dorsal surfaces
tive to the posterior end. of the osteoderms in E. capensis (Ewer, 1965). The slightly med­
Ribs—The preserved dorsal ribs have moderately expanded iolaterally concave ventral sides of the osteoderms are smooth
proximal ends with a short tubercular process and a long, and featureless. The lateral and medial margins of the plates
slender capitular process. Their shafts are gently curved. An are slightly irregular. The shape of the osteoderms suggests
anterior dorsal rib has clearly separated capitular and tubercular that the dorsal armor comprised a pair of paramedian rows.
 Sues et al.—Middle Triassic archosauriform (e2357326-10)

FIGURE 8. Marcianosuchus angustifrons

(SMNS 91318, holotype). Cluster of dorsal ver­
tebrae as preserved. Abbreviations: cd,
depression on lateral surface of centrum; ns,
neural spine; prz, prezygapophysis; pz, postzy­
gapophysis; trp, transverse process. Scale bar
equals 1 cm.

Furthermore, the relative sizes of the plates and the ‘spine tables’ Scapula—Neither scapula preserves a complete blade but
on the presacral vertebrae suggest a one-to-one alignment of that of the left element is more complete (although a
these two features, a common character-state among Archosaur­ section of it was subsequently removed to expose the articular
iformes (Ezcurra, 2016; Nesbitt, 2011). portion of the left mandibular ramus; Fig. 11). The right
Elsewhere, paired rows of dorsal osteoderms are present in element is exposed in lateral view and the left in medial
Euparkeriidae (Ewer, 1965; Sookias et al., 2014b; Sookias, view. Enough of the scapulae is preserved to show that the
2016), phytosaurs (Westphal, 1970), and various pseudosuchians length of the scapula exceeded 75% of the length of the
(Nesbitt, 2011). Proterochampsids have only a single row of humerus (Nesbitt, 2011). The scapular blade is tall dorsoven­
dorsal armor elements (Trotteyn et al., 2013). Unlike those of trally, narrow anteroposteriorly for much of its height, and
Marcianosuchus angustifrons, the osteoderms of Euparkeriidae mediolaterally flattened. Its preserved portion has rather
(Ewer, 1965; Sookias et al., 2014b), Gracilisuchidae (Lecuona straight, nearly parallel anterior and posterior margins and
et al., 2017), and some loricatans (Gower & Schoch, 2009) all only slightly expands anteroposteriorly more distally;
have tapering anterior processes that fit under on the ventral however, the distal (= dorsal) end of the blade is not pre­
surface of the preceding plates. The unsculptured anterior served on either scapula and is possibly expanded as in
region of the plates of M. angustifrons resembles that on dorsal Euparkeria capensis (Ezcurra, 2016). The right scapula is
osteoderms in doswelliid archosauriforms such as Doswellia kal­ slightly constricted just distal to its proximal portion, which
tenbachi (Dilkes & Sues, 2009) and Jaxtasuchus salomoni is widest anteroposteriorly along its contact with the coracoid.
(Schoch & Sues, 2014), but there is no raised anterior ridge Posteroventrally, the scapula bears a posterolaterally and
and the anterior and posterior regions are not separated by a slightly ventrally facing glenoid facet. Just dorsal to this
groove. facet, the posterior edge of the right scapula is thickened
transversely. Its lateral surface bears an abraded feature in
the same position as the tubercle for the origin of

FIGURE 9. Marcianosuchus angustifrons (SMNS 91318, holotype).

Caudal vertebrae. A, anterior caudal vertebra in posterolateral view. B, FIGURE 10. Marcianosuchus angustifrons (SMNS 91318, holotype). A,
mid-caudal vertebra in lateral view. Abbreviations: cr, caudal rib (proxi­ B, two types of dorsal osteoderms. Arrows point anteriorly. Abbrevi­
mal portion only); ncs, neurocentral suture; ns, neural spine; prz, prezyga­ ations: em, dorsal eminence; oz, zone overlapped by preceding osteo­
pophysis; pz, postzygapophysis. Scale bars each equal 1 cm. derm; r, dorsal ridge. Scale bars each equal 5 mm.
 Sues et al.—Middle Triassic archosauriform (e2357326-11)

FIGURE 11. Marcianosuchus angustifrons

(SMNS 91318, holotype). Scapulae and associ­
ated gastralia. The left scapula is exposed in
medial view whereas the right scapula (at the
top of the image) is exposed in lateral view.
Abbreviations: acr, acromial process; gl,
glenoid facet of scapula; gr, gastralium; itm,
insertion of M. triceps lateralis longus. Scale
bar equals 1 cm.

M. triceps as in many other archosauriforms, including Eupar­ ventral surface of the proximal region on the right humerus.
keria capensis and Halazhaisuchus qiaonensis (Sookias et al., The asymmetrical distal end shows little detail on either
2014b) and crocodylians (attachment for M. triceps longus humerus. There is no trace of an ectepicondylar groove and a
lateralis; Meers, 2003). Anterior to the glenoid facet and pos­ supinator process.
terior to the acromial margin, the lateral surface of the Ilium—A fragment of a robust bone near the long edge of
scapula is concave. The distinct acromial region is not raised the sandstone slab is a right ilium lacking its posterior portion
above its ventral margin. The ventral margin of the scapula (Fig. 13). It was obliquely embedded, initially visible only in
is convex. medial view, and distorted, rendering its interpretation diffi­
Coracoid—The left coracoid is exposed in lateral view (Fig. cult. During preparation, both sides of the bone were
12A). It is smaller than the scapula and plate-like. The cora­ exposed. The medial surface of the ilium bears a prominent
coid is longer anteroposteriorly than deep dorsoventrally, with ridge posteriorly, which presumably continued posteriorly
a nearly oval outline in lateral view as in other non-archosaur­ onto the postacetabular process, as in Dorosuchus neoetus
ian archosauriforms (Ezcurra 2016; Nesbitt, 2011). Posteriorly, (Sookias et al., 2014a). No facets for attachment of the
its slightly convex lateral surface bears the dorsoventrally flat­ sacral ribs can be clearly delineated. The imperforate acetabu­
tened ventral ‘lip’ of the posterolaterally facing glenoid facet. lum lacks a supra-acetabular crest and a supra-acetabular but­
Anterior to the glenoid facet and separated from it by a slight tress. Although it is incompletely preserved, the preacetabular
ridge, a dorsoventrally slightly elongated coracoid foramen process of the ilium appears to be short.
opens just below the dorsal margin of the element. The Ischium—An incomplete right ischium lacking its distal end
short postglenoid portion of the coracoid has a rounded pos­ is exposed in medial view near the distal end of the left femur
terior edge. (Fig. 14A). Its plate-like proximal portion has a short, nearly
Humerus—The humerus is represented by the nearly com­ anteroposteriorly extending dorsal and a dorsoventrally deep
plete left bone exposed in dorsal view (Fig. 12B) and much of anterior margin, which presumably contacted the ilium and
the right element (except for the proximal end) exposed in pubis, respectively. The dorsal and ventral edges of the
ventral view on a separate small piece of sandstone (Fig. 12C). ischium converge posteriorly and form a slender, mediolater­
The left element has a length of 67 mm. Its proximal and ally narrow shaft. The medial surface is gently convex dorso­
especially distal ends are expanded relative to the width of the ventrally along the preserved portion of the ischium. The
mid-shaft and flattened (although these features are likely shaft-like distal region of the right ischium bears a longitudi­
affected to some extent by postmortem crushing). The head nal ridge, which extends closer to the dorsal margin. This
and internal tuberosity are indistinct. The shaft is slender. The ridge probably marks the dorsal margin of the symphyseal
moderately developed, medially projecting deltopectoral crest area for contact with the left bone. The ischium differs from
(for the insertion of M. pectoralis and M. deltoideus clavicularis; those of Proterosuchus fergusi (Ezcurra, 2016), Erythrosuchus
Meers, 2003) extends from the lateral margin of the proximal end africanus (Gower, 2003), and Euparkeria capensis (Demuth
distally to about mid-length of the humerus where it merges with et al., 2020) in having a rod-like distal shaft rather than
the shaft. Due to postmortem crushing, it was folded over the being a plate-like bone. This condition resembles that in
 Sues et al.—Middle Triassic archosauriform (e2357326-12)

FIGURE 13. Marcianosuchus angustifrons (SMNS 91318, holotype).

Fragment of right ilium in A, medial and B, oblique dorsolateral view.
Abbreviations: ac, acetabulum; mr, medial ridge; pup, pubic process of
ilium. Scale bar equals 1 cm.

having a flat distal portion without a thickened lateral margin

and being less twisted.
Femur—Both femora are preserved in SMNS 91318. The left
bone is crushed flat and distorted with considerable loss of
bone at both articular ends. The right femur is exposed in poster­
olateral view and much better preserved, lacking only its distal
articular end (Fig. 15A). It closely resembles the femora of Dor­
osuchus neoetus (Sookias et al., 2014a) and Euparkeria capensis
(Ewer, 1965) in the slightly sigmoidal curvature of its slender
shaft, which differs from the more pronounced, smoothly sig­
moidally curved femoral shaft in pseudosuchians such as Batra­
chotomus kupferzellensis (Gower & Schoch, 2009). The poorly
preserved proximal end of the right femur extends anterome­
dially and distally smoothly grades into the shaft. The proximal
and distal articular ends are expanded relative to the shaft and
offset from each other, but, due to crushing and the loss of the
distal condyles, this offset cannot be quantified. The proximal
articular surface of the femur is placed on the proximal end.
On the ventromedial aspect of the proximal third of the femur,
a blade-like crest extends proximodistally and forms a fairly sym­
metrical fourth trochanter for the insertion of M. caudifemoralis
longus (Romer, 1923). The trochanter is not continuous with the
proximal end of the femur and extends parallel to the long axis of
the bone. Its lateral surface is slightly concave. Further distally, a
second ridge, for the insertion of M. adductor femoris (Romer,
1923), extends distally along the margin of the femoral shaft
where the lateral and ventral surfaces meet. Proximally, it
forms a slight lateral eminence on the ventrolateral edge of the
shaft.
FIGURE 12. Marcianosuchus angustifrons (SMNS 91318, holotype). A, The femur is distinctly longer than the humerus (c. 115 mm
left coracoid in lateral view. B, left humerus in dorsal view. C, right [right femur] vs. 67 mm [left humerus]), proportionately more
humerus (missing its proximal end) in ventral view. Abbreviations: cf,
coracoid foramen; dc, deltopectoral crest; hh, proximal head of
humerus; vgl, ventral ‘lip’ of glenoid facet. Scale bars each equal 1 cm.

pseudosuchians such as Batrachotomus kupferzellensis (Gower

& Schoch, 2009) and Arizonasaurus babbitti (Nesbitt, 2005).
Pubis—The right pubis is crushed and has lost most of its bony
substance, but the left pubis is better preserved and exposed in
oblique posterior view (Fig. 14B). Its proximal portion is
narrow mediolaterally. Little of this region is preserved on
either element. More distally, the pubis is expanded mediolater­
ally into a thin bony plate that met its antimere along the midline,
forming a transversely narrow, elongate pubic ‘apron.’ This FIGURE 14. Marcianosuchus angustifrons (SMNS 91318, holotype). A,
‘apron’ slightly increases in transverse width distally. The pubis right ischium in medial view. B, left pubis in oblique posterior view.
differs from that of Euparkeria capensis (Ewer, 1965) by Abbreviation: r, ridge. Scale bars each equal 1 cm.
 Sues et al.—Middle Triassic archosauriform (e2357326-13)

FIGURE 15. Marcianosuchus angustifrons

(SMNS 91318, holotype). A, right femur in
ventral view. B, partial tibia (side and view
uncertain). C, partial ?left ?fibula. Abbrevi­
ations: ad, adductor ridge on femur; em, emi­
nence on femur; ft, fourth trochanter of
femur; hf, head of femur; ?ilf, possible insertion
of M. iliofibularis. Scale bars each equal 2 cm.

so than in Euparkeria capensis (Ewer, 1965). It is clearly PHYLOGENETIC ANALYSIS

shorter than the skull because the estimated length of the
Marcianosuchus angustifrons is referred to Archosauriformes
lower jaw is c. 190 mm, based on the size of the postdentary
based on the presence of the antorbital fenestra. It shares the
bone complex and assuming cranial proportions comparable
possession of interdental plates with other archosauriforms.
to those for Euparkeria capensis (Sookias et al., 2020).
Finally, M. angustifrons shares the presence of paired rows of
Tibia—Much of a tibia is preserved, but little detail is evident
paramedian osteoderms dorsal to the vertebral column with
due to the loss of its distal articular end and its poor state of pres­
Euparkeria capensis and its close relatives (Sookias, 2016;
ervation (Fig. 15B). The length of the fragment is 95 mm. We
Sookias et al., 2014b) as well as with phytosaurs and most pseu­
could not determine whether it represents the left or right dosuchians (Nesbitt, 2011). Some proterochampsids have a
element. Its proximal end is convex and expanded. The shaft is single row of dorsal osteoderms (e.g., Chanaresuchus bonapartei;
slender and nearly straight. Although incomplete distally, the Trotteyn et al., 2013), and certain pseudosuchians have more
tibia was certainly shorter than the femur based on comparisons than one pair of osteoderms per vertebra (e.g., Ticinosuchus
with tibiae of related archosauriforms. This character-state is the ferox; Krebs, 1965). Sookias (2016) and Sookias et al. (2014a,
typical condition for non-archosaurian archosauriforms such as 2020) enumerated the following diagnostic features for Eupar­
Euparkeria capensis (Ewer, 1965) as well as for most pseudosu­ keriidae and its constituent taxa, but most of these character-
chians, except for some non-crocodyliform crocodylomorphs states concern features that are not preserved in SMNS 91318
(Nesbitt, 2011). (with observable states in SMNS 91318 indicated in parentheses):
Fibula—We tentatively identify a fragment of a long, four premaxillary teeth (present); anterolateral surface of
slender, and slightly curved bone lacking its distal end as a maxilla with large, anterolaterally opening foramen (not obser­
left fibula. Its length as preserved is 75 mm. The dorsal vable); exoccipital with a single foramen for CN XII (not obser­
surface of the proximal portion bears a longitudinal ridge, vable); presence of postparietals (not observable); and anterior
possibly for attachment for M. iliofibularis (Romer, 1923), edge of presacral osteoderms with distinct anterior process
which extends closer to the proximal articular end rather (absent).
than more distally. A similar crest is present on the fibula The phylogenetic analysis found 32 most parsimonious trees
of the early-diverging pseudosuchian Nundasuchus songeaensis (MPTs), each with 1356 steps, a consistency index of 0.35, and
(Nesbitt et al., 2014). Nesbitt (2011) considered this position a retention index of 0.68 (Supplementary File 3; for a tree of
of the ridge plesiomorphic for archosauriforms. an analysis with implied weights see Supplementary File 4).
Manus or Pes—Various autopodial bones are preserved but, in The best score was hit 711 times out of the 1000 iterations. In
the absence of articulated or associated manus and pedes, their the strict consensus tree (SCT) of all MPTs (Fig. 16),
specific identities cannot be determined. M. angustifrons was recovered in a polytomy that also comprises
 Sues et al.—Middle Triassic archosauriform (e2357326-14)

FIGURE 16. Strict consensus tree of 32 most parsimonious trees recovered by the TNT analysis of the character-taxon matrix compiled by Sookias
(2016) with the addition of scores for the erpetosuchid Tarjadia ruthae, each with 1333 steps, a consistency index of 0.36, and a retention index of 0.69.
Bootstrap GC frequencies above 50% are listed above and Bremer supports >1 below each branch. The binomen Marcianosuchus angustifrons is
highlighted in bold.
 Sues et al.—Middle Triassic archosauriform (e2357326-15)

FIGURE 17. Marcianosuchus angustifrons (SMNS 91318, holotype). Body silhouette with preserved cranial and appendicular elements and selected
vertebrae and osteoderms. The positions of gastralia, ribs, and vertebrae are conjectural. Silhouette based on the skeletal reconstruction of Euparkeria
capensis by Demuth et al. (2023).

Dorosuchus neoetus, Phytosauria, Euparkeriidae, and Archo­ comparatively short mid- to posterior dorsal vertebrae, the pro­
sauria. This large clade is defined by the following synapomor­ portional differences between the femoral and humeral length
phies (those observable in SMNS 91318 marked by *, both and the femoral and skull length, respectively, and the structure
here and below): foramen on articular present medial to of the dorsal paramedian osteoderms, particularly the absence of
glenoid* (character 150: 0 → 1), distal end of cervical neural an anterior process and the presence of pronounced sculpturing
spines expanded in the middle of its anteroposterior length* on the dorsal surface. Marcianosuchus angustifrons more closely
(character 183: 0 → 1), distal end of dorsal neural spines resembles phytosaurs than euparkeriids in the structure of its
expanded with flat dorsal margin* (character 192: 0 → 1), distal osteoderms and the overlap of successive median osteoderms.
end of fibula angled anterodorsally (asymmetrical) in lateral However, phytosaurian osteoderms differ in being typically not
view (character 336: 1 → 0), posterior corner of the dorsolateral rectangular and having much more pronounced sculpturing on
margin of the astragalus overlaps the calcaneum much more than their dorsal surfaces (Westphal, 1970). Furthermore,
the anterior portion (character 351: 0 → 1), and calcaneal tuber M. angustifrons lacks diagnostic phytosaurian synapomorphies
as broad as tall or somewhat broader than tall (character 367: 0 including: premaxilla longer than maxilla and with six or more
→ 1). In 16 out of 32 MPTs, M. angustifrons was recovered tooth positions; maxillary portion anterior to antorbital fenestra
within Euparkeriidae as the sister taxon to all remaining eupar­ as long or longer than maxillary part posterior to it; maxilla
keriids, which include E. capensis, Osmolskina czatkowicensis, extending anteriorly to nasal; subtriangular quadratojugal; and
Halazhaisuchus qiaoensis, and “Turfanosuchus shageduensis” external nares positioned far posterior to anterior end of the
(Sookias et al., 2014a, b). In these MPTs, Euparkeriidae includ­ snout and directed dorsally (Stocker & Butler, 2013). Our
ing M. angustifrons possesses the following synapomorphies: pre­ results indicate that M. angustifrons presents a distinct, novel
maxilla with four tooth positions* (character 6: 2 → 1; present in body plan (Fig. 17) that among well-known archosauriforms
all trees); presacral osteoderms longer than wide* (character 400: most closely resembles that of E. capensis. The length disparity
0 → 1; present in some trees). In the remaining 16 MPTs, between its humerus and its femur is greater than in the latter
M. angustifrons is found within Phytosauria as the sister taxon taxon, and the short dorsal vertebrae and the proportionately
to all remaining phytosaurs, a clade that in this analysis is rep­ large skull make it likely that M. angustifrons was predominantly
resented by Parasuchus hislopi, Machaeroprosopus pristinus, quadrupedal, as has been inferred for E. capensis (Demuth et al.,
and Smilosuchus gregorii. The analysis found a single synapo­ 2023).
morphy for a clade Phytosauria + M. angustifrons: osteoderms
with pits of irregular size and contour* (character 397: 0 → 1).
The autapomorphies for M. angustifrons common to all MPTs DISCUSSION
are (1) squamosal with distinct (lateral) ridge on dorsal surface Marcianosuchus angustifrons is noteworthy as the first defini­
along edge of supratemporal fossa (character 55: 0 → 1) and tive record of a non-archosaurian archosauriform from the Bunt­
(2) osteoderms with distinct dorsal eminence or ridge (character sandstein Group of Germany. Until now, archosauriform reptiles
399: 0 → 1). In addition, the following autapomorphies for were documented by only a few archosaurian skeletal remains
M. angustifrons are present in the MPTs in which it is recovered from the Middle and Upper Buntsandstein subgroups. The
as a euparkeriid: mid- to posterior dorsal vertebral centra taller ‘sail-backed’ poposauroid pseudosuchian Ctenosauriscus
than long (character 195: 1 → 0) and osteoderms with incised koeneni is known from much of a vertebral column from the
and pitted ornamentation of different size and contour (charac­ upper Olenekian Solling Formation (Middle Buntsandstein Sub­
ter 397: 0 → 1). group) of the Bremketal near Göttingen in Lower Saxony
Tarjadia ruthae, the taxon that was added by us to the charac­ (Butler et al., 2011; Huene, 1914; Krebs, 1969). More recently,
ter-taxon matrix used by Sookias (2016), was consistently recov­ postcranial remains of a possibly congeneric poposauroid were
ered as the sister taxon to Aetosauriformes among non-suchian reported from the lower Anisian Röt Formation (Upper Bunt­
pseudosuchians. This deviates somewhat from recent interpret­ sandstein Subgroup) of the Waldshut region in Baden-Württem­
ations of the phylogenetic position of T. ruthae and other erpeto­ berg (Butler et al., 2011). Ctenosauriscus koeneni is readily
suchids, which recover Erpetosuchidae as the sister group to distinguished from M. angustifrons by its greatly elongated and
Ornithosuchidae (Ezcurra et al., 2017; Foffa et al., 2020). Never­ distally strongly expanded neural spines on the posterior cervi­
theless, its addition excludes an erpetosuchid interpretation of cal, dorsal, sacral, and anterior caudal vertebrae, which form a
M. angustifrons based on this matrix. Instead, M. angustifrons symmetrical dorsal ‘sail’ (Butler et al., 2011). Furthermore, the
is recovered as a non-archosaurian archosauriform that is more large pre- and postzygapophyses of its dorsal vertebrae extend
closely related to archosaurs than to Proterosuchidae, Erythrosu­ well beyond the articular faces of the centra. Ctenosauriscus
chidae, Proterochampsia, and Doswelliidae. Marcianosuchus koeneni is closely related to Arizonasaurus babbitti from the
angustifrons is most notably distinguished from E. capensis by Anisian-age Holbrook Member of the Moenkopi Formation of
the presence of a distinct lateral ridge on the squamosal, the Arizona (Nesbitt, 2005), Hypselorhachis mirabilis from the
 Sues et al.—Middle Triassic archosauriform (e2357326-16)

Anisian-age Lifua Member of the Manda Beds of Tanzania Russian localities and the apparent absence of non-mammalian
(Butler et al., 2009), and Xilousuchus sapingensis from the Ole­ synapsids from the Lower and lower Middle Triassic of Central
nekian-age Heshanggou Formation of north-central China Europe possibly reflect the different paleolatitudinal positions
(Nesbitt et al., 2011). The clade Ctenosauriscidae (as defined of their occurrences (Romano et al., 2020; Schneider et al.,
by Butler et al. [2011]) includes the stratigraphically oldest arch­ 2020) and perhaps more favorable environmental conditions in
osaurs known to date. Russia during the Early and early Middle Triassic.
Two archosauriform taxa, Seemannia palaeotriadica Huene,
1958 and Crenelosaurus nigrosilvanus Ortlam, 1967, were each
CONCLUSION
named on the basis of a single, poorly preserved tooth crown
from the Upper Buntsandstein Subgroup in the Black Forest Marcianosuchus angustifrons is the first non-archosaurian
region of Baden-Württemberg. Both were originally assigned archosauriform from the Upper Buntsandstein Subgroup
to Proterosuchidae, but they lack any diagnostic features and (Middle Triassic: Anisian) of Germany. It presents a novel
should be treated as nomina dubia. Krebs (1969) reported an uni­ body plan that most closely resembles that of Euparkeria capen­
dentified tooth from the upper Middle Buntsandstein near Göt­ sis among well-known non-archosaurian archosauriforms. Like
tingen in Lower Saxony. the latter taxon and Dorosuchus neoetus, it presents a skeletal
Finally, there is an intriguing record of an incomplete cranium structure that is very similar to the plesiomorphic condition of
of a phytosaur, “Mesorhinus” fraasi, which reportedly came from archosaurs (Sookias et al., 2014a).
the basal layers of the Middle Buntsandstein Subgroup (probably Marcianosuchus angustifrons joins the ‘sail-backed’ poposaur­
Volpriehausen Formation) at Bernburg in Saxony-Anhalt oid pseudosuchian Ctenosauriscus koeneni as well as the eupar­
(Jaekel, 1910). Kuhn (1961) noted that the generic name Mesor­ keriid Osmolskina czatkowiciensis (Borsuk-Białynicka &
hinus Jaekel, 1910 was pre-occupied by Mesorhinus Ameghino, Evans, 2003, 2009) and the poorly known archosauriform Colli­
1885 (Mammalia: Litopterna) and proposed Mesorhinosuchus longus rarus (Borsuk-Białynicka & Sennikov, 2009), from Olene­
as its replacement. Unfortunately, the holotype and only kian-age cave fillings at Czatkowice, northwest of Kraków
known specimen of M. fraasi was lost during World War II. (Poland), in documenting the presence of several archosauriform
Despite Jaekel’s efforts to verify its provenance, the geological clades in conterminous Central Europe only a few million years
age of this fossil has been questioned ever since (e.g., Stocker after the end-Permian biotic crisis.
& Butler, 2013). Based on Jaekel’s (1910:fig. 2) photograph, the
cranium definitely belonged to a ‘Paleorhinus-grade’ phytosaur
based on the dorsal position of the external nares on the
DISCLOSURE STATEMENT
rostrum anterior to the antorbital fenestrae.
Unlike in Central Europe, several Olenekian- and Anisian-age No potential conflict of interest was reported by the author(s).
non-archosaurian archosauriform and archosaurian reptiles are
known from the Eastern European Platform and Cis-Uralian
ACKNOWLEDGMENTS
region of Russia (Sennikov, 1995). However, assessing the phylo­
genetic relationships of many of these taxa is difficult because We thank I. Rosin for her skillful preparation of the specimen
they are frequently based on isolated, typically fragmentary and E. Mujal for producing the three-dimensional photogram­
bones, mainly vertebrae (Gower & Sennikov, 2000; Shishkin metric model of the fossil. R. B. Sookias generously provided
et al., 2024). Non-archosaurian archosauriforms from European photographs of specimens of Dorosuchus neoetus and Eupar­
Russia include the chasmatosuchine proterosuchids Chasmatosu­ keria capensis and O. Damuth kindly shared 3D renderings of
chus rossicus from the Rybinskian Gorizont, Gamosaurus the pelvis of the latter taxon. We are indebted to
lozovskii and Jaikosuchus magnus from the Gamian Gorizont, M. D. Ezcurra, C. T. Griffin, and an anonymous reviewer as
Vonhuenia friedrichi from the Vokhminian Gorizont, and Tsyl­ well as Editor G. Bever and Phylogenetics Editor P. Godoy for
mosuchus jakovlevi from the Ustmylian Gorizont (Sennikov, their helpful comments and suggestions. H-DS gratefully
1995); the non-eucrocopodan archosauriform Sarmatosuchus acknowledges support from the MacMillan Fund of the Depart­
otschevi from the Donguz Gorizont (Gower & Sennikov, 1997), ment of Paleobiology. The software TNT is made freely available
and the erythrosuchids Garjainia prima from the Fedorovkian by the Willi Hennig Society.
and Gamian gorizonts (Butler et al., 2019; Ezcurra et al., 2019;
Maidment et al., 2020) and Uralosaurus magnus from the
DATA AVAILABILITY STATEMENT
Donguz Gorizont (Sennikov, 1995). More crownward archo­
sauriforms include the Euparkeria-grade archosauriform Doro­ The data that support the conclusions of this study are avail­
suchus neoetus from the Donguz Gorizont (Sookias et al., able in the manuscript.
2014a); the ctenosauriscid pseudosuchian Bystrowisuchus
flerovi from the Gamian Gorizont (Sennikov, 2012); the possible
AUTHOR CONTRIBUTIONS
pseudosuchians Scolotosuchus basileus from the Gamian Gori­
zont (Sennikov, 2022) and Vjushkovisaurus berdjanensis from RRS and H-DS designed the project. RRS wrote an initial
the Donguz Gorizont (Ochev, 1982); the possible aphanosaurian draft. H-DS prepared the anatomical descriptions and he and
avemetatarsalian Vytshegdosuchus zheshartensis from the SNFS created the figures. H-DS and RRS worked with SNFS
Gamian Gorizont (Ezcurra et al., 2023); and the aphanosaurian on the phylogenetic analysis. All authors edited various drafts
Dongusuchus efremovi from the Donguz Gorizont (Nesbitt of the manuscript and approved the final version.
et al., 2017; Niedźwiedzki et al., 2016). The age of the Vokhmi­
nian Gorizont is Induan (Shishkin et al., 2024). The Rybinskian
SUPPLEMENTARY FILES
Gorizont has been dated as early Olenekian, the Sludkian Gori­
zont is slightly younger than the Rybinskian Gorizont, and the Supplementary File 1.nex: Character-taxon matrix with the
Gamian Gorizont is considered late Olenekian in age (Shishkin addition of Marcianosuchus angustifrons and Tarjadia ruthae in
et al., 2024). The Donguz Gorizont has been dated as Anisian NEXUS format.
(Shishkin et al., 2024). The considerable differences in faunal Supplementary File 2.tnt: Character-taxon matrix with the
composition between the Early and early Middle Triassic age addition of Marcianosuchus angustifrons and Tarjadia ruthae in
assemblages of archosauriform reptiles from the German and TNT format.
 Sues et al.—Middle Triassic archosauriform (e2357326-17)

Supplementary File 3.tre: Most parsimonious trees. Ezcurra, M. D. (2016). The phylogenetic relationships of basal archosaur­
Supplementary File 4.tiff: Tree from phylogenetic analysis with omorphs, with an emphasis on the systematics of proterosuchian
implied weights. archosauriforms. PeerJ, 4, e1778. https://doi.org/10.7717/peerj.1778
Ezcurra, M. D., Bandyopadhyay, S., Sengupta, D. P., Sen, K., Sennikov, A.
G., Sookias, R. B., Nesbitt, S. J., & Butler, R. J. (2023). A new arch­
ORCID osauriform species from the Panchet Formation of India and the
diversification of Proterosuchidae after the end-Permian mass
Hans-Dieter Sues http://orcid.org/0000-0002-9911-8254 extinction. Royal Society Open Science, 10(10), 230387. https://doi.
Stephan N. F. Spiekman http://orcid.org/0000-0002-3197-8752 org/10.1098/rsos.230387
Rainer R. Schoch http://orcid.org/0000-0002-0312-2877 Ezcurra, M. D., & Butler, R. J. (2018). The rise of the ruling reptiles and
ecosystem recovery from the Permo-Triassic mass extinction.
Proceedings of the Royal Society, Series B, 285(1880), 20180361.
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