Journal of Vertebrate Paleontology

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A new typothoracine aetosaur (Archosauria,

Pseudosuchia) from the Upper Triassic of India
with insights on biostratigraphy, diversification,
and paleobiogeography

Atrayee Haldar, Sanghamitra Ray & Saswati Bandyopadhyay

To cite this article: Atrayee Haldar, Sanghamitra Ray & Saswati Bandyopadhyay (06 Oct 2023):
A new typothoracine aetosaur (Archosauria, Pseudosuchia) from the Upper Triassic of India
with insights on biostratigraphy, diversification, and paleobiogeography, Journal of Vertebrate
Paleontology, DOI: 10.1080/02724634.2023.2253292

To link to this article: https://doi.org/10.1080/02724634.2023.2253292

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Published online: 06 Oct 2023.

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Journal of Vertebrate Paleontology e2253292 (22 pages)
© by the Society of Vertebrate Paleontology
DOI: 10.1080/02724634.2023.2253292

ARTICLE

A NEW TYPOTHORACINE AETOSAUR (ARCHOSAURIA, PSEUDOSUCHIA) FROM THE

UPPER TRIASSIC OF INDIA WITH INSIGHTS ON BIOSTRATIGRAPHY, DIVERSIFICATION,
AND PALEOBIOGEOGRAPHY

ATRAYEE HALDAR, 1* SANGHAMITRA RAY, 1 and SASWATI BANDYOPADHYAY 2

1
Department of Geology and Geophysics, Indian Institute of Technology, Kharagpur 721302, India; atrayeehalda@iitkgp.ac.in;
sray@gg.iitkgp.ernet.in;
2
Geological Studies Unit, Indian Statistical Institute, 203 B. T. Road, Kolkata 700108, India; saswati2602@gmail.com

ABSTRACT—A new typothoracine aetosaur is described based on multiple isolated and articulated left paramedian and
lateral osteoderms recovered from the Upper Triassic lower Dharmaram Formation of India. The partial carapace of the
new taxon is reconstructed as strongly discoidal based on the curvature of the paramedian osteoderms with the widest one
positioned dorsal to the mid-dorsal trunk vertebra. Asymmetric lateral osteoderms with acute flexion are considered as
precaudals with the angle of flexion decreasing posteriorly. Phylogenetic analysis recovered the new taxon as deeply
nested within the clades Typothoracinae and Paratypothoracini, and a sister taxon to Kocurypelta silvestris. The
autapomorphic characters involving paramedian osteoderms dorsal to the trunk vertebrae include dorsal surface
ornamented by large, irregular pits surrounding the dorsal eminence and radiating ridges in other areas, straight anterior
margin of the anteromedial corner of the anterior bar in dorsal view, and raised or ridged and ornamented posteromedial
corner. The current study highlights the significance of the new aetosaur, and the age of the lower Dharamaram
Formation is modified here as mid-Norian to Rhaetian based on global correlation with other coeval horizons. The
recovery of this taxon marks the first record of Paratypothoracini from high paleolatitudes of the Gondwanan region. The
study corroborates the earlier findings of a strong Laurasian faunal influx in India during the Late Triassic suggesting
possible land bridges and/or conducive environmental conditions for faunal dispersal.

http://zoobank.org/urn:lsid:zoobank.org:pub:FCC37C0E-107F-4B15-9460-4E802B465865

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

Citation for this article: Haldar, A., Ray, S., & Bandyopadhyay, S. (2023) A new typothoracine aetosaur (Archosauria,
Pseudosuchia) from the Upper Triassic of India with insights on biostratigraphy, diversification, and paleobiogeography.
Journal of Vertebrate Paleontology. https://doi.org/10.1080/02724634.2023.2253292

Submitted: February 7, 2023

Revisions Received: August 18, 2023
Accepted: August 24, 2023

INTRODUCTION Hunt & Lucas, 1991; Long & Murry, 1995; Lucas et al., 2007;
Marsh et al., 2020; Sawin, 1947; Small, 1998, 2002; Small &
The Aetosauria is an extinct clade of extensively armored
Martz, 2013), Greenland (Jenkins et al., 1994), Brazil, Argen­
pseudosuchian archosaurs that reached a high species diversity
tina (Bonaparte, 1969; Casamiquela, 1960, 1980; Desojo
during the Late Triassic (Heckert & Lucas, 2000, 2002; Parker,
et al., 2012), U.K., Germany, Italy, Poland, Scotland (Agassiz,
2016a, b). The aetosaurs are characterized by a small skull with
1844; Czepiński et al., 2021; Dróżdż, 2018; Dzik et al., 2000;
strongly tapering laterally expanded snout (e.g., Stagonolepis
Fraas, 1877; Hunt & Lucas, 1992; Long & Ballew, 1985;
robertsoni), upturned, and predominantly edentulous premax­
Parker, 2018a; Sulej et al., 2012; Teschner et al., 2022; von
illa, long tail, comparatively small forelimbs and an ornamen­
Huene, 1920; Walker, 1961), Morocco (Jalil et al., 1995), and
ted dermal armor or carapace covering the postcranial region
India (Bandyopadhyay & Ray, 2020; Kutty & Sengupta,
(Antczak, 2016; Desojo et al., 2013). The discovery of an
1989). Because of their wide global distribution and restricted
array of dentigerous material suggests that aetosaurs were
occurrences, aetosaurs are considered biostratigraphically
large omnivorous to insectivorous quadrupeds (Desojo &
important (Desojo & Ezcurra 2011; Desojo et al. 2013;
Ezcurra, 2011; Desojo et al., 2013; Paes-Neto et al., 2021a;
Heckert & Lucas, 1999, 2000; Lucas, 2018; Martz & Parker,
Parker, 2016a; Reyes et al., 2021; von Baczko et al., 2018).
2017; Parker, 2016a, b; Parker & Martz, 2011; Parker et al.,
Aetosaurs are documented within the Upper Triassic strata
2008).
of Texas, Arizona, New Jersey, Colorado, New Mexico (Case,
Gondwanan deposits of peninsular India (Fig. 1A) are
1932; Cope, 1875; Gregory, 1953; Heckert & Lucas, 1999;
known for rich vertebrate fossil assemblages (Bandyopadhyay,
2011; Bandyopadhyay & Ray, 2020) that include the three
Upper Triassic horizons, namely the Maleri and lower Dhar­
*Corresponding author.
Color versions of one or more of the figures in the article can be found maram formations of the Pranhita-Godavari Basin, and the
online at www.tandfonline.com/ujvp. Tiki Formation of the Rewa Gondwana Basin (Fig. 1B).

Published online 06 Oct 2023

 Haldar et al. —Typothoracine from India (e2253292-2)

FIGURE 1. A, major Gondwana basins of peninsular India; B, Triassic stratigraphy of Pranhita-Godavari Basin; C, geological map of the northern
part of the Pranhita-Godavari Basin showing the Dharmaram Formation (after Kutty et al., 1987); D, geological map (after Kutty & Sengupta, 1989;
Dasgupta et al., 2017) of the boxed area in (C) showing the fossil locality.

Aetosaurian remains have been reported from the Maleri and
Dharmaram formations (Kutty & Sengupta, 1989; Kutty et al.,
1987, 2007; Novas et al., 2011; von Huene, 1940). The Dhar­
maram osteoderms were subsequently identified as belonging
to Paratypothorax-like and Desmatosuchus-like taxa (Bandyo­
padhyay, 2011; Bandyopadhyay & Ray, 2020; Desojo et al.,
2013; Kutty & Sengupta, 1989), though these lack proper docu­
mentation and detailed description. In the current study, we
focus on comparative description of the Paratypothorax-like
osteoderms and established a new taxon from the lower Dhar­
maram Formation of the Pranhita-Godavari Basin. We also
expand the previous phylogenetic analyses to assess the topo­
logical position of the new taxon within Aetosauria, and
discuss its significance in terms of biostratigraphy and
paleobiogeography.
GEOLOGICAL SETTING
The Pranhita-Godavari Basin is a narrow rectilinear basin
trending NNW–SSE (Fig. 1C) and runs parallel to the basement
fabric in close conformity with the underlying Proterozoic rocks
(Tewari & Maejima, 2010). The basin is slightly asymmetrical
where the thickest sedimentary pile (about 4 km) is found
closer to the western margin (Chakraborty et al., 2003). The
beds trend NW–SE and their overall attitude is 5–15° towards
NE. A nearly continuous and thick Lower (Induan) to Upper
 Haldar et al. —Typothoracine from India (e2253292-3)
(Rhaetian) Triassic succession that comprises at least five for­
mations, each characterized by a distinct vertebrate faunal
assemblage, are known from the Pranhita-Godavari Basin (Ban­
dyopadhyay and Ray, 2020; Fig. 1B). The Dharmaram Formation
(Fig. 1C, D), exposed on the eastern part of the basin, comprises
thick cross-bedded coarse sandstone units at the base followed
by sandstone-red mudstone alternations, conglomerates and
cross-stratified carbonate grainstones. The formation has
yielded two distinct vertebrate faunal assemblages and encom­
passes the Triassic–Jurassic boundary (Bandyopadhyay & Roy­
Chowdhury, 1996). Various vertebrate fossils have been
recovered from the lower Dharmaram Formation that included
a ptychoceratodontid (Bandyopadhyay & Ray, 2020; Kutty &
Sengupta, 1989), a xenacanthid (Nath & Yadagiri, 2007; Prasad
et al., 2008), a Nicrosaurus-like phytosaur, Paratypothorax-like
and Desmatosuchus-like aetosaurs, several sauropodomorphs,
and a neotheropod (Bandyopadhyay, 2011; Bandyopadhyay &
RoyChowdhury, 1996; Bandyopadhyay & Ray, 2020; Kutty
et al., 2007; Novas et al., 2011). Based on an essentially archo­
saur-dominated fauna, the age of the lower Dharmaram For­
mation is considered to be latest Norian to Rhaetian
(Bandyopadhyay & Sengupta, 2006; Bandyopadhyay & Ray,
2020).
MATERIAL AND METHODS
Material
Nine nearly complete paramedian osteoderms were examined,
of which ISIR267/1A–7A belonged to one individual, whereas
ISIR286 is isolated. In addition, seven complete lateral osteo­
derms (ISIR267/1B–7B) were also examined. All the specimens
are housed in the Geology Museum of the Indian Statistical Insti­
tute, Kolkata.
Methods
Nature of Preservation and Fossil Preparation—The speci­
mens were heavily encrusted with hard and thick calcareous
matrices, often masking the dorsal ornamentation. The speci­
mens were washed to remove the sediments, and dried. The
hard, calcareous encrustation was removed manually with the
help of electric engravers/vibrotools/small diamond-tipped drill
machines, and brushes of various sizes following the methods
outlined by Rixon (1976) and Leiggi and May (1994). After
removal of the matrices, lacquer (10% solution of polybuty­
methyl lacrylate in ethyl-methyl ketone) was applied over the
fossil material as a protective layer.
Taxonomic and Positional Nomenclature—As the paramedian
and lateral osteoderms of aetosaurs show considerable antero­
posterior variation within a single carapace (Parker & Martz,
2010), a thorough comparison was carried out for taxonomic
and positional identification of the Dharmaram osteoderms
with emphasis on typothoracine aetosaurs such as Typothorax
coccinarum (Heckert et al., 2010), Paratypothorax andressorum
(Lucas et al., 2006), Tecovasuchus chatterjeei (Martz & Small,
2006), and Kocurypelta silvestris (Czepiński et al., 2021). The
current work follows the positional nomenclature for aetosaur­
ian osteoderms as postulated by Long and Ballew (1985), Long
and Murry (1995), Heckert and Lucas (1999), Parker (2007,
2008), Parker and Martz (2010), Desojo et al. (2013), Parker
(2016b), and Marsh et al. (2020). The dorsal armor or carapace
(Parker 2016b) may be subdivided into four anteroposterior
columns of osteoderms, where those on either side of the
midline are the paramedians, and flanking these are the lateral
osteoderms (Desojo et al., 2013). Moreover, each column is
divided into rows, and given the names of the vertebral series
they cover (Parker, 2016b:8/58, fig. 3). Based on the position of
the paramedian osteoderms on the carapace, those placed ante­
riorly near the back of the skull are termed nuchal osteoderms
(Heckert et al., 2017), followed posteriorly by the cervical osteo­
derms that cover the cervical vertebrae. The set of irregular
osteoderms covering the throat area beneath the cervical ver­
tebrae are known as gular osteoderms (Long & Murry, 1995).
Following Parker (2016b), the paramedian osteoderms dorsal
to the trunk vertebrae are dorsal trunk paramedians, whereas
those on the ventral side and beneath the trunk vertebrae are
termed as ventral trunk osteoderms (Heckert et al., 2010). Fol­
lowed by those on the dorsal sacral region, osteoderms covering
the caudal vertebrae constitute dorsal caudal paramedians (Long
& Ballew, 1985; Parker, 2016b) and those placed ventrally with
respect to the caudal vertebrae are ventral caudal paramedians
(Heckert et al., 2010). Appendicular and palpebral osteoderms
are present in some taxa but not all (Desojo et al., 2013;
Heckert & Lucas, 1999; Schoch, 2007).
Measured Parameters—Several parameters (Fig. 2) of the
lower Dharmaram aetosaurs were measured by digital calipers
with a precision of 0.01 mm. Width (W) and length (L) of the
osteoderms (Fig. 2A, C) are measured as the mediolateral (=
transverse) and anteroposterior (= longitudinal) dimensions,
respectively (Heckert and Lucas, 1999, 2000). Angle of flexion
of the paramedian and lateral osteoderms were measured in
the posterior views (Fig. 2B, D).
Phylogenetic Analysis—The data matrix used in the current
study is predominantly based on those of Parker (2016a),
Reyes et al. (2021), and Paes-Neto et al. (2021a), and is heavily
biased towards characters pertaining to the cranial elements
and osteoderms (Supplementary File 1), as postcranial skeletal
elements of many of these taxa are not known. Our phylogenetic
analysis includes 28 taxa and 98 characters (Supplementary File
2.NEXUS) in TNT version 1.5 (Goloboff et al., 2008) using
New Technology Search with default settings for sectorial
search, ratchet, tree drift, and tree fusing, where Postosuchus
kirkpatricki (Chatterjee, 1985) has been used as the operational
outgroup. The constraint for the number of times the minimum
tree length was set as 100. Systematic resampling has been
carried out with 1000 replicates and a removal probability of 5
to assess the support for each clade. Two separate analyses
were performed, where (i) all the characters were treated as
unordered and (ii) 13 characters (3, 4, 14, 20, 22, 23, 28, 64, 70,
73, 76, 79, 83) were treated as ordered (= additive, Goloboff
et al., 2008) as given by Reyes et al. (2021). Sources for infor­
mation for the taxa are as follows: Adamanasuchus eisenhardtae
(Lucas et al., 2007), Aetosauroides scagliai (Casamiquela, 1960;
Paes-Neto et al. 2021b), Aetosaurus ferratus (Fraas, 1877;
Marsh, 1896), Apachesuchus heckerti (Spielmann & Lucas,
2012), Calyptosuchus wellesi (Long & Ballew, 1985), Coahoma­
suchus kahleorum (Heckert & Lucas, 1999), Coahomasuchus
chathamensis (Heckert et al., 2017; Hoffman et al., 2018), Desma­
tosuchus smalli (Parker, 2005), Desmatosuchus spurensis (Case,
1920), Gorgetosuchus pekinensis (Heckert et al., 2015), Kocury­
pelta silvestris (Czepiński et al., 2021), Longosuchus meadei
(Sawin, 1947), Lucasuchus hunti (Long & Murry, 1995), Neoaeto­
sauroides engaeus (Bonaparte, 1969), Paratypothorax sp. (Long
& Murry, 1995), Paratypothorax andressorum (Long & Ballew,
1985), Redondasuchus rineharti (Spielmann et al., 2006), Rioarri­
basuchus chamaensis (Zeigler et al., 2002), Tecovasuchus chatter­
jeei (Martz & Small, 2006), Typothorax coccinarum (Hunt &
Lucas, 1990), Scutarx deltatylus (Parker, 2016b), Sierritasuchus
macalpini (Parker et al., 2008), Stenomyti huangae (Small &
Martz, 2013), Stagonolepis robertsoni (Agassiz, 1844), Stagonole­
pis olenkae (Dróżdż, 2018; Sulej, 2010), and new Indian taxon
(ISIR 267, ISIR286; this study). In the current analysis, Aetobar­
bakinoides brasiliensis (Desojo et al., 2012) is omitted as there is
little information of the osteoderms because of poor preservation
and lack of lateral osteoderms, though the taxon is well
 Haldar et al. —Typothoracine from India (e2253292-4)
represented by endoskeletal elements (Parker, 2016a); previous
authors considered Aetobarbakinoides as a wild card taxon
(Heckert et al., 2015; Parker, 2016a). Polesinesuchus aurelioi is
omitted based on a recent study by Paes-Neto et al. (2021a)
that considered it as a junior synonym of Aetosauroides scagliai.
Additionally, Kocurypelta silvestris known from Poland (Cze­
piński et al., 2021) is scored for the first time. The following
three characters have been modified (Supplementary File 1)—
character 53: pattern on dorsal surface of the paramedian osteo­
derms, character 65: ornamentation on posteromedial surface of
the paramedian osteoderms dorsal to the trunk vertebrae, and
character 70: posteromedial corner of paramedian osteoderms
dorsal to the trunk vertebrae. We added two new characters
into the analysis (Supplementary File 1: Fig. S1): character 97:
ventral flexion at the center of ossification of the paramedian
osteoderms positioned anterior dorsal to the trunk vertebrae;
character 98: ventral flexion at the center of ossification of the
paramedian osteoderms positioned mid-posterior dorsal to the
trunk vertebrae.
Institutional Abbreviations—ISI, Indian Statistical Institute,
Kolkata, India; PEFO, Petrified Forest National Park, Arizona,
U.S.A.; SMNS, Staatliches Museum für Naturkunde, Stuttgart,
Germany; UMMP, University of Michigan, Museum of Paleon­
tology, Ann Arbor, MI, U.S.A.
SYSTEMATIC PALEONTOLOGY
DIAPSIDA Osborn, 1903
ARCHOSAURIA Cope 1869, sensu Gauthier & Padian
1985
PSEUDOSUCHIA Zittel 1887–1890, sensu Gauthier &
Padian 1985
AETOSAURIA Marsh 1884, sensu Parker, 2007
STAGONOLEPIDIDAE Lydekker 1887, sensu Heckert
& Lucas 2000
TYPOTHORACINAE von Huene 1915, sensu Parker
2016a
VENKATASUCHUS, gen. nov.
VENKATASUCHUS ARMATUM, gen. et sp. nov.
(Figs. 3–9)
Holotype—ISIR267/1–7, eight nearly complete paramedian
osteoderms, with associated lateral osteoderms.
Etymology—Generic name is after N. Venkata Raja Reddy, an
enthusiast who was helpful in finding vertebrate fossils from the
Pranhita-Godavari Basin, and from ‘suchus,’ the Greek word for
FIGURE 2. Schematic line drawings showing
measured parameters of a left, A–B, parame­
dian osteoderm in A, dorsal; B, posterior
views; and C–D, lateral osteoderm in C,
dorsal; and D, posterior views. Abbreviations:
ab, anterior bar; abL, length of the anterior
bar; aW, anterior width; de, dorsal eminence;
dfL, length of dorsal flange; dfW, width of the
dorsal flange; L, length measured along the
dorsal eminence; LL, length of the lateral
margin; MW, medial width; pW, posterior
width; vfL, length of the ventral flange; vfW,
width of the ventral flange; W, width measured
along dorsal eminence.
crocodile. The specific name is in Latin, where ‘armatum’ means
‘armored.’
Referred Specimen— ISIR286, a complete paramedian
osteoderm.
Differential Diagnosis—The taxon differs from all other aeto­
saurs by a unique combination of the following characters: extre­
mely wide dorsal trunk paramedian osteoderms (shared with all
typothoracines), raised anterior bar (shared with most aetosaur­
ians except Desmatosuchus), dorsal surface ornamentation con­
sisting of irregular large- to medium-sized pits and radial ridges
surrounding the dorsal eminence (or center of ossification) that
does not contact the posterior margin of osteoderm in the
trunk region (shared with Desmatosuchus spurensis, Lucasuchus,
Rioarribasuchus, Tecovasuchus, and dorsal trunk region of Para­
typothorax andressorum), posterior margin bears transverse, pos­
teroventrally sloping flange (bevel) (shared with Tecovasuchus,
Kocurypelta, and a few specimens of Paratypothorax andres­
sorum), presence of ventral strut (shared with Redondasuchus
rineharti, Typothorax coccinarum, Tecovasuchus, a few speci­
mens of Paratypothorax andressorum, Paratypothorax sp., Calyp­
tosuchus, and Adamanasuchus), straight lateral edge of the
dorsal trunk paramedian osteoderms in dorsal view (shared
with Desmatosuchus spurensis, Stagonolepis robertsoni, Aeto­
sarus ferratus, and Apachesuchus), dorsal eminence strongly
offset medially (shared with Rioarribasuchus, Tecovasuchus,
and Paratypothorax andressorum), highly reduced and narrow
triangular dorsal flange of lateral osteoderms (shared with Rioar­
ribasuchus, Tecovasuchus, and Paratypothorax sp.), ventrally
concave embayment of the posterior face of dorsal spine of
lateral osteoderm (shared with Typothorax coccinarum and
Longosuchus), elongated flattened horn-like dorsal eminence
of lateral osteoderm (shared with Rioarribasuchus, Tecovasu­
chus, Kocurypelta, Paratypothorax sp., and Paratypothorax
andressorum), strongly acute angle of flexion between dorsal
and lateral flanges of lateral osteoderm (shared with Redondasu­
chus rineharti, Rioarribasuchus, Tecovasuchus, and Paraty­
pothorax andressorum), distinct flexion in posterior view at the
center of ossification of paramedian osteoderm (shared with
Calyptosuchus, Stagonolepis robertsoni, Stagonolepis olenkae,
Scutarx deltatylus, Redondasuchus rineharti, Typothorax cocci­
narum, and Rioarribasuchus).
The taxon differs from other members of Typothoracinae and
Paratypothoracini by a unique combination of the following
characters: straight or sigmoidal lateral edge of the paramedian
osteoderms (shared with Tecovasuchus and Kocurypelta), a pro­
minent and rounded ridge extending along the entire
 Haldar et al. —Typothoracine from India (e2253292-5)

mediolateral width of the paramedian osteoderm in continuation Martz & Small, 2006:figs 1, 2). Other similar features included
with the dorsal eminence (shared with Kocurypelta), straight a narrow, weakly raised anterior bar with a distinct anterolateral
anterior margin of anterior bar on the medial side of the parame­ projection, and medial offset of the dorsal eminence resulting in
dian osteoderm (shared with Paratypothorax). a relatively shorter medial width (about 40% of the total width)
Autapomorphic characters involving paramedian osteoderms in comparison to the longer lateral portion. The lateral osteo­
dorsal to the trunk vertebrae: dorsal surface ornamented by derms of Venkatasuchus are asymmetric with a longer ventral
large, irregular pits surrounding the dorsal eminence and radiat­ flange with respect to the dorsal flange and an acute angle of
ing ridges in other areas, straight anterior margin of the antero­ flexion showing similarity with those of typothoracines
medial corner of the anterior bar in dorsal view, and raised or (Heckert et al., 2010:fig. 4D; Lucas et al., 2006: fig. 6; Martz &
ridged and ornamented posteromedial corner. Small, 2006:fig. 3; Parker, 2016a:supplemental information, fig.
Locality, Horizon, and Age—Near the village of Rampur (19° 13I–J). Such similarities suggest that Venkatasuchus is positioned
9′54″N, 79°35′6″E; Fig. 1D), Adilabad district, Telangana, India; within the Typothoracinae (sensu Parker, 2016a), a well-sup­
lower Dharmaram Formation of the Pranhita-Godavari Basin; ported clade within Aetosaurinae of the family Stagonolepididae
Upper Triassic: middle Norian to Rhaetian (Bandyopadhyay & (sensu Heckert & Lucas, 2000).
Sengupta, 2006; Bandyopadhyay & Ray, 2020; Kutty et al.,
2007; Novas et al., 2011; current study).
Comments—The specimens (ISIR267 and ISIR286) recovered DESCRIPTION
from the lower Dharmaram Formation of the Pranhita-Godavari
Paramedian Osteoderms
Basin, India, are placed within the monophyletic clade Aeto­
sauria (Heckert et al., 1996; Parrish, 1994) based on greater General Features—The left paramedian osteoderms (ISIR267/
width with respect to the length, sculptured ornamentation, 1A–7A, Figs. 3–6) recovered from the lower Dharmaram For­
and more specifically presence of dorsal eminence, and trans­ mation are moderately well preserved, though often with incom­
verse flexion close to the medial margin of the osteoderm plete/broken anterior and posterior margins. These are
(Parker et al., 2022). Although isolated dermal plates are difficult quadrangular, much wider than long and have a high width-to-
to assign correctly at the generic level using autapomorphies length ratio (W:L) of ≥ 4:1, except for ISIR267/1A (W:L = 3.7:1,
(Martz & Small, 2006), these may be taxonomically identified Table 1). Two of these osteoderms were found as articulated or
based on a unique combination of charaters (Long & Ballew, overlapping (ISIR267/7A), where remnants of the preceding
1985) such as osteoderm proportions, pattern of ornamentation osteoderm were found attached to ISIR267/1A suggesting that
and presence or absence of several morphological traits. As a these belonged to the same individual of an aetosaur.
result, type specimens of various valid aetosaur taxa consist pre­ Dorsal View—The osteoderms have a predominantly straight
dominantly of osteoderms (Parker et al., 2008). A few such lateral and medial margin (Fig. 3). In most of the specimens,
genera include Stagonolepis robertsoni (Agassiz, 1844), the anterior margin is partially preserved (Fig. 3A–J).
Typothorax coccinarum (Cope, 1875), and Paratypothorax However, ISIR267/6A shows a thick, weakly raised and distinct
andressorum (Long & Ballew, 1985). anterior bar (Fig. 3G, H) constituting 36% of the osteodermal
Aetosauria consists of two main stem-based clades, Stagonole­ length measured at the center of ossification. The posterior
pidoidea and Aetosaurinae (Heckert & Lucas, 2000; Parker, margin of the anterior bar is often incised by ornamentation.
2007, 2016a; Reyes et al., 2021), which are supported by several The anterior bar preserved in ISIR267/5A (Fig. 3E, F) and
synapomorphies. In most of the taxa belonging to Stagonolepi­ ISIR267/6A (Fig. 3G, H) broadens out laterally and projects
doidea (sensu Reyes et al., 2021), the paramedian osteoderms into a short triangular anterolateral process which is gently
have width/length proportion <3, well-developed spikes or pro­ rounded instead of being a sharp point similar to that of Paraty­
minent bosses as dorsal eminences. In contrast, the dorsal para­ pothorax andressorum (Lucas et al., 2006:fig. 3A) and
median osteoderms of Venkatasuchus exhibit an extremely high Typothorax (Martz, 2002: fig. 4.28c, d), though in other speci­
width:length ratio (W:L ≥ 4:1, Table 1), which is similar to that mens, the anterolateral and anteromedial corners are missing.
of Typothorax coccinarum (W:L = 4:1, Desojo et al., 2013:fig. In all the specimens, there is a low mediolaterally extended
7G; Long & Murry, 1995:fig. 100A–D), Paratypothorax sp. (W: dorsal eminence, which is offset medially resulting in a short
L = 4.6:1, Long & Murry, 1995:fig. 114A–E), Paratypothorax mediodorsal surface constituting only 41–45% of the width of
andressorum (W:L = 4.8:1, Desojo et al., 2013:fig. 7J; Lucas the paramedian osteoderms. The dorsal eminence is situated pos­
et al., 2006:fig. 3) and Tecovasuchus chatterjeei (W:L = 4.4:1, teriorly, but does not contact the posterior margin (Fig. 3) similar
to that seen in Desmatosuchus spurensis (Parker, 2008:fig. 24A–
K), Lucasuchus hunti (Parker & Martz, 2010:figs. 5A, 6B), Rioar­
TABLE 1. Measured parameters of the available paramedian ribasuchus (=Heliocanthus) chamaensis (Parker, 2007:fig. 3A;
osteoderms recovered from the lower Dharmaram Formation. All Zeigler et al. 2002:fig. 2D, E), Tecovasuchus chatterjeei (Martz
measurements are in mm, if not mentioned otherwise. Abbreviations: & Small, 2006:fig. 2A, E), and Paratypothorax andressorum
ABL, length of anterior bar along the dorsal eminence; L, length (Lucas et al., 2006:fig. A–D). The dorsal eminence is placed on
measured along dorsal eminence; MW, medial width; W, width a prominent and raised transverse ridge extending parallel
measured along dorsal eminence.
along, and 15–20 mm anterior to the posterior margin.
Ornamentation on Dorsal Surface—The dorsal ornamentation
Registration is more prominent in the specimens ISIR267/1A–4A (Fig. 3A–F)
Angle of
number W L W/L ABL/L MW/W flexure (°) in comparison to the faint and fine ornamentation of ISIR267/
6A–7A (Fig. 3G–J). It consists of a combination of fine ridges,
1 ISI/R267/1A 302.3 82.3 3.7 0.14 0.32 155 elongated grooves, and circular to elliptical pits (Figs. 3A–J,
2 ISI/R267/2A 328.6 - - 0.21 0.42 155
3 ISI/R267/3A 338 76.5 4.42 0.2 0.41 150 4A, C), especially surrounding the dorsal eminence. In general,
4 ISI/R267/4A 310 69 4.5 0.22 0.44 155 the ornamentation is more incised on the part medial to the
5 ISI/R267/5A 310 72 4.31 0.23 0.45 160 dorsal eminence in contrast to that on the lateral side. The
6 ISIR267/6A 296.2 67.3 4.4 0.36 0.44 155 dorsal eminence is surrounded by randomly arranged, dense
7 ISI/R267/7A/a 274.2 61.2 4.5 0.16 0.4 155 pits of varying sizes (Fig. 4A, B). The alternating ridges and
8 ISIR267/7A/b 251.8 61 4.13 - 0.34 155
9 ISI/R286 310 67.6 4.6 0.16 0.36 - grooves radiate from the center of ossification or dorsal emi­
nence, and within the grooves multiple deep pits of varying
 Haldar et al. —Typothoracine from India (e2253292-6)

FIGURE 3. Venkatasuchus armatum. Partial and nearly complete left paramedian osteoderms and their interpretative line drawings in dorsal views,
A–B, ISIR267/3A; C–D, ISIR267/4A; E–F, ISIR267/5A; G–H, ISIR267/6A; I–J, ISIR267/7A, two articulated osteoderms. Arrow indicates lateral side.
The boxed areas I, II, and III are magnified in Figure 4. Abbreviations: ab, anterior bar; alp, anterolateral projection bpm, beveled posterior margin;
de, dorsal eminence; pdr, posterodorsal ridge. Scale bar equals 50 mm.

sizes are present (Fig. 4A). These pits may be either isolated, con­
densed and surrounding the dorsal eminence or present inside
the grooves in between the ridges (Taborda et al., 2015). The iso­
lated pits are generally large, circular to sub-circular and bor­
dered by a thick rounded or oval margin whereas the pits
present inside the grooves are usually small and elongated or
elliptical (Fig. 4C). In the specimens, ISIR 267/2A–3A, the pits
become larger and deeply incised near the dorsal eminence
and the ridges between the large pits show an anastomosing
pattern (Fig. 4A, B). On the medial side, the ridges radiate at a
high acute angle (nearly 40°) with an anteromedial inclination
with respect to the dorsal eminence (Fig. 4A, B), whereas later­
ally the number of ridges are less, and either at low acute angle
(less than 10°) with an anterolateral inclination or sub-parallel
to the posterior margin (Fig. 4B). The ridges often bifurcate
near the medial edge (Fig. 4C). As mentioned earlier, a mediolat­
erally extended transverse ridge is present posteriorly on the
dorsal surface of the osteoderm. Beyond this posterodorsal
ridge is a thick, sharp edged posteroventrally sloping flange
that gives the posterior margin a beveled appearance. This
beveled area is ornamented by sub-circular pits, grooves, and
ridges (Figs. 3, 4B, C). Such ornamentation of the beveled area
is not seen in other aetosaurs, except for the presence of
shallow grooves in the beveled area of Tecovasuchus (Martz &
Small, 2006:figs. 1B–F, 2B–F).
Ventral View—The ventral surface of all the paramedian
osteoderms is smooth and devoid of any ornamentation. Five
of the examined paramedian osteoderms (ISIR267/1A–5A)
possess a thick, wide, anteroposteriorly convex ventral keel/
strut extending mediolaterally along the osteodermal width
(Fig. 5A–J) and situated equidistant from the anterior and pos­
terior margins. The strut is thickest along the line of arching, flat­
tens out gradually on the lateral side, and terminates abruptly
medially (Fig. 5A–J). In ISIR267/6 (Fig. 5G, H) and the two
articulated osteoderms (ISIR267/7A, Fig. 5I–J), the ventral
strut is flattened and/or nearly absent especially in ISIR267/7A
indicating a degree of variation in the prominence of the
ventral strut within the trunk region.
 Haldar et al. —Typothoracine from India (e2253292-7)

FIGURE 4. Venkatasuchus armatum. Boxed areas (I–III) of Figure 3 at high magnification showing distinct patterns of ornamentation composed of
circular pits and radiating ridges on selected paramedian osteoderms in dorsal views. A, ISIR267/3A, medial side; B, ISIR267/4A, lateral side; C,
ISIR267/6A, medial side. Abbreviations: ab, anterior bar; bpm, beveled posterior margin; de, dorsal eminence; pdr, posterodorsal ridge. Pits and
ridges are indicated by arrows and arrowheads, respectively. Scale bars equal 50 mm.

Posterior View—All the osteoderms, except for ISIR267/1A
(Fig. 6A, B), are widely arched (Fig. 6C–J) at the center of ossi­
fication or dorsal eminence with the angle of flexure ranging
between 155–160° (Table 1). The arching is asymmetric and
offset medially keeping parity with that of the dorsal eminence.
On the other hand, ISIR267/1A (Fig. 6A, B) exhibits a ventral
arching where the center of arching is not at the center of ossifi­
cation but shifted laterally. This possibly is a taphonomic artifact,
caused by overburden sediment pressure.
Lateral Osteoderms
There are seven well-preserved, isolated asymmetric left
lateral osteoderms (Fig. 7A–X) in the collection that were recov­
ered in association with the paramedian osteoderms (ISIR267/
1A–7A). These lateral osteoderms possess a reduced dorsal
flange, long ventral flange and curved horn-like structure,
where the two flanges meet at an acute angle. Although respect­
ive positions were not known, measured parameters (Fig. 2)
reveal that the lateral osteoderms may be subdivided into two
groups based on the ratio between the dorsal flange width and
length (dfW:dfL), and angle of flexure between the two flanges
(Table 2). These included group I, comprising four lateral osteo­
derms (ISIR267/1B–2B), where dfW:dfL < 1:1 and angle of
flexure ranges from 50–40° (Fig. 7A–P). Group II, on the other
hand comprises three lateral osteoderms (ISIR267/4B) having
dfW:dfL > 1:1, and relatively low angle of flexure (30–25°; Fig.
7Q–X). In both the groups, the two flanges meet laterally to
form a short, dorsoventrally compressed, anteriorly convex or
curved and posteriorly embayed horn-like structure as seen in
the typothoracines (Heckert et al., 2010; Lucas et al., 2006;
Martz & Small, 2006).
The dorsal flange is triangular, and smaller than the ventral
flange. In many of the lateral osteoderms, the dorsal surface is
ornamented with fine ridges alternating with grooves radiating
from the lateral boss or eminence (Fig. 7A, B). The medial
edge of the dorsal flange is keeled in all the specimens and
bears a slightly developed cut-off at the anteriormost corner of
the medial edge, except for ISIR267/1B–2B (Fig. 7A–P). The
ventral flange is plate-like, broad (vfW/vfL ≥ 2, Table 2) and
much wider than the dorsal flange (vlfW/dfW ≥ 1.4). The exter­
nal surface of the ventral flange is rough with faint radial
ridges/corrugations. In a few of the lateral osteoderms examined,
the external surface of the flanges shows profusion of minute pits/
foramina, as exemplified by ISIR267/4B (Fig. 8A).
ISIR286
An isolated quadrangular, left paramedian osteoderm,
ISIR286 (Fig. 8B–H), was recovered from the same locality as
the paramedian osteoderms of ISIR267, and is comparable to
the latter based on high width (W:L = 4.6:1; Table 1), medially
offset center of ossification (MW/W = 0.36; Table 1), ornamenta­
tion pattern on the dorsal surface, and beveled posterior edge.
ISIR286 has an overall sinuous or wavy structure, and straight
medial and sigmoidal lateral margins, respectively. The specimen
exhibits a narrow and strongly developed anterior bar (ABL/L =
0.16) though it is incompletely preserved, especially in the lateral
direction. The anterior margin of the anterior bar is straight (Fig.
8B, C) and not scalloped as seen in Desmatosuchus spurensis
(Parker, 2008:figs. 24D–G) and Longosuchus meadei (Parker &
Martz, 2010:figs. A, C). Other features included a small anterolat­
eral projection, reduced dorsal eminence, narrow posterodorsal
ridge, and broadly convex ventral strut. As in ISIR267, the
 Haldar et al. —Typothoracine from India (e2253292-8)

FIGURE 5. Venkatasuchus armatum. Partial and nearly complete left paramedian osteoderms and their interpretative line drawings in ventral views,
A–B, ISIR267/3A; C–D, ISIR267/4A; E–F, ISIR267/5A; G–H, ISIR267/6A; I–J, ISIR267/7A, two articulated osteoderms. Arrow indicates lateral side.
Abbreviation: vs, ventral strut. Scale bars equal 50 mm.

center of ossification rests on a narrow posterodorsal ridge
extending parallel to the posterior margin of the osteoderm.
However, the dorsal eminence is low and contrasts with that of
ISIR267. The dorsal ornamentation, on the other hand, is
similar to that of ISIR267 and consists of fine ridges alternating
with moderately incised grooves that radiate from the center of
ossification. These radial grooves are more prominent laterally,
and contain circular and elliptical pits (Fig. 8D), and form an
incised posterior margin of the anterior bar. Anteriorly the
ridges are at high angle to the anterior margin. Posterior to the
posterodorsal ridge, the osteoderm forms a thin, posteroventrally
sloping narrow flange (about 20 mm in length), which gives the
posterior margin a beveled appearance. Small pits and ridges
and grooves radiating posteriorly from the center of ossification
characterize the posterodorsal ridge and the beveled posterior
margin. The ventral surface of ISIR286 (Fig. 8E, F) is smooth
and devoid of any ornamentation. The ventral strut is thick and
widely convex at the middle and covers almost the entire
length of the osteoderm. In posterior view (Fig. 8G, H), the
osteoderm shows a sinuous structure with a wide convexity at
the center of ossification, possibly because of postmortem distor­
tion (Fig. 8H).
PHYLOGENETIC ANALYSIS
The results of the analysis are similar to those of the earlier
studies (e.g., Heckert et al., 2015; Hoffman et al., 2018; Paes-
Neto et al., 2021a; Parker, 2007, 2016a; Reyes et al., 2021;
Schoch & Desojo, 2016) though there are a few significant differ­
ences. The resultant tree topology shows better resolution and
higher clade support for the phylogenetic analysis using unor­
dered characters (Fig. 9A) in comparison to that of the ordered
characters (Supplementary File 3: Fig. S2), and hence the latter
is not considered here. The analysis involving unordered charac­
ters has yielded 19 most parsimonious trees (Supplementary File
4) with a tree length of 261 steps, consistency index (CI) of 0.540
and retention index (RI) of 0.683. The strict consensus tree (Fig.
9A) is poorly resolved with polytomies present within the Aeto­
saurinae and Stagonolepidoidea although the major clades of
Typothoracinae and Desmatosuchini are distinct and resolved.
 Haldar et al. —Typothoracine from India (e2253292-9)

FIGURE 6. Venkatasuchus armatum. Partial and nearly complete left paramedian osteoderms and their interpretative line drawings in posterior views
(column wise), A–B, ISIR267/1A; C–D, ISIR267/4A; E–F, ISIR267/5A; G–H, ISIR267/6A; I–J, ISIR267/7A, two articulated osteoderms. Arrow indi­
cates lateral side. Arrowheads indicate dorsal eminence or center of ossification. Scale bars equal 50 mm.

The 50% majority rule tree (Fig. 9B) giving an unambiguous
aetosaur inter-relationship is better resolved and recovered
Aetosauroides scagliai as the earliest-diverging non-stagonolepi­
did aetosaurian, as seen in previous analyses (Desojo et al., 2012;
Heckert et al., 2015; Parker, 2016a; Reyes et al., 2021). The fully
resolved resampled tree (Fig. 9C) is conformable with that of
Heckert et al. (2015), Parker (2016a), Schoch and Desojo
(2016), Hoffman et al. (2018), Reyes et al. (2021), and Paes-
Neto et al. (2021a). The clades of Aetosauria (Node A) and Sta­
gonolepididae (Node B) are supported by 10 and three unam­
biguous synapomorphies, respectively (Supplementary File 5).
In contrast to previous literature (Heckert and Lucas, 1999;
Parker, 2007, 2016a) Stenomyti huangae is recovered as an
early-diverging taxon within Stagonolepidoidea (Node C), the
latter being defined by six unambiguous synapomorphies (Sup­
plementary File 5). The most well resolved clade is Desmatosu­
chini (Node E), which is defined by 11 synapomorphies. A
major clade, Aetosaurinae (Node F) is supported by two synapo­
morphic characters where Coahomasuchus chathamensis is
recovered as an early-diverging aetosaurine and the later-diver­
ging/derived clades of Typothoracinae (node G) and Paraty­
pothoracini (node H) are well resolved and defined by two and
seven unambiguous synapomorphies, respectively (Supplemen­
tary File 5). The clade Typothoracinae comprises Apachesuchus
heckerti as an early-diverging taxon whereas the new taxon, Ven­
katasuchus armatum is deeply nested within Paratypothoracini
and is sister taxon to Kocurypelta silvestris (Node K, Fig. 9C)
based on characters related to the paramedian osteoderms
dorsal to the trunk vertebrae such as a weakly raised anterior
bar (character 52) and a prominent and rounded posterodorsal
ridge extending along the entire mediolateral width in continu­
ation with the dorsal eminence (character 65). Venkatasuchus
armatum is characterized by several autapomorphic features
related with the paramedian osteoderms dorsal to the trunk
vertebrae, including ornamentation on the dorsal surface com­
prising large, irregular pits surrounding DE and radiating
ridges in other areas (character 53), straight anterior margin of
the anteromedial corner of the anterior bar in dorsal view (chara­
ter 67), posteromedial corner is raised/ridged and ornamented
(character 70).
DISCUSSION
Carapace Reconstruction
Reconstruction of the carapace of Venkatasuchus is based on
the associated osteoderms of ISIR267. The positional identifi­
cation of the available paramedian (ISIR267/1A–7A) and
lateral (ISIR267/1B–7B) osteoderms of Venkatasuchus follows
Martz (2002) and Martz and Small (2006). Partially complete
and articulated skeletons of various taxa (e.g., Stagonolepis, Des­
matosuchus, Typothorax, Calyptosuchus) indicate that each pair
of paramedian and lateral osteoderms correspond to a single
trunk vertebra (Heckert et al., 2010; Martz, 2002; Walker,
1961). However, this 1:1 correspondence may not be true for cer­
vical vertebrae, as is evident from Desmatosuchus and Longosu­
chus (Parker, 2008). In these taxa, five or six osteoderm plates are
seen over nine cervical vertebrae (Long & Ballew, 1985; Paes-
Neto et al., 2021a; Parker, 2008, 2018 a, b). Thus, reconstruction
of a carapace based on isolated and semi-articulated paramedian
and lateral osteoderms is limited.
As phylogenetic analysis (current study) revealed that
Venkatasuchus belongs to the clades Typothoracinae and
Paratypothoracini (Fig. 9), its partial carapace comprising wide
paramedian (W:L ≥ 4:1) and strongly asymmetric lateral
osteoderms is reconstructed based on the carapace of
Typothorax coccinarum (W:L ≥ 4:1; Heckert et al., 2010) along
with that of Rioarribasuchus (W:L ≥ 3.5:1; Parker, 2007, 2016a;
 Haldar et al. —Typothoracine from India (e2253292-10)

FIGURE 7. Venkatasuchus armatum. Nearly complete left lateral osteoderms and their interpretative drawings in dorsal, ventral, medial, and lateral
views, A–H, ISIR267/1B; I–P, ISIR267/2B; Q–X, ISIR267/4B. Abbreviations: df, dorsal flange; vf, ventral flange. Scale bars equal 50 mm.

Zeigler et al., 2002), Tecovasuchus (W:L ≥ 4:1; Martz & Small,
2006) and Paratypothorax sp. (PEFO 3004, W:L ≥ 4:1; Long &
Murry, 1995; Lucas et al., 2006:figs. 3, 4), as these taxa exhibit
similar body plan because of their proportionately wide parame­
dian osteoderms. All the available paramedian osteoderms of
Venkatasuchus are extremely wide (W ≥ 250 mm, Table 1) with
the widest being ISIR267/3A (W = 338 mm), suggesting that
these were positioned dorsal to the trunk vertebrae and ranged
in position from anterior through to the mid-trunk region with
respect to the carapace (Fig. 10A, B). This resulted in a strongly
 Haldar et al. —Typothoracine from India (e2253292-11)

TABLE 2. Measured parameters of the available lateral osteoderms recovered from the lower Dharmaram Formation. All measurements are in
mm, if not mentioned otherwise. Abbreviations: dfL, length of dorsal flange; dfW, width of dorsal flange; vfL, length of ventral flange; vfW, width
of ventral flange.

Registration number dfL dfW dfW/dfL vfL vfW vfW/vfL vfW/dfw Angle of flexure (°)
1 ISIR267/1B 58.6 54 0.92 36.5 - - - 50
2 ISIR267/2B 52 53.3 0.98 36 76.3 2.12 1.43 50
3 ISIR267/3B 35.3 - - 35.3 - - - 50
4 ISIR267/4B 53.7 52 0.97 38.4 95 2.48 1.83 40
5 ISIR267/5B 53.6 59.6 1.11 47.2 102 2.16 1.7 25
6 ISIR267/6B 56.6 60 1.06 54.7 92.6 1.69 1.54 30
7 ISIR267/7B 50.5 56 1.11 44.8 - - - 30

discoidal carapace as in Typothorax and Paratypothorax
(Heckert et al., 2010; Long & Ballew, 1985).
The two articulated and overlapping paramedian osteoderms
(ISIR267/7A) are considered as placed anteriorly with respect
to the other paramedians based on their minimum width, less
pronounced dorsal eminence, lightly incised ornamentation
pattern composed mainly of radiating ridges and weakly devel­
oped ventral strut as suggested by Martz (2002) and also evi­
denced from well preserved complete carapace of
Calyptosuchus (Case, 1932; Long & Ballew, 1985; Parker,
2018b), Aetosaurus (Schoch, 2007), Typothorax coccinarum
(Heckert et al., 2010), Coahomasuchus chathamensis (Heckert
et al., 2017), and Coahomasuchus kahleorum (Heckert &
Lucas, 1999). ISIR267/7A is successively followed by ISIR267/
6A, ISIR267/5A, ISIR267/1A, and ISIR267/4A in the posterior
direction as suggested by their increase in width, relatively
strong sculptured ornamentation on the dorsal surface, which is
composed of deeply incised large pits surrounding a low, pyrami­
dal dorsal eminence, and relatively well-developed ventral strut.
ISIR267/2A and ISIR267/3A are considered to have occupied
the middle portion of the dorsal carapace because these are the
widest available paramedians having strong ornamentation and
prominent ventral keel.
A tapering and short dorsal flange relative to a long ventral
flange is seen in the pre-caudal lateral osteoderms of Typothorax,
Tecovasuchus, and Paratypothorax (Heckert et al., 2010; Long &

FIGURE 8. Venkatasuchus armatum. A, ISIR267/4B, boxed area of Figure 7 at high magnification showing pits/foramina (arrows) and faint corruga­
tions (arrowheads) on the dorsal flange of a lateral osteoderm; B–H, ISIR286, a complete, isolated paramedian osteoderm and its interpretative line
drawings. B–C, dorsal views; D, boxed area of B at higher magnification showing radiating ridges (white arrowhead) on the dorsolateral surface; E–F,
ventral; and G–H, posterior views. Arrow and black arrowhead indicate lateral direction and postmortem distortion, respectively. Abbreviations: ab,
anterior bar; de, dorsal eminence; pdr, posterodorsal ridge; vs, ventral strut. Scale bars equal 50 mm.
 Haldar et al. —Typothoracine from India (e2253292-12)

FIGURE 9. Cladograms showing aetosaur inter-relationships. A, strict consensus tree of 19 MPTs; B, 50% majority-rule tree with values of Bremer
support given above the nodes; C, resampled tree with group present/contradicted (GC) values indicated by numbers above the nodes. The new Indian
taxon is given in bold.

Ballew, 1985; Long & Murry, 1995; Martz, 2002; Martz & Small, Taxonomic Identity
2006). This contrasts with the lateral osteoderms of the caudal
The paramedian osteoderms dorsal to the trunk vertebrae in
region, where the strong asymmetry between the dorsal and
Venkatasuchus differ from those of other typothoracine taxa
ventral flanges is lacking (Heckert et al., 2010). The lateral osteo­
including Apachesuchus (Spielmann & Lucas, 2012), Redondasu­
derms (ISIR267/1B–7B) examined in the current study have
chus (Spielmann et al., 2006), Typothorax (Long & Murry, 1995),
similar configuration as the pre-caudal region of the aforemen­
Rioarribasuchus (Zeigler et al., 2002), Paratypothorax (Lucas
tioned genera, where the flanges meet at an acute angle (angle
et al., 2006), Tecovasuchus (Martz & Small, 2006), and Kocury­
of flexure = 50–25°, Table 2) showing a tight flexion, unequal
pelta (Czepiński et al., 2021) based on a combination of several
flanges, and prominent dorsal eminence or horn-like structure.
diagnostic features such as the nature of the anterior bar,
This is also corroborated by a comparable lateral length (LL)
highly incised/sculptured ornamentation, beveled posterior
of the paramedian osteoderms with the length of the dorsal
margin, and presence of arching at the center of ossification. In
flange (dfL) of the lateral osteoderms (e.g., ISIR267/6A [LL =
contrast to Venkatasuchus, the dorsal paramedian osteoderms
58 mm] and ISIR267/1B [dfL = 58.6 mm]; and ISIR267/4A [LL
of Typothorax are characterized by a distinctly raised anterior
= 52 mm] and ISIR267/4B [dfL = 53.7 mm]), even though
bar, densely packed and deep interconnecting pits, absence of
lateral edges of most of the paramedians are not preserved or
distinct dorsal eminence except for a faint expression in the
partially broken. Three of the lateral osteoderms (ISIR267/1B–
anterior and mid-trunk paramedian osteoderms, faint expression
3B) have equal angle of flexure (50°), whereas ISIR267/4B and
in a few, absence of beveled posterior margin (Long & Murry,
ISIR267/6B–7B have low angles of flexure (= 40° and 30°,
1995:figs. 100A–D). Paratypothorax, on the other hand, is differ­
respectively). The minimum flexion is shown by ISIR267/5B (=
entiated from Venkatasuchus by long radiating depressions alter­
25°). Based on the angle of flexure, we hypothesized that
nating with distinct ridges from the center of ossification/a high
ISIR2671B–3B are anteriorly placed, ISIR267/4B and ISIR267/
dorsal eminence on the dorsal paramedian osteoderm, and
7B are intermediate in position, and followed posteriorly by
absence of arching (Long & Murry, 1995:fig. 113A; Lucas et al.,
ISIR267/5B–6B.
 Haldar et al. —Typothoracine from India (e2253292-13)

2006:fig. 3A, B). However, the absence or presence of ventral
strut represents intraspecific variation within the carapace of
Paratypothorax as is evident from its absence in Paratypothorax
sp. (DMNH 9942, Long & Murry, 1995) and presence in SMNS
syntypes and UMMP 8858 (Martz & Small, 2006:fig. 8B).
Although the dorsal paramedian osteoderms of Venkatasuchus
(Fig. 3) show similarity with those of Tecovasuchus chatterjeei
(Martz & Small, 2006:figs. 2A–F) in terms of ornamentation
and ornamented beveled posterior margin, there are distinct
differences. These include an indistinct dorsal eminence, a
weakly developed posterodorsal ridge, and absence of arching
in Tecovasuchus. Similarity is also observed between the lateral
osteoderms of Venkatasuchus (Fig. 7) and T. chatterjeei (Martz
& Small, 2006:fig. 3) based on a curved horn-like structure and
a longer ventral flange in comparison to the dorsal flange.
However, differences in the lateral osteoderms lie in the pres­
ence of distinct ornamentation in the form of pits and ridges in
the case of Tecovasuchus in contrast to that of Venkatasuchus.
In the latter, minute pits and faint corrugations on the dorsal
flange are observed in a few osteoderms.
The dorsal paramedian osteoderms of Kocurypelta silvestris
(Czepiński et al., 2021:fig. 7A) differ from those of Venkatasu­
chus armatum in having nearly parallel ridges as ornamenta­
tion, absence of a mixed pattern comprising oblique and
radiating ridges along with subcircular to elongate pits as in
Venkatasuchus (Fig. 3), a strongly developed mediolaterally

FIGURE 10. Venkatasuchus armatum. Reconstruction of the partial carapace based on available paramedian and lateral osteoderms, A, position of
the osteoderms with respect to the entire carapace is shown in gray; B, restored paramedian and associated lateral osteoderms (in successive order).
Arrow indicates anterior end. 1, ISIR267/7A, ISIR267/3B; 2, ISIR267/6A, ISIR267/1B; 3, ISIR267/5A, ISIR267/2B; 4, ISIR267/1A, ISIR267/7B; 5,
ISIR267/4A, ISIR267/4B; 6, ISIR267/2A, ISIR267/6B; 7, ISIR267/3A, ISIR267/5B. Scale bar equals 100 mm.
 Haldar et al. —Typothoracine from India (e2253292-14)
expanded posterodorsal ridge, and wide, incised beveled
flange. On the other hand, the lateral osteoderms associated
with the trunk paramedian osteoderms of Kocurypelta (Cze­
piński et al., 2021:fig. 7D) differ from those of Venkatasuchus
(Fig. 7) based on the presence of prominent radial ridges
and an almost circular lateral flange.
Hence, a combination of multiple diagnostic features defines
the lower Dharmaram aetosaur. In addition, reconstruction
shows a wide discoidal carapace (Fig. 10) where the trunk ver­
tebrae, ranging from the anterior to mid-trunk region, were
covered by at least seven exceptionally wide paramedian osteo­
derms having a narrow anterior bar, mediolaterally extended
low dorsal eminence at the center of ossification and offset medi­
ally, ridges and alternating grooves, along with circular to ellipti­
cal pits radiate from the dorsal eminence, a prominent and thick
posterodorsal ridge in continuation with the dorsal eminence and
parallel to the posterior margin, thick beveled posterior margin,
wide arching at the center of ossification, and a transverse ventral
strut. The associated lateral osteoderms were flat, horn-shaped,
asymmetric with short dorsal and long ventral flanges, often
faintly striated and have an acute angle of flexure. All these fea­
tures imply that the specimens of ISIR267 may be assigned to a
new taxon, Venkatasuchus armatum, belonging to the clade
Typothoracinae (sensu Parker, 2016a) of the aetosaurian family
Stagonolepididae (sensu Heckert & Lucas, 2000).
The single isolated left paramedian osteoderm, ISIR286,
recovered from the same locality as the holotype material
(ISIR267) is assigned to Typothoracinae (Parker, 2007, 2016a)
based on a high W:L ratio, which is comparable with the dorsal
paramedian osteoderms of Typothorax, Paratypothorax, and
Tecovasuchus (Heckert & Lucas, 2000; Hunt & Lucas, 1992;
Long & Ballew, 1985; Martz, 2002; Martz & Small, 2006).
ISIR286 is referred to Venkatasuchus armatum based on
similar W:L ratio, medially offset center of ossification, ornamen­
tation pattern on the dorsal surface, and beveled posterior edge
(Table 2; Figs. 3, 8B–H). However, there are several features that
distinguish ISIR286 from ISIR267 such as a reduced dorsal emi­
nence, a relatively narrow posterodorsal ridge, pronounced
ridges and grooves radiating from the ossification center
towards the lateral side, near absence of ornamentation,
especially incised circular to elliptical pits and radiating ridges
on the medial side, absence of a pronounced anterolateral pro­
jection and a broadly convex and indistinct ventral strut. These
areas of variation within the osteoderms could be due to intras­
pecific variations such as positional/regional variation on the car­
apace, ontogeny, individual and sexual dimorphism (Desojo &
Ezcurra, 2011; Paes-Neto et al., 2021b; Taborda et al., 2015).
As a result, ISIR286 is considered as belonging to another indi­
vidual of Venkatasuchus, until more material is recovered.
Phylogenetic Perspectives
Interspecific variation within the Aetosauria has been widely
studied (Harris et al., 2003; Heckert & Lucas, 1999, 2000;
Hoffman et al., 2018; Long & Murry, 1995; Paes-Neto et al.,
2021b; Parker 2007, 2016a, b; Parrish, 1994; Reyes et al., 2021;
Schoch & Desojo, 2016). Three major clades, Stagonolepididae,
Aetosaurinae, and Stagonolepidoidea, were identified within
Aetosauria with Aetosauroides scagliai as the earliest-diverging
taxon (Da-Silva et al., 2014; Desojo et al., 2012; Heckert &
Lucas, 2000; Paes-Neto et al., 2021a; Parker, 2016a; Reyes
et al., 2021). Analysis by Parker (2007) recovered well-resolved
Desmatosuchinae and Typothoracinae with the Aetosaurinae
being weakly supported. Since then several analyses (Desojo &
Ezcurra, 2011; Desojo et al., 2012; Heckert et al., 2015) have
expanded on the matrix compiled by Parker (2007) by adding
several new taxa and characters resulting in various phylogenetic
hypotheses. Parker (2016a) recognized five major clades,
Stagonolepididae, Aetosaurinae, Stagonolepidinae, Desmatosu­
chinae, and Typothoracinae, based on a comprehensive analysis
involving a maximum number of cranial, dermal, and endoskele­
tal (non-osteoderm) postcranial characters. This was supported
by later studies (Hoffman et al., 2018; Schoch & Desojo, 2016).
Recently Reyes et al. (2021) and Paes-Neto et al. (2021a) differ­
entiated two major clades within Stagonolepidae, the Stagonole­
pidoidea and Aetosaurinae, and added 16 new cranial characters
and modified a few existing characters from the data matrix com­
piled by Parker (2016a).
In the current study, the positions of Aetosaurus ferratus, Coa­
homasuchus kahleorum, and Coahomasuchus chathamensis are
well resolved (Fig. 9C), and these are recovered as early-diver­
ging taxa within the clade Stagonolepididae (Node B) and Aeto­
saurinae (Node F), respectively. The latter also included
Typothoracinae (Node G) similar to the previous studies by
Parker (2016a), Reyes et al. (2021), and Paes-Neto et al.
(2021a). Aetosaurinae is the stem-based clade that contains all
taxa more closely related to Aetosaurus ferratus than to the last
common ancestor of Aetosaurus and Desmatosuchus where
Aetosaurus is an early-diverging taxon and lying at the base of
Aetosauria (Heckert & Lucas, 1999). The present study corrobo­
rates Heckert and Lucas (1999), Parker (2007, 2016a), and
Parker et al. (2008), which recovered Aetosaurinae as a more
inclusive clade including all non-Desmatosuchines with Coaho­
masuchus and Aetosaurus at its base. Aetosaurus ferratus is con­
sidered as immature as evidenced from histological study and
hypothesized as a juvenile form of Paratypothorax based on
close resemblance of the cranial anatomy of the two taxa
(Scheyer et al., 2014; Schoch & Desojo, 2016). On the other
hand, marked differences in their dorsal trunk osteoderms and
their co-occurrence in the same middle Norian horizon of
Germany raise two distinct possibilities: (i) Aetosaurus ferratus
is a juvenile of a close relative of Paratypothorax or (ii) the
former represents the juvenile stage of Paratypothorax
(Scheyer et al., 2014; Schoch and Desojo, 2016). However,
most of the phylogenetic analyses, including the current study,
retain Aetosaurus ferratus and Paratypothorax as two distinct
taxa within Aetosaurinae, where A. ferratus is found to be an
early-diverging taxon (e.g., Paes-Neto et al., 2021a; Parker,
2016a; Reyes et al., 2021; Schoch & Desojo, 2016).
Typothoracinae (Node G) is well resolved and supported by a
GC value of 80. This clade was first described by Parker (2007)
containing all taxa closely related to Typothorax and Paraty­
pothorax than to Stagonolepis, Aetosaurus, and Desmatosuchus.
The current study recovered Apachesuchus heckerti as an early-
diverging taxon within Typothoracinae as observed in previous
studies (Parker, 2016a, Reyes et al., 2021). Parker (2007)
placed Rioarribasuchus, Paratypothorax, and Tecovasuchus
within the clade Paratypothoracisini, which is nested within
Typothoracinae; this clade was later emended as Paratypothora­
cini (Parker, 2016a). Paratypothoracini (Node H, GC value = 90,
Fig. 9C) comprises (Rioarribasuchus chamaensis +(Paraty­
pothorax sp. + Paratypothorax andressorum) + (Tecovasuchus
chatterjeei + (Venkatasuchus armatum + Kocurypelta silvestris)))
and is defined by seven unambiguous synapomorphies (Sup­
plementary File 5). The new aetosaur from India, Venkatasuchus
armatum, is deeply nested within Typothoracinae and Paraty­
pothoracini (Fig. 9C) and is recovered as a sister taxon to Kocur­
ypelta silvestris (Czepiński et al., 2021) based on three
synapomorphic characters (Supplementary File 5).
Although synapomorphies define Stagonolepidoidea (Sup­
plementary File 5), the GC value is low (= 15) and the clade exhi­
bits distinct differences in topology with respect to previous
literature. These include recovery of Stenomyti huangae at the
base, contrary to Parker (2016a), Hoffman et al. (2018), Reyes
et al. (2021), and Paes-Neto et al. (2021a). However, this study
recovers Neoaetosauroides engaeus, Adamanasuchus
 Haldar et al. —Typothoracine from India (e2253292-15)
eisenhardtae, Calyptosuchus wellesi, Scutarx deltatylus, and Sta­
gonolepis olenkae within Desmatosuchinae, similar to the ana­
lyses performed by Parker (2016a), Reyes et al. (2021), and
Paes-Neto et al. (2021a). Adamanasuchus is in a sister taxa
relationship with Scutarx and Calyptosuchus with a low GC
value of 13 and based on the width:length ratio (3.0–3.5) of the
widest paramedian osteoderm dorsal to the trunk vertebrae
(character 64). The clade Desmatosuchini is recovered here as
deeply nested within Desmatosuchinae, and comprises several
taxa as suggested by Harris et al. (2003), Parker (2007), and
Parker (2016a). The clade Desmatosuchini is highly robust with
GC = 98, and defined by 11 unambiguous synapomorphies (Sup­
plementary File 5). Hence, the current study reiterates the robus­
ticity of the aetosaurian clades Typothoracinae and
Desmatosuchini, which are the most stable and well supported
(GC values > 80) within Aetosauria. The new taxon, Venkatasu­
chus armatum, is recovered as deeply nested within Paraty­
pothoracini (GC value = 90) as a late-diverging taxon with
respect to (Paratypothorax sp. + Paratypothorax andressorum)
+ Tecovasuchus chatterjeei)), sister taxon to Kocurypelta silvestris
(GC value = 91), and is defined by several autapomorphic
characters.
BIOSTRATIGRAPHIC SIGNIFICANCE
Highly diverse and rich aetosaurian fossil records are known
from different Upper Triassic horizons of North America
(Heckert & Lucas, 1999, 2000; Long & Ballew, 1985; Long &
Murry, 1995; Lucas et al., 2006; Martz et al., 2013; Parker, 2007,
2008, 2016b; Walker, 1961) and South America (Argentina and
Brazil) (Bonaparte, 1969; Casamiquela, 1960; Da-Silva et al.
2014; Desojo & Ezcurra, 2011; Desojo et al., 2012, 2013;
Heckert et al., 2021; Paes-Neto et al., 2021a). In comparison,
aetosaurs are scarce in other parts of the world and are currently
reported from Scotland, Poland, Germany, Greenland, Morocco,
and India (Burmeister, 2000; Czepiński et al., 2021; Dróżdż, 2018;
Jalil et al., 1995; Kutty & Sengupta, 1989; Lucas et al., 2006; Sulej,
2010; Teschner et al., 2022). Well established radioisotopic and
magnetostratigraphic data are available for many of the strati­
graphic units encompassing the fossiliferous horizons (Abdala
& Rebeiro, 2010; Irmis et al., 2010, 2011); however, such data
are lacking from many of the Gondwanan strata. Previous
studies have correlated several Upper Triassic horizons of
North America based on the lowest and highest occurrences of
the aetosaurs, phytosaurs, and metoposaurids (Lucas et al.,
1998, 2010; Parker & Martz, 2011). The Late Triassic estimated
Land Vertebrate teilzones and holochronozones are based on
phytosaur biostratigraphy (Martz & Parker, 2017). Aetosaur
fossils are extremely common, and in some Upper Triassic
localities of North America even more than the phytosaurs and
their unique osteoderms exhibit a great taxonomic utility
(Parker & Martz, 2017). This is more well established for the
osteoderms of Typothoracine and Desmatosuchine aetosaurs
(Desojo et al., 2013; Lucas et al., 2006; Martz & Small, 2006;
Parker, 2007, 2008, 2016a) than the early-diverging forms, as
most of the osteodermal characters are plesiomorphic for the
clade (Desojo & Ezcurra, 2011; Desojo et al., 2013; Heckert
et al., 2017, 2021; Paes-Neto et al., 2021b; Parker, 2016a;
Parker & Martz, 2017; Small & Martz, 2013). Hence, our study
expands the understanding of aetosaur biostratigraphy based
on their lowest and highest occurrences in conjunction with
those of the phytosaur taxa.
The Santa Rosa Formation of the Dockum Group in the Texas
Panhandle and northeastern New Mexico is correlated with the
Shinarump Member and upper part of the Mesa Redondo
Member of the Chinle Formation in northern Arizona
(Heckert & Lucas, 1999; Long & Murry, 1995; Lucas, 2018;
Parker & Martz, 2010). U-Pb geochronology suggests a
depositional age of 228–225 Ma for the Mesa Redondo and Shi­
narump members of the Chinle Formation, thus providing an
approximate age for the Santa Rosa Formation (Ramezani
et al., 2014; Rasmussen et al., 2021). Several aetosaurian taxa
are known from the Colorado City Member (= lower portion
of the Cooper Canyon Formation), Dockum Group, Texas such
as Coahomasuchus kahleorum, Longosuchus meadei, and Luca­
suchus hunti, suggesting that the Desmatosuchines first appeared
during the early Norian in the southwestern U.S.A. (Heckert &
lucas, 1999; Parker and Martz, 2010). However, the documen­
tation of Gorgetosuchus and Coahomasuchus chathamensis in
the Pekin Formation of the Newark Supergroup suggests that
the Desmatosuchines and Aetosaurines were present during
the late Carnian (Fig. 11). Paleomagnetostratigraphic correlation
of the Pekin Formation with the other Newark Supergroup sec­
tions suggests an age of 231 Ma (Heckert et al., 2015, 2017;
Whiteside et al., 2011). In addition, the Blue Mesa and lower
Sonsela members of the Chinle Formation in northern Arizona
and the Los Esteros Member of the Santa Rosa Formation, the
Tecovas Formation, the lower Trujilo and the lower Cooper
Canyon formations of the Dockum Group in northeastern New
Mexico and Western Texas, respectively, are correlated based
on Malerisaurus-like azendohsaurids and non-mystriosuchine
leptosuchomorph phytosaurs (Martz & Parker, 2017; Nesbitt
et al., 2022), and are considered to be early to middle Norian
(225–214 Ma; Ramezani et al., 2011, 2014). The upper Blue
Mesa Member of Chinle Formation, Arizona and the Tecovas
Formation of Dockum Group, Texas have yielded several aeto­
saurian taxa including Tecovasuchus chatterjeei (Martz & Small,
2006; Parker, 2016a), which currently represents the earliest
occurrence of Paratypothoracini and also could tentatively be
the oldest typothoracine. On the other hand, the highest strati­
graphic occurrences of Desmatosuchinae aetosaurs are from
the Middle unit of the Cooper Canyon Formation (e.g., Post
Quarry) of Western Texas and the Martha Butta Beds of the
Sonsela Member (Lucas, 2010, 2018; Martz & Parker, 2017;
Parker, 2016b; Parker & Martz, 2011). Typothoracinae aetosaurs
such as Redondasuchus rineharti and Apachesuchus heckerti are
also known from the Redonda Formation of New Mexico, the
age of which is latest Norian-Rhaetian (Martz et al., 2013; Spiel­
mann & Lucas, 2012). Hence, the highest and lowest occurrences
of typothoracine aetosaurs mark an early Norian to Rhaetian
age.
In South America, Hyperodapedon-dominated beds of the
lower Ischigualasto Formation, Argentina, exhibit a U-Pb
depositional age of 229.20 + 0.11/−0.15 to 227.94 + 0.83/−1.67
Ma (Desojo et al., 2020). The early-diverging aetosaur, Aetosaur­
oides scagliai, is known from the lower levels of the Ischigualasto
Formation in Argentina and the Santa Maria Formation of Brazil
(Desojo & Ezcurra, 2011; Paes-Neto et al., 2021a) suggesting the
age of the Santa Maria Formation to be late Carnian (Casami­
quela, 1960, 1961; Desojo & Ezcurra, 2011; Martínez et al.,
2011; Paes-Neto et al., 2021a, b). Desojo et al. (2020) and Paes-
Neto et al. (2021a) correlate the Hyperodapedon-dominated
horizon in Argentina with Hyperodapedon Assemblage Zone
of the Santa Maria Formation of Brazil. The latter has yielded
two aetosaurs, Aetosauroides scagliai and Aetobarbakinoides
brasiliensis (Desojo et al., 2012, 2013; Langer, 2005). Neoaeto­
sauroides engaeus is known from the Upper part of Los Colora­
dos Formation overlying the Ischigualasto Formation in
Argentina, which is early–middle Norian based on magnetostra­
tigraphic correlation (Bonaparte, 1969; Desojo & Báez, 2005,
2007; Desojo et al., 2013). Hence, the Carnian-Norian boundary
marks the highest stratigraphic occurrence of a few early-diver­
ging aetosaurs such as Aetosauroides scagliai and Aetobarbaki­
noides brasiliensis (Fig. 11).
Aetosaurs are known from the Irohalene Member of the Time­
zgadiouine Formation of Morocco and the Isalo II beds of
 Haldar et al. —Typothoracine from India (e2253292-16)

FIGURE 11. Time calibrated phylogenetic tree showing ranges of the aetosaur taxa. Ranges of the aetosaurs (boxes) are given by different colors to
show their region of occurrences: black, Laurasia; red, Gondwana; blue, both Laurasia and Gondwana. Abbreviations: Alau, Alaunian; AZ, Assem­
blage Zone; Cord, Cordevolian; CPE, Carnian Pluvial Event; F, Formation; Hyp, Hyperodapedon; Seva, Sevatian; Tey, Teyumbaita; Tuva, Tuvalian;
Sources of information: Long & Murry (1995), Lucas et al. (2006), Parker & Martz (2011), Desojo et al. (2013), Parker (2016b), Martz & Parker (2017).

Madagascar, from where members of Paratypothoracini (Jalil et
al., 1995; Lucas et al. 2006) and Desmatosuchinae (Burmeister,
2000) are known, respectively. Based on other associated fauna
(e.g., phytosaur Parasuchus, rhynchosaur Hyperodapedon, and
metoposaurid temnospondyls), these horizons have been corre­
lated with the lower Ischigualasto Formation of Argentina and
lower Maleri Formation of India and is therefore considered as
late Carnian (Datta & Ray, 2023; Parker & Martz, 2017). In
addition, the Lossiemouth Sandstone Formation of Scotland,
and the Krasiejów locality in Poland have yielded Stagonolepis
olenkae, Stagonolepis robertsoni, and Aetosaurus ferra­
tus (Antczak, 2016; Dróżdż, 2018) along with the rhynchosaur
Hyperodapedon and phytosaur Parasuchus resulting in the corre­
lation of these strata with the Hyperodapedon Assemblage Zone
of the Santa Maria Formation, Brazil, and lower Ischigualasto For­
mation of Argentina, Middle Wolfville Formation, Nova Scotia,
and Popo Agie Formation, Wyoming of North America, and the
lower Maleri and Tiki formations of India (Datta et al., 2021;
Fitch et al., 2023; Langer et al., 2007; Martínez et al., 2011), indicat­
ing a range from late Carnian to earliest Norian. It is evident that
most of the Upper Triassic aetosaur-bearing horizons have also
yielded rhynchosaurs (Hyperodapedon), phytosaurs, and meto­
posaurids, although not exclusively. Langer et al. (2017) suggested
a late Carnian–earliest Norian age for the horizons where Hyper­
odapedon is abundant.
Previous studies showed that the vertebrate fossil record of the
lower Dharmaram Formation of India is archosaur-dominated
and includes an undescribed phytosaur, aetosaurs, a sauropodo­
morph Jaklapallisaurus asymmetrica, and an indeterminate
neotheropod (Bandyopadhyay, 2011; Bandyopadhyay & Ray,
2020; Kutty et al., 2007; Novas et al., 2011). Currently, there is
no record of Hyperodapedon, and/or metoposaurids from the
lower Dharmaram Formation. Bandyopadhyay and Ray (2020)
correlated the lower Dharmaram Formation with the mid- to
late Norian lower Stubensandtein Formation (Löwenstein) of
Germany based on the report of a Nicrosaurus-like phytosaur,
though the latter is yet to be described in detail. On the other
hand, the upper Maleri Formation, underlying the lower Dhar­
maram Formation (Fig. 1), contains a distinctive fossil fauna
comprising chigutisaurid temnospondyls (Kutty & Sengupta,
1989) and several sauropodomorphs (Novas et al., 2011),
though Hyperodapedon and metoposaurids are absent. Based
 Haldar et al. —Typothoracine from India (e2253292-17)

FIGURE 12. Late Triassic Pangean map (after Scotese, 2001) showing distribution of aetosaurs, phytosaurs, and metoposaurids. The white arrows
show the possible paleomigratory routes of the terrestrial Typothoracine aetosaurs. 1, Southwestern North America; 2, Eastern North America; 3–
5, Central Europe; 6, Greenland; 7, Morocco; 8, Algeria; 9, Turkey; 10, Argentina; 11, Zimbabwe; 12, Madagascar; 13, Rewa Basin, India; 14, Pran­
hita-Godavari Basin, India; 15, Brazil; 16, Chile; 17, Italy. Sources of information: Long & Murry (1995), Heckert & Lucas (1999), Lucas et al. (2006),
Desojo et al. (2013), Martz et al. (2013), Brusatte et al. (2015), Parker (2016a, b), Lucas (2018, 2020), Datta et al. (2021), Datta & Ray (2023).

on the resemblance of the lower Dharmaram Formation with a AETOSAUR DIVERSIFICATION AND
post-Ischigualastian assemblage, the horizon was correlated PALEOBIOGEOGRAPHY
with the Los Colorados Formation, the Trossingen Formation
Aetosaurian taxa are mostly known from Laurasian regions
of the German Keuper, the lower Elliot Formation, and the
(Fig. 11) of North America and Europe (Desojo et al., 2013).
Chinle Formation, and its age was considered as latest Norian–
Except for South America in the west, aetosaurs are poorly rep­
Rhaetian (Bandyopadhyay, 2011; Bandyopadhyay & Ray,
resented in central and eastern Gondwana, with no reports from
2020). However, presence of a new member of Paratypothoracini
Antarctica and Australia (Desojo et al., 2013; Lucas et al., 2006;
(current study) suggests an early/middle Norian–Rhaetian age
Parker, 2016b; Reyes et al., 2021). The oldest record of the Aeto­
for the lower Dharmaram Formation, though more study is
sauria is that of the early-diverging taxon Aetosauroides scagliai
required, especially on the undescribed Desmatosuchus-like
known from late Carnian strata of the Ischigualasto Formation of
aetosaurs and Nicrosaurus-like phytosaurs for precise global cor­
Argentina and Hyperodapedon Assemblage Zone, Santa Maria
relation. Additionally, the overlap between a Paratypothorax-
Formation of Brazil (Paes-Neto et al., 2021a). In most of the
like and Desmatosuchus-like taxon suggest at least a mid-
Upper Triassic horizons, aetosaurs un-exclusively co-existed
Norian age based on the refined biostratigraphy of the Chinle
with the rhynchosaurs, phytosaurs, and metoposaurids, especially
Formation and Dockum Group, U.S.A. (e.g., Sonsela Member
during the late Carnian (Datta et al., 2021; Langer, 2005; Lucas,
of Chinle Formation of northern Arizona and western New
2018). It may be hypothesized that the Carnian Pluvial Event
Mexico; Tecovas Formation and Trujilo Formation of Dockum
(CPE, 234–232 Ma; Dal Corso et al., 2020; Desojo et al., 2013)
Group of northeastern New Mexico and Texas Panhandle;
marking a phase of increased precipitation and humidity
middle Cooper Canyon Formation of Dockum Group of
during the Late Triassic (Benton et al., 2018; Dal Corso et al.,
western Texas) which are in similar paleolatitudes to these taxa
2020; Griffin et al., 2022; Tanner, 2000, 2003) was instrumental
of India.
 Haldar et al. —Typothoracine from India (e2253292-18)

in the initial aetosaurian radiation as hypothesized for metopo­ as conducive environmental and ecological conditions supported
saurid temnospondyls (Buffa et al., 2019; Lucas, 2020) and phyto­ dispersal of tetrapods from the Laurasian and Gondwanan pro­
saurs (Datta & Ray, 2023). Many of the early-diverging taxa vinces to India, and vice versa. In contrast, the restricted occur­
including Aetosauroides and the stagonolepidoid such as Gorge­ rence of Laurasian fauna in other Gondwanan provinces may
tosuchus are absent from the fossil record after the Carnian– be because of topographic, environmental or climatic barriers,
Norian boundary (Fig. 11). Typothoracinae likely originated and requires further study.
during the early Norian as suggested from their lack of fossil
record during the Carnian, and diversified during the mid-to-
late Norian to early Rhaetian. This is best exemplified in the CONCLUSIONS
southwestern U.S.A. (Heckert et al., 2010; Lucas et al., 2006;
1. Examination of multiple isolated and partially articulated
Martz & Small, 2006; Spielmann et al., 2006), particularly
left paramedian and lateral osteoderms recovered from a
within the Petrified Forest National Park, Arizona (Reyes
single locality reveals a new typothoracine aetosaur, Venka­
et al., 2021). On the other hand, stagonolepidoids originated
tasuchus armatum, from the Upper Triassic of India. The
during the late Carnian, crossed the Carnian–Norian boundary
taxon is characterized by wide paramedian osteoderms
and diversified during early-mid Norian (Fig. 11). This pattern
dorsal to the trunk region that comprise a narrow, weakly
of aetosaurian diversification is also corroborated by Reyes
raised anterior bar with distinct anterolateral projection,
et al. (2021).
mediolaterally extended dorsal eminence offset medially
The Late Triassic distribution of the fossil localities of aeto­
from the center of ossification, ornamentation composed of
saurs (Fig. 12) shows distinct clustering comprising: (i) low paleo­
randomly arranged pits and radiating ridges alternating
latitudinal regions of Laurasia (North America) and Gondwanan
with grooves, a thick, transverse posterodorsal ridge,
region of Morocco from where both early- and late-diverging
beveled posterior margin, weakly to well-developed ventral
forms have been reported; and (ii) at high paleolatitude in the
strut, and a wide flexion at the center of ossification. The
(a), northern hemisphere (Europe) from where late-diverging
lateral osteoderms, on the other hand, are asymmetric,
forms are known, and (b) southern hemisphere encompassing
have short and long dorsal and ventral flanges, and a
South America (early-diverging forms). In addition, these
curved horn-like structure where the two flanges meet at
localities correlate with those in which phytosaurs and metopo­
an acute angle. Faint ornamentations in the form of radiating
saurid temnospondyls are highly abundant (Fig. 12), though aeto­
ridges and a few circular pits especially on the dorsal flange
saurs are relatively scarce in the Gondwanan regions (Fig. 11)
are visible.
and restricted to South America (Chile, Brazil, and Argentina;
2. The partial carapace of V. armatum is reconstructed as
Desojo et al., 2013), except for the current study from India.
strongly discoidal based on the curvature of the paramedian
However, site inaccessibility and collection bias cannot be
osteoderms as well as their high width:length ratio with the
ruled out. Moreover, Typothoracinae are reported, but poorly
widest one positioned dorsal to the mid-trunk vertebra.
documented from Morocco (Lucas et al., 2006) and India
Other osteoderms are arranged according to the degree of
(current study), though these were prolific in the Laurasian pro­
sculptured ornamentation, prominence of dorsal eminence,
vinces, especially in the southwestern U.S.A. and Europe
and presence of a strongly or weakly developed ventral
(Desojo et al., 2013; Heckert et al., 1996; Hunt & Lucas, 1990,
strut. Asymmetric lateral osteoderms with acute flexion are
1992; Lucas et al., 2006; Martz & Small., 2006; Schoch &
considered as pre-caudals with the angle of flexion decreasing
Desojo, 2016). Additionally, desmatosuchine-like taxa are
posteriorly along with strongly unequal flanges.
reported from India, though this material is yet to be studied in
3. Phylogenetic analysis recovered Venkatasuchus as a late
detail.
diverging taxon within the clades of Typothoracinae and Para­
Previous studies have proposed a mixed faunal affinity for
typothoracini, and a sister taxon to Kocurypelta silvestris. Ven­
India, which included both Gondwanan and Laurasian fauna
katasuchus is defined by several autapomorphic characters
(Chatterjee et al., 2017). During the Late Triassic, a strong Laur­
involving paramedian osteoderms dorsal to the trunk ver­
asian affinity can be seen for the semi-aquatic phytosaurs (Datta
tebrae such as dorsal surface ornamented by large, irregular
& Ray, 2023) and aquatic metoposaurids (Sengupta, 2002)
pits surrounding the dorsal eminence and radiating ridges in
known from India because of their possible migration along
other areas, straight anterior margin of the anteromedial
the circum-Tethyan shoreline. Such dispersal may be hypoth­
corner of the anterior bar in dorsal view, raised or ridged
esized from increased waterways because of the rifting of
and ornamented posteromedial corner.
Pangea (Datta & Ray, 2023). For the lower Dharmaram fauna,
4. The similarity of the lower Dharamaram vertebrate fauna
the association of a Nicrosaurus-like phytosaur (Bandyopadhyay
with that of other coeval horizons around the world indicates
& Ray, 2020) and a member of Paratypothoracini (current study)
a mid-Norian–Rhaetian age. Typothoracine aetosaurs were
shows a similar Laurasian faunal influx whereas the dinosaurian
relatively scarce in the Gondwanan region during the Late
fauna (Novas et al., 2011) points towards Gondwanan influence
Triassic with the early-diverging forms known essentially
during early/middle Norian–Rhaetian. Such mixed (Laurasian
from South America. Thus, the recovery of Venkatasuchus
and Gondwanan) faunal association is also evident from the
marks the first record of the late-diverging Paratypothoracini
fossil tetrapods of the Tiki Formation (late Carnian- to early/
from the Gondwanan region.
middle Norian) of the Rewa Gondwana Basin, and the Maleri
5. The current study proposes a strong Laurasian faunal influx
Formation (late Carnian to late Norian) of the Pranhita-Goda­
irrespective of aquatic or terrestrial animals in peninsular
vari Basin (Bandyopadhyay & Ray, 2020; Chatterjee et al.,
India during the Late Triassic suggesting possible land
2017). Thus, India exhibits a unique mixed vertebrate faunal
bridges and conducive environmental conditions.
association that persisted throughout the entire span of the
Late Triassic (late Carnian to Rhaetian), irrespective of
whether the animals were terrestrial or semi-aquatic. In a
ACKNOWLEDGMENTS
recent study on dispersal of early-diverging dinosaurs, Griffin
et al. (2022) suggested that dispersal of the dinosauromorphs The fossil specimens were collected by T. S. Kutty, Geological
may be attributed to relaxation of paleolatitude-driven climatic Studies Unit, Indian Statistical Institute, Kolkata. We thank
barriers during the Late Triassic. Hence, it is hypothesized that D. Mukherjee of the same Institute, and D. Datta, Department
possible land bridges and/or absence of physical barrier as well of Earth Sciences, Indian Institute of Technology, Roorkee for
 Haldar et al. —Typothoracine from India (e2253292-19)
advice and help throughout the study. We thank A. Huttenlocker,
T. Sulej, P. L. Godoy, and an anonymous reviewer for thorough
revision and constructive suggestions. D. Pradhan and
L. Mahankur are gratefully acknowledged for preparation and
removal of matrices from the fossil specimens. The financial assist­
ance was provided by Science and Engineering Research Board,
India (SERB/CRG/000388) to SR. The Indian Institute of Tech­
nology Kharagpur and Indian Statistical Institute Kolkata are
acknowledged for providing infrastructural facilities to AH and
SR, and SB, respectively.
DISCLOSURE STATEMENT
No potential conflict of interest was reported by the authors.
AUTHOR CONTRIBUTIONS
SR, SB conceptualized and designed the project, AH, SR col­
lected and analyzed data, AH drafted the manuscript, SR carried
out interpretative line drawings, acquired funding and adminis­
tered the project. All authors edited the manuscript.
SUPPLEMENTARY FILES
Supplementary File 1.docx: Details of the characters used for
the phylogenetic analysis.
Supplementary File 2.nex: Combined characters and data
matrix used for phylogenetic analysis.
Supplementary File 3.docx: Resampled MPTs (ordered).
Supplementary File 4.tre: MPTs of the parsimony analysis
(unordered).
Supplementary File 5.docx: List of synapomorphies.
ORCID
Atrayee Haldar http://orcid.org/0000-0001-9031-4174
Sanghamitra Ray http://orcid.org/0000-0002-9231-3498
Saswati Bandyopadhyay http://orcid.org/0000-0001-8273-3184
LITERATURE CITED
Abdala, F., & Rebeiro, A. M. (2010). Distribution and diversity patterns
of Triassic cynodonts (Therapsida, Cynodontia) in Gondwana.
Palaeogeography, Palaeoclimatology, Palaeoecology, 286, 202–217.
Agassiz, L. (1844). Monographie des poisons fossils du Vieux Grés
Rouge ou Systéme Dévonien (Old Red Sandstone) des Iles
Britanniques ed de Russie. Jent et Gassman, Neuchâtel, 171 pp.
Antczak, M. (2016). Late Triassic aetosaur (Archosauria) from Krasiejów
(SW Poland): new species or an example of individual variation?
Geological Journal, 51(5), 779–788.
Bandyopadhyay, S. (2011). Non-marine Triassic vertebrates of India; 33–
46 in J.O. Calyo, J. Porifiri, B. González, and D. D. Santos (eds.),
Paleontología y Dinosarios desde América Latina. EDIUNCE,
Editorial de la Universidad Nacional de Cuyo, Mendoza,
Argentina.
Bandyopadhyay, S., & Ray, S. (2020). Gondwana vertebrate faunas of
India: their diversity and intercontinental relationships. Episodes
Journal of International Geoscience, 43(1), 438–460.
Bandyopadhyay, S., & RoyChowdhury, T. K. (1996). Beginning of the
continental Jurassic in India—a palaeontological approach.
Museum of Northern Arizona Bulletin, 60, 371–378.
Bandyopadhyay, S., & Sengupta, D. P. (2006). Vertebrate faunal turnover
during the Triassic-Jurassic transition: an Indian scenario. New
Mexico Museum of Natural History and Science Bulletin, 37, 77–85.
Benton, M. J., Bernardi, M., & Kinsella, C. (2018). The Carnian Pluvial
Episode and the origin of dinosaurs. Journal of the Geological
Society, 175(6),1019–1026.
Bonaparte, J. F. (1969). Dos nuevas “faunas” de reptiles Triásicos de
Argentina; 283–306 in A. J. Amos (ed.), Gondwana Stratigraphy,
IUGS Symposium, Buenos Aires, 1–15 October 1967. Paris:
United Nations Educational Scientific and Cultural Organization.
Brusatte, S. L., Butler, R. J., Mateus, O., & Steyer, J. S. (2015). A new
species of Metoposaurus from the Late Triassic of Portugal and
comments on the systematics and biogeography of metoposaurid
temnospondyls. Journal of Vertebrate Paleontology, 35(3),
e912988.
Buffa, V., Jalil, N. E., & Steyer, J. S. (2019). Redescription of
Arganasaurus (Metoposaurus) azerouali (Dutuit) comb. nov. from
the Upper Triassic of the Argana Basin (Morocco), and the first
phylogenetic analysis of the Metoposauridae (Amphibia,
Temnospondyli). Papers in Palaeontology, 5(4), 699–717.
Burmeister, K. C. (2000). Paleogeographic and biostratigraphic impli­
cations of new early Mesozoic terrestrial vertebrate fossils from the
Poamay site: central Morondava Basin, Madagascar [Doctoral dis­
sertation, University of California, Santa Barbara].
Casamiquela, R. M. (1960). Sobre dos nuevos estagonolepoideos argen­
tinos. Ameghiniana, 2(1), 3–9.
Casamiquela, R. M. (1961). Dos nuevos estagonolepoideos argentinos
(de Ischigualasto, San Juan). Revista de la Asociación Geológica
Argentina, 16(3–4), 143–204.
Casamiquela, R. M. (1980). Nota sobre restos de un reptil aetosauroideo
(Thecodontia, Aetosauria) de Quimal, Cordillera de Domeyko,
Antofagasta. Prueba de la existencia del Neotriásico continental
en los Andes del Norte de Chile. In Actas Congreso Argentino de
Paleontología y Bioestratigrafía (No. 2, pp. 135–142).
Case, E. C. (1920). Preliminary description of a new suborder of phyto­
saurian reptiles with a description of a new species of
Phytosaurus. The Journal of Geology, 28(6), 524–535.
Case, E. C. (1932). A perfectly preserved segment of the armor of a phy­
tosaur, with associated vertebrae. Contributions from the Museum
of Paleontology. University of Michigan, 4, 57–80.
Chakraborty, C., Mandal, N., & Ghosh, S. K. (2003). Kinematics of the
Gondwana basins of peninsular India. Tectonophysics, 377, 299–324.
Chatterjee, S. (1985). Postosuchus, a new thecodontian reptile from the
Triassic of Texas and the origin of tyrannosaurs. Philosophical
Transactions of the Royal Society of London. B, Biological
Sciences, 309(1139), 395–460.
Chatterjee, S., Scotese, C. R., & Bajpai, S. (2017). Indian plate and its epic
voyage from Gondwana to Asia: Its tectonic, paleoclimatic, and
paleobiogeographic evolution. Geological Society of America
Special Paper, 529, 1–147.
Cope, E. D. (1869). Synopsis of the extinct Batrachia, Reptilia, and Aves
of North America. Transactions of the American Philosophical
Society, 14, 1–252.
Cope, E. D. (1875). Report on the geology of that part of New Mexico
examined during the field season of 1875. Annual Report of the
Wheeler Survey, 61–116.
Czepiński, Ł., Dróżdż, D., Szczygielski, T., Tałanda, M., Pawlak, W.,
Lewczuk, A., … & Sulej, T. (2021). An Upper Triassic terrestrial
vertebrate assemblage from the forgotten Kocury locality
(Poland) with a new aetosaur taxon. Journal of Vertebrate
Paleontology, 41(1), e1898977.
Dal Corso, J., Bernardi, M., Sun, Y., Song, H., Seyfullah, L. J., Preto, N.,
Gianolla, P., Ruffell, A., Kustatscher, E., Roghi, G., Merico, A.,
Hohn S., Schimdt A. R., Marzoli, A., Newton R. J., Wignall, P. B.,
& Benton, M. J. (2020). Extinction and dawn of the modern world
in the Carnian (Late Triassic). Science Advances, 6(38), eaba0099.
Da-Silva, L. R, Desojo, J. B., Cabreira, S. F., Aires, A. S., Mueller, R. T.,
Pacheco, C. P., & Da-Silva, S. D. (2014). A new aetosaur from the
Upper Triassic of the Santa Maria Formation, southern Brazil.
Zootaxa, 3764(3), 240–278.
Dasgupta, S., Ghosh, P., & Gierlowski-Kordesch, E. H. (2017). A discon­
tinuous ephemeral stream transporting mud aggregates in a conti­
nental rift basin: the Late Triassic Maleri Formation, India.
Journal of Sedimentary Research, 87(8), 838–865.
Datta, D., & Ray, S. (2023). A giant phytosaur (Diapsida, Archosauria)
from the Upper Triassic of India with new insights on phytosaur
migration, endemism and extinction. Papers in Palaeontology, 9
(1), e1476.
Datta, D., Ray, S., & Bandyopadhyay, S. (2021). Cranial morphology of a
new phytosaur (Diapsida, Archosauria) from the Upper Triassic of
India: implications for phytosaur phylogeny and biostratigraphy.
Papers in Palaeontology, 7(2), 675–708.
Desojo, J. B., & Báez, A. M. (2005). The postcranial skeleton of
Neoaetosauroides (Archosauria: Aetosauria) from the Upper
Triassic of west-central Argentina. Ameghiniana, 42(1), 115–126.
 Haldar et al. —Typothoracine from India (e2253292-20)
Desojo, J. B., & Baez, A. M. (2007). Cranial morphology of the
Late Triassic South American archosaur Neoaetosauroides
engaeus: evidence for aetosaurian diversity. Paleontology, 50(1),
267–276.
Desojo, J. B., & Ezcurra, M. D. (2011). A reappraisal of the taxonomic
status of Aetosauroides (Archosauria, Aetosauria) specimens from
the Late Triassic of South America and their proposed synonymy
with Stagonolepis. Journal of Vertebrate Paleontology, 31(3), 596–
609.
Desojo, J. B., Ezcurra, M. D., & Kischlat, E. E. (2012). A new aetosaur
genus (Archosauria: Pseudosuchia) from the early Late Triassic of
southern Brazil. Zootaxa, 3166,1–33.
Desojo, J. B., Fiorelli, L. E., Ezcurra, M. D., Martinelli, A. G., Ramezani,
J., Da Rosa, Á., & Langer, M. C. (2020). The Late Triassic
Ischigualasto Formation at Cerro Las Lajas (La Rioja,
Argentina): fossil tetrapods, high-resolution chronostratigraphy,
and faunal correlations. Scientific Reports, 10(1), 1–34.
Desojo, J. B., Heckert, A. B., Martz, J. W., Parker, W. G., Schoch, R. R.,
Small, B. J., & Sulej, T. (2013). Aetosauria: a clade of armoured
pseudosuchians from the Upper Triassic continental beds.
Geological Society of London, Special Publications, 379(1), 203–239.
Dróżdż, D. (2018). Osteology of a forelimb of an aetosaur Stagonolepis
olenkae (Archosauria: Pseudosuchia: Aetosauria) from the
Krasiejów locality in Poland and its probable adaptations for a
scratch-digging behavior. PeerJ, 6, e5595.
Dzik, J., Sulej, T., Kaim, A., & Niedžwiedzki, R. (2000). Póznotriasowe
cmentarzysko ladowych czworonogów w Krasiejowie na Šlasku
Opolskim (Late Triassic graveyard of large Triassic tetrapods in
the Opole Silesia). Przeglacedild Geologiczny, 48(3), 226–235.
Fitch, A. J., Haas, M., C’Hair, W., Ridgley, E., Ridgley, B., Oldman, D.,
Reynolds C., & Lovelace, D. M. (2023). A New Rhynchosaur
Taxon from the Popo Agie Formation, WY: Implications for a
Northern Pangean Early-Late Triassic (Carnian) Fauna. Diversity,
15(4), 544.
Fraas, O. (1877). Aetosaurus ferratus Fraas, die gepanzerte Vogel-Echse
aus dem Stubensandstein bei Stuttgart. Wiirttemburgische
Naturwissenschaftliche, 33, 1–27.
Gauthier, J. A., & Padian, K. (1985). Phylogenetic, functional, and aero­
dynamic analyses of the origin of birds and their flight; 185–197 in
Hecht, M. K., Ostrom, H. M., Viohl, J. and Wellnhofer, P. (eds.),
The Beginnings of Birds. Proceedings of the International
Archaeopteryx Conference. Freundo des Jura Museums, Eichstatt,
Germany.
Goloboff, P. A., Farris, J. S., & Nixon, K. C. (2008). TNT, a free program
for phylogenetic analysis. Cladistics, 24(5), 774–786.
Gregory, J. T. (1953). Typothorax and Desmatosuchus. Postilla, 16, 1–27.
Griffin, C. T., Wynd, B. M., Munyikwa, D., Broderick, T. J., Zondo, M.,
Tolan, S., & Taruvinga, H. R. (2022). Africa’s oldest dinosaurs
reveal early suppression of dinosaur distribution. Nature, 609
(7926), 313–319.
Harris, S. R., Gower, D. J., & Wilkinson, M. (2003). Intraorganismal hom­
ology, character construction, and the phylogeny of aetosaurian
archosaurs (Reptilia, Diapsida). Systematic Biology, 52(2), 239–252.
Heckert, A. B., & Lucas, S. G. (1999). A new aetosaur (Reptilia:
Archosauria) from the Upper Triassic of Texas and the phylogeny
of aetosaurs. Journal of Vertebrate Paleontology, 19(1), 50–68.
Heckert, A. B., & Lucas, S. G. (2000). Taxonomy, phylogeny, biostratigra­
phy, biochronology, paleobiogeography, and evolution of the Late
Triassic Aetosauria (Archosauria: Crurotarsi). Zentralblatt für
Geologie und Paläontologie, Teil I, 1998(11–12), 1539–1587.
Heckert, A. B., & Lucas, S. G. (2002). South American occurrences of the
Adamanian (Late Triassic: latest Carnian) index taxon Stagonolepis
(Archosauria: Aetosauria) and their biochronological significance.
Journal of Paleontology, 76(5), 852–863.
Heckert, A. B., Hunt, A. P., & Lucas, S. G. (1996). Redescription of
Redondasuchus reseri, a Late Triassic aetosaur (Reptilia:
Archosauria) from New Mexico (USA), and the biochronology
and phylogeny of aetosaurs. Geobios, 29(5), 619–632.
Heckert, A. B., Fraser, N. C., & Schneider, V. P. (2017). A new species of
Coahomasuchus (Archosauria, Aetosauria) from the Upper Triassic
Pekin Formation, Deep River Basin, North Carolina. Journal of
Paleontology, 91(1), 162–178.
Heckert, A. B., Martínez, R. N., & Celeskey, M. D. (2021). Anatomical
details of Aetosauria (Archosauria: pseudosuchia) as revealed by
an articulated posterior skeleton from the Upper Triassic
Ischigualasto Formation, San Juan Province, Argentina.
Ameghiniana, 58(6), 464–484.
Heckert, A. B., Schneider, V. P., Fraser, N. C., & Webb, R. A. (2015). A
new aetosaur (Archosauria, Suchia) from the Upper Triassic Pekin
Formation, Deep River Basin, North Carolina, USA, and its impli­
cations for early aetosaur evolution. Journal of Vertebrate
Paleontology, 35(1), e881831.
Heckert, A. B., Lucas, S. G., Rinehart, L. F., Celeskey, M. D., Spielmann,
J. A., & Hunt, A. P. (2010). Articulated skeletons of the aetosaur
Typothorax coccinarum Cope (Archosauria: Stagonolepididae)
from the Upper Triassic Bull Canyon Formation (Revueltian:
early-mid Norian), eastern New Mexico, USA. Journal of
Vertebrate Paleontology, 30(3), 619–642.
Hoffman, D. K., Heckert, A. B., & Zanno, L. E. (2018). Under the armor:
X-ray computed tomographic reconstruction of the internal skel­
eton of Coahomasuchus chathamensis (Archosauria: Aetosauria)
from the Upper Triassic of North Carolina, USA, and a phyloge­
netic analysis of Aetosauria. PeerJ, 6, e4368.
Hunt, A. P., & Lucas, S. G. (1990). Re-evalution of “Typothorax” meadei,
a Late Triassic aetosaur from the United States. Paläontologische
Zeitschrift, 64(3), 317–328.
Hunt, A. P., & Lucas, S. G. (1991). Rioarribasaurus, a new name for a Late
Triassic dinosaur from New Mexico (USA). Paläontologische
Zeitschrift, 65(1), 191–198.
Hunt, A. P., & Lucas, S. G. (1992). The first occurrence of the aetosaur
Paratypothorax andressi (Reptilia, Aetosauria) in the western
United States and its biochronological significance.
Paläontologische Zeitschrift, 66(1), 147–157.
Irmis, R. B., Martz, J. W., Parker, W. G., & Nesbitt, S. J. (2010). Re-evaluat­
ing the correlation between Late Triassic terrestrial vertebrate biostra­
tigraphy and the GSSP-defined marine stages. Albertiana, 38, 40–52.
Irmis, R. B., Mundil, R., Martz, J. W., & Parker, W. G. (2011). High-res­
olution U–Pb ages from the Upper Triassic Chinle Formation (New
Mexico, USA) support a diachronous rise of dinosaurs. Earth and
Planetary Science Letters, 309(3–4), 258–267.
Jalil, N., Lucas, S. G., & Hunt, A. P. (1995). Biochronological significance
of aetosaurs and phytosaurs (Reptilia, Archosauromorpha) in the
Triassic Zarzaitine Series of Algeria. Neues Jahrbuch für Geologie
und Paläontologie-Monatshefte, 173–181.
Jenkins, F. A. Jr., Shubin N. H., Amaral W. W., Gatesy S. M., Schaff C. R.,
Clemmensen L. B., Downs W. R., Davidson A. R., Bonde N., &
Osbaeck F. F. (1994). Late Triassic continental vertebrates and
depositional environments of the Fleming Fjord Formation,
Jameson Land, East Greenland. Meddelelser om GrØnland
Geoscience, 32, 1–25.
Kutty, T. S., & Sengupta D. P., (1989). Late Triassic formations of the
Pranhita-Godavari valley and their vertebrate faunal sequence-a
reappraisal. Indian Journal of Earth Sciences, 16(4), 189–206.
Kutty, T. S., Jain S. L., & Chowdhury, T. R. (1987). Gondwana sequence of
the northern Pranhita–Godavari Valley: its stratigraphy and ver­
tebrate faunas. The Palaeobotanist, 36, 214–219.
Kutty, T. S., Chatterjee, S., Galton, P. M., & Upchurch, P. (2007). Basal
sauropodomorphs (Dinosauria: Saurischia) from the Lower
Jurassic of India: their anatomy and relationships. Journal of
Paleontology, 81(6), 1218–1240.
Langer, M. C. (2005). Studies on continental Late Triassic tetrapod bio­
chronology. II. The Ischigualastian and a Carnian global correlation.
Journal of South American Earth Sciences, 19(2), 219–239.
Langer, M. C., Ribeiro, A. M., Schultz, C. L., & Ferigolo, J. (2007). The
continental tetrapod-bearing Triassic of south Brazil. New Mexico
Museum of Natural History and Science Bulletin, 41, 201–218.
Langer, M. C., da Rosa, Á. A., & Montefeltro, F. C. (2017).
Supradapedon revisited: geological explorations in the Triassic of
southern Tanzania. PeerJ, 5, e4038.
Leiggi, P., & May, P. (eds.). 1994. Vertebrate Paleontological Techniques,
Vol. 1. Cambridge University Press, Cambridge, 344 pp.
Long, R. A., & Ballew K. L. (1985). Aetosaur dermal armor from the Late
Triassic of south western North America, with special reference to
material from the Chinle Formation of Petrified Forest National
Park; 45–68 in E. H. Colbert, and R. R. Johnson (eds.), The
Petrified Forest Through the Ages, 75th Anniversary Symposium
November 7, 1981. Museum of Northern Arizona Bulletin 54.
Long, R. A., & Murry P. A. (1995). Late Triassic (Carnian and Norian)
tetrapods from the southwestern United States. New Mexico
Museum of Natural History and Science Bulletin, 4, 1–254.
 Haldar et al. —Typothoracine from India (e2253292-21)
Lucas, S. G. (2010). The Triassic timescale based on nonmarine tetrapod
biostratigraphy and biochronology. Geological Society, London,
Special Publications, 334(1), 447–500.
Lucas, S. G. (2018). Late Triassic terrestrial tetrapods: Biostratigraphy,
biochronology and biotic events; 351–405 in L. H. Tanner (ed.),
The Late Triassic World, Springer International Publishing,
Switzerland, 805 pp.
Lucas, S. G. (2020). Biochronology of Late Triassic Metoposauridae
(Amphibia, Temnospondyli) and the Carnian pluvial episode.
Annales Societatis Geologorum Poloniae, 90, 409–418.
Lucas, S. G., Heckert, A. B., & Huber, P. (1998). Aetosaurus
(Archosauromorpha) from the Upper Triassic of the Newark
Supergroup, eastern United States, and its biochronological signifi­
cance. Palaeontology, 41(6), 1215–1230.
Lucas, S. G., Heckert, A. B., & Rinehart, L. F. (2006). The Late Triassic
aetosaur Paratypothorax. In Harris J. D., Lucas S. G., Spielmann J.
A., Lockley M. G., Milner A. R. C., & Kirkland J. I. (eds). The
Triassic-Jurassic Terrestrial Transition, New Mexico Museum of
Natural History and Science Bulletin. Albuquerque: New Mexico
Museum of Natural History and Science, 37, 575–580.
Lucas, S. G., Hunt, A. P., Heckert, A. B., & Spielmann, J. A. (2007).
Global Triassic tetrapod biostratigraphy and biochronology: 2007
status. New Mexico Museum of Natural History and Science
Bulletin, 41, 229–240.
Lucas, S. G., Rinehart, L. F., Krainer, K., Spielmann, J. A., & Heckert, A.
B. (2010). Taphonomy of the Lamy amphibian quarry: a Late
Triassic bonebed in New Mexico, USA. Palaeogeography,
Palaeoclimatology, Palaeoecology, 298(3–4), 388–398.
Lydekker, R. (1887). The fossil vertebrata of India. Records of the
Geological Survey of India, 20, 51–80.
Marsh, O. C. (1884). The classification and affinities of dinosaurian rep­
tiles. Nature, 31, 68–69.
Marsh, O. C. (1896). ART. VIII.–A New Belodont Reptile
(STEGOMUS) from the Connecticut River Sandstone. American
Journal of Science (1880–1910), 2(7), 59.
Marsh, A. D., Smith, M. E., Parker, W. G., Irmis, R. B., & Kligman, B. T.
(2020). Skeletal anatomy of Acaenasuchus geoffreyi Long and
Murry, 1995 (Archosauria: Pseudosuchia) and its implications for
the origin of the aetosaurian carapace. Journal of Vertebrate
Paleontology, 40(4), e1794885.
Martinez, R. N., Sereno, P. C., Alcober, O. A., Colombi, C. E., Renne, P.
R., Montañez, I. P., & Currie, B. S. (2011). A basal dinosaur from the
dawn of the dinosaur era in southwestern Pangaea. Science, 331
(6014), 206–210.
Martz, J. W. (2002). The morphology and ontogeny of Typothorax cocci­
narum (Archosauria, Stagonolepididae) from the Upper Triassic of
the American Southwest [Doctoral dissertation, Texas Tech
University].
Martz, J. W., & Parker W. G. (2017). Revised formulation of the Late
Triassic land vertebrate “Faunachrons” of western North
America: recommendations for codifying nascent systems of ver­
tebrate biochronology. In K. E. Zeigler K. E. & Parker W. G.
(Eds.), Terrestrial depositional systems: Deciphering complexities
through multiple stratigraphic methods, (pp. 39–124). Amsterdam:
Elsevier.
Martz, J. W., & Small, B. J. (2006). Tecovasuchus chatterjeei, a new aeto­
saur (Archosauria: Stagonolepididae) from the Tecovas Formation
(carnian, Upper Triassic) of Texas. Journal of Vertebrate
Paleontology, 26(2), 308–320.
Martz, J. W., Mueller, B., Nesbitt, S. J., Stocker, M. R., Parker, W. G.,
Atanassov, M., Fraser, N., Weinbaum, J., & Lehane, J. R. (2013).
A taxonomic and biostratigraphic re-evaluation of the Post
Quarry vertebrate assemblage from the Cooper Canyon
Formation (Dockum Group, Upper Triassic) of southern Garza
County, western Texas. Earth and Environmental Science
Transactions of the Royal Society of Edinburgh, 103(3–4), 339–364.
Nath, T. T., & Yadagiri, P. (2007). A new mammal-like reptile
(Cynodontia) from Upper Triassic Maleri Formation of Pranhita-
Godavari Valley, Andhra Pradesh. Geological Society of India, 69
(1), 57–60.
Nesbitt, S. J., Stocker, M. R., Ezcurra, M. D., Fraser, N. C., Heckert, A. B.,
Parker, W. G., & Marsh, A. D. (2022). Widespread azendohsaurids
(Archosauromorpha, Allokotosauria) from the Late Triassic of
western USA and India. Papers in Palaeontology, 8(1), e1413.
Novas, F. E., Ezcurra, M. D., Chatterjee, S., & Kutty, T. S. (2011). New
dinosaur species from the Upper Triassic Upper Maleri and
Lower Dharmaram formations of central India. Earth and
Environmental Science Transactions of the Royal Society of
Edinburgh, 101(3–4), 333–349.
Osborn, H. F. (1903). On the primary division of the Reptilia into two
sub-classes, Synapsida and Diapsida. Science, 17(424), 275–276.
Paes-Neto, V. D., Desojo, J. B., Brust, A. C., Ribeiro, A. M., Schultz, C. L.,
& Soares, M. B. (2021a). The first braincase of the basal aetosaur
Aetosauroides scagliai (Archosauria: Pseudosuchia) from the
Upper Triassic of Brazil. Journal of Vertebrate Paleontology, 41(2),
e1928681.
Paes-Neto, V. D., Desojo, J. B., Brust, A. C. B., Schultz, C. L., Da-Rosa,
A. A. S., & Soares, M. B. (2021b). Intraspecific variation in the
axial skeleton of Aetosauroides scagliai (Archosauria: Aetosauria)
and its implications for the aetosaur diversity of the Late Triassic
of Brazil. Anais da Academia Brasileira de Ciências, 93, suppl. 2,
1–36.
Parker, W. G. (2005). A new species of the Late Triassic aetosaur
Desmatosuchus (Archosauria: Pseudosuchia). Comptes Rendus
Palevol, 4(4), 327–340.
Parker, W. G. (2007). Reassessment of the aetosaur ‘Desmatosuchus’ cha­
maensis with a reanalysis of the phylogeny of the Aetosauria
(Archosauria: Pseudosuchia). Journal of Systematic Palaeontology,
5(1), 41–68.
Parker, W. G. (2008). Description of new material of the aetosaur
Desmatosuchus spurensis (Archosauria: Suchia) from the Chinle
Formation of Arizona and a revision of the genus Desmatosuchus.
PaleoBios, 28(1), 1–40.
Parker, W. G. (2016a). Revised phylogenetic analysis of the Aetosauria
(Archosauria: Pseudosuchia); assessing the effects of incongruent
morphological character sets. PeerJ, 4, e1583.
Parker, W. G. (2016b). Osteology of the Late Triassic aetosaur Scutarx
deltatylus (Archosauria: Pseudosuchia). PeerJ, 4, e2411.
Parker, W. G. (2018a). Anatomical notes and discussion of the first
described aetosaur Stagonolepis robertsoni (Archosauria: Suchia)
from the Upper Triassic of Europe, and the use of plesiomorphies
in aetosaur biochronology. PeerJ, 6, e5455.
Parker, W. G. (2018b). Redescription of Calyptosuchus (Stagonolepis)
wellesi (Archosauria: Pseudosuchia: Aetosauria) from the Late
Triassic of the Southwestern United States with a discussion of
genera in vertebrate paleontology. PeerJ, 6, e4291.
Parker, W. G., & Martz, J. W. (2010). Using positional homology in aeto­
saur (Archosauria: Pseudosuchia) osteoderms to evaluate the taxo­
nomic status of Lucasuchus hunti. Journal of Vertebrate
Paleontology, 30(4), 1100–1108.
Parker, W. G., & Martz, J. W. (2011). The Late Triassic (Norian)
Adamanian–Revueltian tetrapod faunal transition in the Chinle
Formation of Petrified Forest National Park, Arizona. Earth and
Environmental Science Transactions of the Royal Society of
Edinburgh, 101(3–4), 231–260.
Parker, W.G., and Martz, J.W., (2017), Building local biostratigraphic
models for the Upper Triassic of western North America:
Methods and considerations. In Zeigler, K. E. and Parker, W. G.
(eds). Terrestrial depositional systems: Deciphering complexities
through multiple stratigraphic methods. Elsevier, 1–38.
Parker, W. G., Stocker, M. R., & Irmis, R. B. (2008). A new desmatosu­
chine aetosaur (Archosauria: Suchia) from the Upper Triassic
Tecovas Formation (Dockum Group) of Texas. Journal of
Vertebrate Paleontology, 28(3), 692–701.
Parker, W. G., Nesbitt, S. J., Irmis, R. B., Martz, J. W., Marsh, A. D.,
Brown, M. A., Stocker, M. R., & Werning, S. (2022). Osteology
and relationships of Revueltosaurus callenderi (Archosauria:
Suchia) from the Upper Triassic (Norian) Chinle Formation of
Petrified Forest National Park, Arizona, United States. The
Anatomical Record, 305(10), 2353–2414.
Parrish, J. M. (1994). Cranial osteology of Longosuchus meadei and the
phylogeny and distribution of the Aetosauria. Journal of
Vertebrate Paleontology, 14(2), 196–209.
Prasad, G. V., Singh, K., Parmar, V., Goswami, A., & Sudan, C. S. (2008).
Hybodont shark teeth from the continental Upper Triassic deposits
of India. 413-432. In Arratia, G., Schultze, H.-P., & Wilson, M. V. H.
(eds). Mesozoic fishes, 4: Homology and Phylogeny. Friedrich Pfeil,
Munich. 502 pp.
 Haldar et al. —Typothoracine from India (e2253292-22)
Ramezani, J., Hoke, G. D., Fastovsky, D. E., Bowring, S. A., Therrien, F.,
Dworkin, S. I., Atchley, S. C., & Nordt, L. C. (2011). High-precision
U-Pb zircon geochronology of the Late Triassic Chinle Formation,
Petrified Forest National Park (Arizona, USA): Temporal con­
straints on the early evolution of dinosaurs. Geological Society of
America Bulletin, 123(11–12), 2142–2159.
Ramezani, J., Fastovsky, D. E., & Bowring, S. A. (2014). Revised chron­
ostratigraphy of the lower Chinle Formation strata in Arizona and
New Mexico (USA): high-precision U-Pb geochronological con­
straints on the Late Triassic evolution of dinosaurs. American
Journal of Science, 314(6), 981–1008.
Rasmussen, C., Mundil, R., Irmis, R. B., Geisler, D., Gehrels, G. E.,
Olsen, P. E., & Parker, W. G. (2021). U-Pb zircon geochronology
and depositional age models for the Upper Triassic Chinle
Formation (Petrified Forest National Park, Arizona, USA):
Implications for Late Triassic paleoecological and paleoenviron­
mental change. Geological Society of America Bulletin, 133(3–4),
539–558.
Reyes, W. A., Parker, W. G., & Marsh, A. D. (2021). Cranial anatomy and
dentition of the aetosaur Typothorax coccinarum (Archosauria:
Pseudosuchia) from the Upper Triassic (Revueltian–mid Norian)
Chinle Formation of Arizona. Journal of Vertebrate Paleontology,
40(6), e1876080.
Rixon, A. E. (1976). Fossil animal remains: their preparation and conser­
vation. The Athlone Press, University of London, London 304 pp.
Sawin, H. J. (1947). The pseudosuchian reptile Typothorax meadei.
Journal of Paleontology, 21(3), 201–238.
Scheyer, T. M., Desojo, J. B., & Cerda, I. A. (2014). Bone histology of
phytosaur, aetosaur, and other archosauriform osteoderms
(Eureptilia, Archosauromorpha). The Anatomical Record, 297(2),
240–260.
Schoch, R. R. (2007). Osteology of the small archosaur Aetosaurus from
the Upper Triassic of Germany. Neues Jahrbuch Für Geologie Und
Paläontologie-Abhandlungen, 246, 1–35.
Schoch, R. R., & Desojo, J. B. (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, 73–95.
Scotese, C. R. (2001). Atlas of Earth History. PALEOMAP Project.
Arlington, University of Texas, 52 pp.
Sengupta, D. P. (2002). Indian metoposaurid amphibians revised.
Paleontological Research, 6(1), 41–65.
Small, B. J. (1998). The occurrence of Aetosaurus in the Chinle Formation
(Late Triassic, USA) and its biostratigraphic significance. Neues
Jahrbuch für Geologie und Paläontologie-Monatshefte, 5(1998),
285–296.
Small, B. J. (2002). Cranial anatomy of Desmatosuchus haplocerus
(Reptilia: Archosauria: Stagonolepididae). Zoological Journal of
the Linnean Society of London, 136(1), 97–111.
Small, B. J., & Martz, J. W. (2013). A new aetosaur from the Upper
Triassic Chinle Formation of the Eagle Basin, Colorado, USA.
Geological Society, London, Special Publications, 379(1), 393–412.
Spielmann, J. A., & Lucas, S. G. (2012). Tetrapod fauna of the Upper
Triassic Redonda Formation, east central New Mexico: the charac­
teristic assemblage of the Apachean land-vertebrate faunachron.
New Mexico Museum of Natural History and Science Bulletin,
55,1–119.
Spielmann, J. A., Hunt, A. P., Lucas, S. G., & Heckert, A. B. (2006).
Revision of Redondasuchus (Archosauria: Aetosauria) from the
Upper Triassic Redonda Formation, New Mexico, with description
of a new species. New Mexico Museum of Natural History and
Science Bulletin, 37, 583–587.
Sulej, T. (2010). The skull of an early Late Triassic aetosaur and the evol­
ution of the stagonolepidid archosaurian reptiles. Zoological
Journal of the Linnean Society of London, 158(4), 860–881.
Sulej, T., Niedźwiedzki, G., & Bronowicz, R. (2012). A new Late
Triassic vertebrate fauna from Poland with turtles, aetosaurs, and
coelophysoid dinosaurs. Journal of Vertebrate Paleontology, 32(5),
1033–1041.
Taborda, J. R., Heckert, A. B., & Desojo, J. B. (2015). Intraspecific
variation in Aetosauroides scagliai Casamiquela (Archosauria:
Aetosauria) from the Upper Triassic of Argentina and Brazil:
an example of sexual dimorphism? Ameghiniana, 52(2), 173–
187.
Tanner, L. H. (2000). Palustrine-lacustrine and alluvial facies of the
(Norian) Owl Rock Formation (Chinle Group), Four Corners
region, southwestern U.S.A.: implications for Late Triassic paleocli­
mate. Journal of Sedimentary Research, 70(6), 1280–1289.
Tanner, L. H. (2003). Pedogenic features of the Chinle Group, Four
Corners region: evidence of Late Triassic aridification. In Lucas, S.
G., Semken, S. C., Berglof, W., & Ulmer-Scholle, D. S. (eds).
Geology of the Zuni Plateau: New Mexico Geological Society
Guidebook, 54, 269–280.
Teschner, E. M., Konietzko-Meier, D., & Klein, N. (2022). Growth and
limb bone histology of aetosaurs and phytosaurs from the Late
Triassic Krasiejów locality (sw Poland) reveals strong environ­
mental influence on growth pattern. Contributions to Zoology, 91
(3), 199–232.
Tewari, R. C., & Maejima, W. (2010). Origin of Gondwana basins of
peninsular India. Journal of Geosciences, 53, 43–49.
von Baczko, M. B., Taborda, J. R., & Desojo, J. B. (2018).
Paleoneuroanatomy of the aetosaur Neoaetosauroides engaeus
(Archosauria: Pseudosuchia) and its paleobiological implications
among archosauriforms. PeerJ, 6, e5456.
von Huene, F. (1915). On reptiles of the New Mexican Trias in the Cope
collection. Bulletin of the American Museum of Natural History, 34,
485–507.
von Huene, F. (1920). Osteologie von Aëtosaurus ferratus O. Fraas. Acta
Zoologica, 1(3), 465–491.
von Huene, F. (1940). The tetrapod fauna of the upper Triassic Maleri
bets. Palaeontologica Indica, new series, 32(1), 1–42.
Walker, A. D. (1961). Triassic reptiles from the Elgin area: Stagonolepis,
Dasygnathus and their allies. Philosophical Transactions of the
Royal Society of London. Series B, Biological Sciences, 244(709),
103–204.
Whiteside, J. H., Grogan, D. S., Olsen, P. E., & Kent, D. V. (2011).
Climatically driven biogeographic provinces of Late Triassic tropi­
cal Pangea. Proceedings of the National Academy of Sciences, 108
(22), 8972–8977.
Zeigler, K. E., Heckert, A. B., & Lucas, S. G. (2002). A new species of
Desmatosuchus (Archosauria: Aetosauria) from the Upper
Triassic of the Chama Basin, north-central New Mexico. New
Mexico Museum of Natural History and Science Bulletin, 21, 215–
219.
Zittel, K. A. von. 1887 (1887–1890). Handbuch der
Paläontologie. I. Abtheilung. Paläozoologie. 3: Vertebrata (Pisces,
Amphibia, Reptilia, Aves). R. Oldenbourg, München & Leipzig,
900 pp.
Handling Editor: Adam Huttenlocker.
Phylogenetics Editor: Pedro L. Godoy.