Received: 10 April 2023 Revised: 4 August 2023 Accepted: 22 August 2023

DOI: 10.1002/ar.25317

COMMENTARY

New information on the mandibular anatomy of

Agudotherium gassenae, a Late Triassic non-mammaliaform
probainognathian from Brazil

Leonardo Kerber 1,2 |
vio A. Pretto 1,2

Fla | Rodrigo T. Müller 1,2
1
Centro de Apoio à Pesquisa Paleontol

ogica da Quarta Colônia, Universidade Federal de Santa Maria, São João do Polêsine, Brazil
2
Programa de P

os-Graduação em Biodiversidade Animal, Universidade Federal de Santa Maria, Santa Maria, Brazil

Correspondence
Leonardo Kerber, Centro de Apoio à
Pesquisa Paleontol
ogica da Quarta
Colônia, Universidade Federal de Santa
Maria, São João do Polêsine, RS, Brazil.
Email: leonardokerber@gmail.com
Funding information
Conselho Nacional de Desenvolvimento
Científico e Tecnol

ogico, Grant/Award
Numbers: 303034/2022-0, 309178/2022-3,
309414/2019-9, 404095/2021-6,
406902/2022-4, 422568/2018-0; Fundação
de Amparo à Pesquisa do Estado do Rio
Grande do Sul, Grant/Award Numbers:
21/2551-0000619-6, 21/2551-0002030-0;
Proyectos de Investigaci

on Científica y
Tecnol
ogica (PICT), Grant/Award
Number: 2020-SERIEA-01498
Abstract
Agudotherium gassenae is a poorly known non-mammaliaform probainog-
nathian cynodont from the Late Triassic of southern Brazil. It is known only by
mandibular remains, and its affinities within Probainognathia are unclear.
Furthermore, its phylogenetic affinities were never investigated through com-
putational analyses. In this study, we described new lower jaw remains exca-
vated from the type locality and performed the first phylogenetic investigation
of this taxon. The new specimen provides further anatomical information. The
rostral region of the lower jaw was poorly preserved in the type series, leading
to the interpretation that A. gassenae had three lower incisors. The new speci-
men demonstrates the presence of four incisors. The phylogenetic analysis posi-
tioned A. gassenae as the sister group of Prozostrodontia. This hypothesis differs
from that previously presented in the former description of the taxon, in which
it was considered a non-mammaliaform prozostrodont by means of character-
state comparisons.

KEYWORDS
Carnian, Niemeyer site, Norian, Paleobiodiversity, Prozostrodontia

1 | INTRODUCTION (Abdala et al., 2020; Martinelli & Soares, 2016; Schultz

Eucynodontia is one of the most diverse and abundant
tetrapod clades in the Brazilian Triassic strata (Schultz
et al., 2020). Non-mammaliaform probainognathians are
eucynodonts with a wide range of body shapes and life-
styles, including small to large-sized carnivorous, omnivo-
rous, or herbivorous forms (Abdala, 2021; Hopson &
Kitching, 2001; Liu & Olsen, 2010; Martinelli &
Soares, 2016). The southern Brazilian Santa Maria Superse-
quence within the Paran
a Basin holds a significant portion
of the Triassic probainognathian diversity known to date
et al., 2020). Agudotherium gassenae Stefanello et al. (2020)
is a probainognathian recently described based on lower
jaws collected in the Niemeyer site (Agudo, State of Rio
Grande do Sul, Brazil), which was originally considered a
prozostrodont with uncertain phylogenetic relationships
(Stefanello et al., 2020).
Here, we describe the mandibular anatomy of a new
specimen assigned to this small mammal forerunner.
Furthermore, a phylogenetic analysis was performed to
analyze the relationships of A. gassenae within the Pro-
bainognathia clade.

Anat Rec. 2023;1–9. wileyonlinelibrary.com/journal/ar © 2023 American Association for Anatomy. 1

2 COMMENTARY

2 | MATERIALS AND METHODS Universidade Cato
lica do Rio Grande do Sul (Porto
Alegre, Brazil), using 80 kV and 88 μA. The scanning
2.1 | Provenance resulted in 2995 tomographic slices with a voxel size of
17.014 μm. The software AvizoTM (3D) was used to seg-
The specimen CAPPA/UFSM 0377 was collected at the ment the 3D models of the specimen.
Niemeyer site in Agudo, Rio Grande do Sul, Brazil (see
Pavanatto et al., 2018; Figure 1). The horizon of this
locality is part of the Candel
aria Sequence, Upper Trias-
sic (Santa Maria Supersequence; Horn et al., 2014;
Zerfass et al., 2003). Whereas this locality has not been
radioisotopically dated, current biostratigraphic
approaches suggested an age from the late Carnian to
early Norian (Martinelli et al., 2021; Miron et al., 2020;
Müller, 2021; Novas et al., 2021; Pavanatto et al., 2018;
Schultz et al., 2020).

2.2 | CT scanning and 3D modeling

The specimen is preserved within a sedimentary matrix

that can hardly be removed without damaging its frag-
ile dentition (Figure 2). For this reason, we employed
digital preparation to access its morphological informa-
tion. The specimen CAPPA/UFSM 0377 was scanned F I G U R E 2 Left lower jaws of Agudotherium gassenae in lateral
employing a μCT scan Skyscan™ 1173 at the Labor- view: CAPPA/UFSM 0377 (a) and the holotype CAPPA/UFSM
at

orio de Sedimentologia e Petrologia of the Pontifícia 0262 (b).

F I G U R E 1 Location of the
Niemeyer site (Agudo, Brazil). Surface
distribution of the geologic units in the
area (a) and aerial photograph of the
fossiliferous site (b).
COMMENTARY
2.3 | Phylogenetic analysis
Agudotherium gassenae was scored (Supporting informa-
tion S1) in the data matrix presented in Kerber et al.
(2023) based on all three available specimens (CAPPA/
UFSM 0262, 0208, 0377). This dataset is an updated ver-
sion of the phylogenetic matrix of Liu and Olsen (2010),
which was modified by a series of authors (e.g., Kerber
et al., 2022, 2023; Martinelli, Eltink, et al., 2017; Marti-
nelli, Soares, et al., 2017; Soares et al., 2014; Stefanello
et al., 2023; Wallace et al., 2019). Here, we corrected some
miscodings introduced by Wallace et al. (2019). Hence, we
returned the codifications from previous versions
(i.e., Martinelli, Eltink, et al., 2017; Martinelli, Soares,
et al., 2017). The states of the characters 107 and 108 (former
106 and 107) change from 1 to 2 in Morganucodon spp. and
Sinocodonon rigney. Tritylodontids had the state of charac-
ter 107 (former 106) changed from 2 to 3. Paleoneurological
characters were coded for Lumkuia fuzzi based on
Benoit (2023).
The final data matrix includes 158 characters and
40 operational taxonomic units. It was processed in the
software TNT v.1.5 (Goloboff & Catalano, 2016). Procyno-
suchus delaharpeae was used to root the most parsimoni-
ous trees (MPTs), which were recovered with a heuristic
search (random addition sequence + tree bi-
section reconnection), with 1000 replicates of Wagner
trees (with random seed = 0) and using tree bi-
section reconnection and branch swapping (holding
10 trees save per replication). All characters received the
same weight and were treated as unordered (i.e., non-
additive). An additional analysis was performed enfor-
cing A. gassenae within Prozostrodontia to assess the
number of extra steps necessary to support the former
hypothesis of Stefanello et al. (2020). This analysis fol-
lowed the same procedures as the first analysis.
2.4 | Institutional abbreviations
CAPPA/UFSM, Centro de Apoio à Pesquisa Paleon-
tol

ogica da Quarta Colônia da Universidade Federal de
Santa Maria, São João do Polêsine, Rio Grande do Sul,
Brazil.
3 | R E SUL T S
3.1 | Systematic paleontology
Cynodontia Owen, 1861.
Eucynodontia Kemp, 1982.
Probainognathia Hopson, 1990.
3
Agudotherium Stefanello et al., 2020.
Agudotherium gassenae Stefanello et al., 2020.
Holotype: CAPPA/UFSM 0262, a left lower jaw with
teeth.
Paratype: CAPPA/UFSM 0208, a right lower jaw
with teeth.
Referred specimen: CAPPA/UFSM 0377, mandible
including right and left dentaries with incisors, canines,
and postcanine teeth.
Locality and horizon: Niemeyer Site (29
400 3000 S;

0 00
53 13 59 W), municipality of Agudo, State of Rio Grande
do Sul, Brazil. Candel
aria Sequence, Santa Maria Super-
sequence (see Schultz et al., 2020; Langer et al., 2018).
The age of these strata has been interpreted as late Car-
nian to early Norian (see above).
3.1.1 | Description and comparison
General comments: CAPPA/UFSM 0377 is similar to
both specimens of the type series (Stefanello et al., 2020).
For this reason, here, we only highlight the most impor-
tant features of the specimen.
Dentary: The specimen is represented by a lower
jaw, including both dentary bones (Figures 2–4). The
symphysis of the dentary is not fused, similar to
CAPPA/UFSM 0262, CAPPA/UFSM 0208, Bonacynodon
schultzi, and prozostrodonts (Liu & Olsen, 2010;
Martinelli et al., 2016; Stefanello et al., 2020, 2023), and
distinct from non-prozostrodont probainognathians,
such as L. fuzzi, Chiniquodon theothonicus, Aleodon
spp., Protheriodon estudianti, Candelariodon barberenai,
Trucidocynodon riograndensis, and Probainognathus jen-
seni (Abdala & Giannini, 2002; Bonaparte et al., 2006;
Hopson & Kitching, 2001; Martinelli et al., 2016; Marti-
nelli, Soares, et al., 2017; Oliveira et al., 2010;
Romer, 1970). The dentary body of CAPPA/UFSM 0377
is more robust than in CAPPA/UFSM 0262 and CAPPA/
UFSM 0208. It increases in height caudally (less than in
the type series; see below; Figure 3a,d), resembling the
mandibular anatomy of L. fuzzi, Riograndia guaibensis,
and Irajatherium hernandezi (Abdala & Ribeiro, 2010;
fig. 5; Kerber et al., 2022), and differing from the pattern
found in Prozostrodon brasiliensis and Brasilodon quad-
rangularis (Martinelli, Soares, et al., 2017; Pacheco
et al., 2018), in which the height of the body of the den-
tary is similar along its extension. In C. barberenai, the
dentary is deeper than in A. gassenae and the height of
the dentary body is uniform along the main axis (see
Martinelli, Soares, et al., 2017). The height of the den-
tary body of P. jenseni is proportionally similar to A. gas-
senae, but it does not increase in height caudally (see
4 COMMENTARY
Martinelli & Soares, 2016; fig. 5). The coronoid process
is broken, and its base is located at the level of the pc10
(Figure 3a).
The rostralmost region of the dentary of CAPPA/
UFSM 0377 is better preserved than in CAPPA/UFSM
0262 and CAPPA/UFSM 0208 (Figure 3a–d). The dorsal-
most area of this region surpasses the alveolar level of the
postcanines (Figure 3a). On the rostral face of this region,
the dentary is perforated by two foramina (Figure 3a,c).
On the lateral aspect of the body of the dentary, the men-
tal foramen is located caudally to the canine (Figure 3a),
as in the holotype and paratype.
On the lateral face of the dentary, a shallow masse-
teric fossa is located on the caudal region of the den-
tary, ventrally to the coronoid process (Figure 3a,d).
The fossa is not as marked as observed in P. jenseni, R.
F I G U R E 3 Lower jaw and dentition
of CAPPA/UFSM 0377. (a–c) Left
dentary (with a translucent dentary on
the right showing the mandibular canal
in red) in lateral (a), medial (b), and
rostrolateral (c) views. (d) Right dentary
and teeth, in lateral and medial views.
(e) Left pc9 in labial and lingual views.
(f) Left pc10 in labial and lingual views.
a–e, cusps a–e; ca, canine; i 1–4, first to
fourth lower incisors; f, foramina; msf,
masseteric fossa; Mg, Meckelian groove;
mf, mental foramen; pc1-10, first to
tenth postcanine; sy, symphysis.
guaibensis, and B. quadrangularis (see Abdala & Ribeiro,
2010; fig. 5; Martinelli & Soares, 2016, figure 5). On the
medial aspect, CAPPA/UFSM 0377 displays a Meckelian
groove at the mid-height on the dentary (Figure 3b), simi-
lar to CAPPA/UFSM 0262. The mandibular canal (passage
of the mandibular nerve) is rostrocaudally oriented
(Figures 3a,b, and 4a,b). A small branch separates from
the main canal near the base of the canine tooth and exits
the dentary through the mental foramen (Figures 3a and
4c). In the rostralmost region of the dentary, the canal is
branched in small canaliculi. On the anterior face of the
dentary, three ramifications were observed (Figure 3c).
Two of them exit the dentary via two foramina, and the
other extends to the base of the incisors (Figure 3c).
Teeth: The most remarkable trait observed in
CAPPA/UFSM 0377, which is not observed in CAPPA/
COMMENTARY
F I G U R E 4 Digital sections of the lower jaw of Agudotherium
gassenae. (a,b) Sagittal and parasagittal sections of the left dentary
of CAPPA/UFSM 0377 and transverse plane of the right dentary
(above). (c) Transverse sections of the left dentary. (d,e) Sagittal
and parasagittal sections of the right dentary of CAPPA/UFSM
0262. 1–10, 1 to 10th postcanines; c, lower canine; cr, canine root;
dl, dental lamina; ea, empty alveolus; i1-4, first to fourth incisors;
mc, mandibular canal; mf, mental foramen; rpc, replacement
postcanine.
UFSM 0262 and CAPPA/UFSM 0208, is the incisor
morphology (Figure 3a–c). They are conical and well-
developed, and their apex is generally higher than those
of postcanines (Figure 3a–c). The first three incisors have
5
approximately the same size, while the fourth one is
smaller. In the original description, Stefanello et al.
(2020) described the presence of three incisors. However,
in CAPPA/UFSM 0377, in which the rostral region of the
lower jaw is better preserved, there are four incisors
(Figures 3a–c and 4b). Four lower incisors are present in P.
estudianti and P. brasiliensis (Bonaparte & Barberena, 2001;
Martinelli, Soares, et al., 2017; Pacheco et al., 2018;
Stefanello et al., 2023), but not in Alemoatherium huebneri,
which has five incisors, possibly representing a juvenile
condition (Martinelli, Eltink, et al., 2017), while other pro-
bainognathians have three, such as Aleodon spp., T. rio-
grandensis, R. guaibensis, I. hernandezi, and adult
individuals of B. quadrangularis (Bonaparte et al., 2005;
Bonaparte & Barberena, 2001; Kerber et al., 2022;
Martinelli et al., 2016; Oliveira et al., 2010, 2011; Soares
et al., 2011). More incisors are present in juvenile forms of
the latter taxon (Martinelli, 2017). In CAPPA/UFSM 0377,
there is no diastema between the fourth incisor and the
canine (Figure 3a).
The canine is well developed, linguo-labially com-
pressed (Figure 4c), without mesial or distal serrations,
which are present in ecteniniids, B. schultzi, and variably
in P. brasiliensis (Martinelli et al., 2016; Oliveira
et al., 2010; Stefanello et al., 2023). The canine of CAPPA/
UFSM 0377 is more robust than those of the type series.
The tips of both canines are missing (Figure 3a–d).
CAPPA/UFSM 0377 has 10 postcanines (Figures 3a–d
and 4a,b), unlike the holotype and paratype, which have
only seven (Stefanello et al., 2020). On the left side, the
pc8-10 are better preserved (Figure 3a,b,e,f). The postcanine
roots are constricted by a longitudinal groove (Figures 3e
and 4a–c). The pc8 has a small cusp a, followed by a dor-
sally projected cusps a–c. The pc9 is better preserved than
the pc8 (Figure 3e). It shows a small cusp b, followed suc-
cessively by cusps a, c, and d. The cusp a is sharp and is the
largest one. It is slightly curved distally (Figure 3e) but not
as the pattern shared by chiniquodontids and ecteniniids, in
which the curvature is more accentuated (Abdala &
Giannini, 2002; Martinelli, Kammerer, et al., 2017; Martinez
et al., 1996; Oliveira et al., 2010). Both teeth are similar to
the distal postcanines of the holotype and paratype. Mesio-
lingually, there is a weak cingulum (cusp e), representing a
pattern less complex than in C. barberenai, P. brasiliensis,
Botucaraitherium belarminoi, and B. quadrangularis (Soares
et al., 2014; Martinelli, Soares, et al., 2017; Pacheco
et al., 2018). The pc10 is an erupting tooth (Figure 3f). Its
cusps are sharper than in the other postcanines. The cusp a
is smaller than in the pc8-9, and the cusps a–c are more
curved distally than in the other postcanines (Figure 3a).
The cusp a is slightly lingually oriented. On the right side,
pc1-8 are present, but most have broken crowns and do not
provide new anatomical data (Figure 3d).
6 COMMENTARY
3.2 | Phylogenetic analysis
The phylogenetic analysis recovered two MPTs with
499 steps (CI = 0.454, RI = 0.774; Figure 5). Agudother-
ium gassenae emerges as the sister group of Prozostro-
dontia. The clade including A. gassenae + prozostrodonts
is supported by two character-state transformations in
the strict consensus tree: unfused dentary symphysis
(83:1 ! 0) and lower postcanine teeth with roots
constricted by a longitudinal groove (108:0 ! 1). The
Ladinian/early Carnian probainognathian P. estudianti is
the sister-taxon of A. gassenae + prozostrodonts.
The constrained analysis recovered two MPTs
with 501 steps (CI = 0.453, RI = 0.773). In the strict
consensus tree, there is a polytomy at the base of
Prozostrodontia, which is composed A. gassenae, Therioh-
erpeton cargnini, Prozostrodontidae (P. brasiliensis
+ Pseudotherium argentinus), and a clade with late diverging
prozostrodonts.
4 | DISCUSSION AND
C O N C L U S IO N
4.1 | Ontogenetic remarks
CAPPA/UFSM 0377 is larger than the holotype and para-
type, and we assumed that it is ontogenetically older than
the individual(s) of the type series. In this case, some
ontogenetic observations are pertinent. The dentary of
CAPPA/UFSM 0377 is more robust, and the difference in
height between the rostral region of the dentary ramus
and the caudal region is less evident than in CAPPA/
F I G U R E 5 Abbreviated strict
consensus tree depicting the position of
Agudotherium gassenae (bold). The
numbers on nodes indicate the
Bootstrap values and Bremer support of
the main clades.
UFSM 0262 and CAPPA/UFSM 0208. Hence, the dentary
body becomes deeper as the individual grows.
The canine becomes more robust, and postcanines
are added during ontogeny (seven in the type series and
10 in the new specimen). CAPPA/UFSM 0262, the holo-
type, has two replacement postcanines replacing the pc2
and pc5, respectively (Stefanello et al., 2020). On the
other hand, CAPPA/UFSM 0377 does not show any
replacement teeth, which may indicate that the replace-
ment ceased (or was ceasing) at this ontogenetic stage.
Considering that A. gassenae had delayed ankylosis as
the pattern of tooth attachment (LeBlanc et al., 2018), in
which the tooth goes through an early stage of gomphosis
and ends with ankylosis, the pc6 and pc9 seem to be
among the latest postcanines that erupted. In such teeth,
there is an alveolar space formerly occupied by periodon-
tal ligament in life (gomphosis; LeBlanc et al., 2018).
4.2 | Morphological and phylogenetic
implications
CAPPA/UFSM 0377 is referred to A. gassenae according
to the shared diagnostic suite of traits proposed for the
taxon (Stefanello et al., 2020). Whereas this assignation
expands the fossil record of A. gassenae, it is a still poorly
known cynodont. Nevertheless, the new specimen pro-
vides new anatomical information. Its dental formula
presents 4 incisors, 1 canine, and 7–10 lower postcanines.
In the type series, the rostral region of the dentary was
not well preserved, and A. gassenae was interpreted as
having three lower incisors (Stefanello et al., 2020). In the
description of this taxon, Stefanello et al. (2020) suggested a
COMMENTARY
possible relationship with sectorial-toothed non-mamma-
liaform prozostrodonts, such as B. quadrangularis, B. belar-
minoi, or I. hernandezi, mainly due to the presence of three
incisors (although three incisors are present in non-prozos-
trodont probainognathians, they are shared by all non-
mammaliaform Norian prozostrodonts with preserved
incisors) and unfused dentary symphysis. Although it
presents some traits more commonly distributed among
prozostrodonts (e.g., non-fused symphysis, postcanine roots
with a constriction), it also has lower postcanines with less
complex morphology, with weak mesiolingual cingulum
(not complete), not evidently developed as in P. brasiliensis
and B. belarminoi (Pacheco et al., 2018; Soares et al., 2014).
Based on the available morphological information on this
taxon, our phylogenetic analysis recovered it as the sister
group of Prozostrodontia (a node-based clade; Liu &
Olsen, 2010). Indeed, the constrained analysis demands
two extra steps to recover A. gassenae within
Prozostrodontia.
We consider the results of this phylogenetic analysis to
be preliminary, considering that little is known about the
anatomy of A. gassenae. Even so, its phylogenetic position
contributes to the discussion of an unappreciated diversity
of probainognathians that flourished during the Late Trias-
sic in the branch leading to prozostrodonts (Bonaparte
et al., 2006; Martinelli et al., 2016; Martinelli, Soares,
et al., 2017). However, the main forces that drove this evolu-
tionary context are poorly investigated. The Carnian Pluvial
episode is usually regarded as one of the factors that
impacted the diversification of tetrapods and its subsequent
success (Bernardi et al., 2018; Griffin et al., 2022). Con-
versely, such humid period seems to have impacted far
more groups than previously thought, including the probai-
nognathian cynodonts, which have achieved a wide range
of early diverging and distinct phylogenetic groups (see
Stefanello et al., 2023). Agudotherium gassenae favors this
scenario, carrying a unique set of traits among this plethora
of forms representing the diversity of cynodonts before the
origin and diversification of mammaliaforms and mammals.
A U T H O R C ON T R I B U T I O NS
Leonardo Kerber: Conceptualization; data curation;
formal analysis; funding acquisition; investigation; method-
ology; project administration; resources; software; supervi-
sion; validation; visualization; writing—original draft;
writing – review and editing. Fla
vio A. Pretto:
Conceptualization; data curation; investigation; validation;
visualization; writing—original draft; writing—review and
editing. Rodrigo T. Müller: Conceptualization; data cura-
tion; formal analysis; funding acquisition; investigation;
methodology; validation; visualization; writing—original
draft; writing—review and editing.
7
ACKNOWLEDGMENTS
Agustin G. Martinelli and Jun Liu, for their valuable
comments that significantly improved this article's first
version. We also extend our gratitude to the Willi Hennig
Society for the gratuity of TNT software.
F U N D I N G IN F O R M A T I O N
Conselho Nacional de Desenvolvimento Científico e Tec-
nol

ogico (Leonardo Kerber [422568/2018-0, 309414/2019-9,
309178/2022-3, and 406902/2022-4], Rodrigo T. Müller
[CNPq 404095/2021-6, 303034/2022-0, and 406902/2022-4]),
Fundação de Amparo à Pesquisa do RS (Leonardo
Kerber [21/2551-0002030-0] and Fl
avio A. Pretto
(21/2551-0000619-6), and PICT (2020-SERIEA-01498;
Leonardo Kerber).
C O N F L I C T O F I N T E R E S T S T A TE M E N T
The authors declare no competing financial interests.
DA TA AVAI LA BI LI TY S T ATE ME NT
Data (CT-Scan) are available on request from the
authors.
ORCID
Leonardo Kerber https://orcid.org/0000-0001-8139-1493
avio A. Pretto https://orcid.org/0000-0001-8091-7932
Fl

Rodrigo T. Müller https://orcid.org/0000-0001-8894-
9875
RE FER EN CES
Abdala, F. (2021). Permo-Jurassic cynodonts: The early road to
mammalness. In Encyclopedia of geology (pp. 206–226). Else-
vier. https://doi.org/10.1016/b978-0-12-409548-9.12020-2
Abdala, F., Gaetano, L. C., Martinelli, A. G., Soares, M. B.,
Hancox, P. J., & Rubidge, B. S. (2020). Non-mammaliaform
cynodonts from western Gondwana and the significance of
Argentinean forms in enhancing understanding of the group.
Journal of South American Earth Sciences, 104, 102884. https://
doi.org/10.1016/j.jsames.2020.102884
Abdala, F., & Giannini, N. P. (2002). Chiniquodontid cyno-
donts: Systematic and morphometric considerations.
Palaeontology, 45(6), 1151–1170. https://doi.org/10.1111/
1475-4983.00280
Abdala, F., & Ribeiro, A. M. (2010). Distribution and diversity pat-
terns of Triassic cynodonts (Therapsida, Cynodontia) in Gond-
wana. Palaeogeography, Palaeoclimatology, Palaeoecology, 286
(3–4), 202–217. https://doi.org/10.1016/J.PALAEO.2010.01.011
Benoit, J. (2023). Synchrotron scanning reveals the deep evo-
lutionary root of the mammalian brain: The surprisingly
advanced endocast morphology of Lumkuia fuzzi
(Cynodontia: Probainognathia). Palaeontologica Africana,
56, 103–110.
Bernardi, M., Gianolla, P., Petti, F. M., Mietto, P., & Benton, M. J.
(2018). Dinosaur diversification linked with the Carnian pluvial
8 COMMENTARY
episode. Nature Communications, 9(1), 1499. https://doi.org/10.
1038/s41467-018-03996-1
Bonaparte, J. F., & Barberena, M. C. (2001). On two carnivorous
cynodonts from southern Brazil. Bulletin of the Museum of
Comparative Zoology at Harvard College, 156, 59–80.
Bonaparte, J. F., Martinelli, A. G., & Schultz, C. L. (2005). New
information of Brasilodon and Brasilitherium (Cynodontia, Pro-
bainognathia) from the Late Triassic of southern Brazil. Revista
Brasileira de Paleontologia, 8(1), 25–46.
Bonaparte, J. F., Soares, M. B., & Schultz, C. L. (2006). A new non-
mammalian cynodont from the Middle Triassic of southern
Brazil and its implications for the ancestry of mammals. In
H. D. Harris, S. G. Lucas, J. A. Spielmann, M. G. Lockley,
A. R. C. Milner, & J. L. Kirkland (Eds.), The Triassic-Jurassic
terrestrial transition (Vol. 37, pp. 599–607). New Mexico
Museum of Natural History & Science.
Goloboff, P. A., & Catalano, S. A. (2016). TNT version 1.5, including
a full implementation of phylogenetic morphometrics. Cladis-
tics, 32(3), 221–238. https://doi.org/10.1111/cla.12160
Griffin, C. T., Wynd, B. M., Munyikwa, D., Broderick, T. J.,
Zondo, M., Tolan, S., Langer, M. C., Nesbitt, S. J., &
Taruvinga, H. R. (2022). Africa's oldest dinosaurs reveal early
suppression of dinosaur distribution. Nature, 609(7926), 313–
319. https://doi.org/10.1038/s41586-022-05133-x
Hopson, J. A. (1990). Cladistic analysis of therapsid relationships.
Journal of Vertebrate Paleontology, 10(3), 28A.
Hopson, J. A., & Kitching, J. W. (2001). A probainognathian cyno-
dont from South Africa and the phylogeny of nonmammalian
cynodonts. Bulletin of the Museum of Comparative Zoology at
Harvard College, 156(1), 5–35.
Horn, B. L. D., Melo, T. M., Schultz, C. L., Philipp, R. P.,
Kloss, H. P., & Goldberg, K. (2014). A new third-order sequence
stratigraphic framework applied to the Triassic of the Paran
a
Basin, Rio Grande do Sul, Brazil, based on structural, stratigraphic
and paleontological data. Journal of South American Earth Sci-
ences, 55, 123–132. https://doi.org/10.1016/J.JSAMES.2014.07.007
Kemp, T. S. (1982). Mammal-like reptiles and the origin of mam-
mals. Academic Press.
Kerber, L., Martinelli, A. G., Müller, R. T., & Pretto, F. A. (2022). A
new specimen provides insights into the anatomy of Irajather-
ium hernandezi, a poorly known probainognathian cynodont
from the Late Triassic of southern Brazil. Anatomical Record,
305(11), 3113–3132. https://doi.org/10.1002/ar.24830
Kerber, L., Roese-Miron, L., Bubadué, J. M., & Martinelli, A. G.
(2023). Endocranial anatomy of the early prozostrodonts
(Eucynodontia: Probainognathia) and the neurosensory evolu-
tion in mammal forerunners. The Anatomical Record, 1–32.
https://doi.org/10.1002/ar.25215
Langer, M. C., Ramezani, J., & Da Rosa, Á. A. S. (2018). U-Pb age
constraints on dinosaur rise from south Brazil. Gondwana
Research, 57, 133–140. https://doi.org/10.1016/J.GR.2018.01.005
LeBlanc, A. R. H., Brink, K. S., Whitney, M. R., Abdala, F., &
Reisz, R. R. (2018). Dental ontogeny in extinct synapsids reveals a
complex evolutionary history of the mammalian tooth attachment
system. Proceedings of the Royal Society B: Biological Sciences,
285(1890), 20181792. https://doi.org/10.1098/rspb.2018.1792
Liu, J., & Olsen, P. (2010). The phylogenetic relationships of Eucy-
nodontia (Amniota: Synapsida). Journal of Mammalian Evolu-
tion, 17(3), 151–176. https://doi.org/10.1007/s10914-010-9136-8
Martinelli, A. G. (2017). Contribuição ao conhecimento dos cino-
dontes probainogn
atios (Therapsida, Cynodontia, Probainog-
nathia) do Tri
assico da América do Sul e seu impacto na origem
dos Mammaliaformes. Universidade Federal do Rio Grande
do Sul.
Martinelli, A. G., Eltink, E., Da-Rosa, Á. A. S., & Langer, M. C.
(2017). A new cynodont from the Santa Maria formation, South
Brazil, improves Late Triassic probainognathian diversity.
Papers in Palaeontology, 3(3), 401–423. https://doi.org/10.1002/
spp2.1081
Martinelli, A. G., Escobar, J. A., Francischini, H., Kerber, L.,
Müller, R. T., Rubert, R., Schultz, C. L., & Da-Rosa, Á. A. S.
(2021). New record of a stahleckeriid dicynodont (Therapsida,
Dicynodontia) from the Late Triassic of southern Brazil and
biostratigraphic remarks on the Riograndia assemblage zone.
Historical Biology, 33(11), 3101–3110. https://doi.org/10.1080/
08912963.2020.1850715
Martinelli, A. G., Kammerer, C. F., Melo, T. P., Paes Neto, V.,
Ribeiro, A. M., Da-Rosa, A. A. S., Schultz, C. L., & Soares, M. B.
(2017). The African cynodont Aleodon (Cynodontia, Probainog-
nathia) in the Triassic of southern Brazil and its biostrati-
graphic significance. PLoS One, 12(6), 1–54. https://doi.org/10.
1371/journal.pone.0177948
Martinelli, A. G., & Soares, M. B. (2016). Evolution of south Ameri-
can cynodonts. In F. L. Agnolin, G. L. Lio, F. Briss
on Egli, N. R.
Chimento, & F. Novas (Eds.), Contribuciones del • Historia evo-
lutiva y paleobiogeogr
afica de los vertebrados de América del Sur.
Museo Argentino de Ciencias Naturales ‘Bernardino Rivadavia’
(Vol. 6, pp. 183–197). Asociaci
on Paleontol
ogica Argentina.
Martinelli, A. G., Soares, M. B., & Cibele, S. (2016). Two new cyno-
donts (Therapsida) from the middle-early Late Triassic of Brazil
and comments on South American probainognathians. PLoS
One, 11(10), 1–43. https://doi.org/10.1371/journal.pone.0162945
Martinelli, A. G., Soares, M. B., de Oliveira, T. V., Rodrigues, P. G., &
Schultz, C. L. (2017). The Triassic eucynodont Candelariodon bar-
berenai revisited and the early diversity of stem prozostrodontians.
Acta Palaeontologica Polonica, 62, 527–542. https://doi.org/10.
4202/APP.00344.2017
Martinez, R. N., May, C. L., & Forster, C. A. (1996). A new carnivo-
rous cynodont from the Ischigualasto formation (Late Triassic,
Argentina), with comments on eucynodont phylogeny. Journal
of Vertebrate Paleontology, 16(2), 271–284. https://doi.org/10.
1080/02724634.1996.10011314
Miron, L. R., Pavanatto, A. E. B., Pretto, F. A., Müller, R. T., Dias-
da-Silva, S., & Kerber, L. (2020). Siriusgnathus niemeyerorum
(Eucynodontia: Gomphodontia): The youngest south American
traversodontid? Journal of South American Earth Sciences, 97,
102394. https://doi.org/10.1016/j.jsames.2019.102394
Müller, R. T. (2021). A new theropod dinosaur from a peculiar Late
Triassic assemblage of southern Brazil. Journal of South Ameri-
can Earth Sciences, 107, 103026. https://doi.org/10.1016/J.
JSAMES.2020.103026
Novas, F. E., Agnolin, F. L., Ezcurra, M. D., Temp Müller, R.,
Martinelli, A. G., & Langer, M. C. (2021). Review of the fossil
record of early dinosaurs from South America, and its phyloge-
netic implications. Journal of South American Earth Sciences,
110, 103341. https://doi.org/10.1016/j.jsames.2021.103341
Oliveira, T., Soares, M. B., & Schultz, C. (2010). Trucidocynodon rio-
grandensis gen. Nov. et sp. nov. (Eucynodontia), a new
COMMENTARY
cynodont from the Brazilian upper Triassic (Santa Maria forma-
tion). Zootaxa, 2382, 1–71.
Oliveira, T. V., Martinelli, A. G., & Soares, M. B. (2011). New infor-
mation about Irajatherium hernandezi Martinelli, Bonaparte,
Schultz & Rubert 2005 (Eucynodontia, Tritheledontidae) from
the upper Triassic (Caturrita formation, Paran
a Basin) of
Brazil. Paläontologische Zeitschrift, 85(1), 67–82. https://doi.
org/10.1007/s12542-010-0078-5
Owen, R. (1861). Palaeontology, or a systematic summary of extinct
animals and their geological relations (2nd ed.). Adam and
Black.
Pacheco, C. P., Martinelli, A. G., Pavanatto, A. E. B.,
Soares, M. B., & Dias-da-Silva, S. (2018). Prozostrodon brasilien-
sis, a probainognathian cynodont from the Late Triassic of
Brazil: Second record and improvements on its dental anatomy.
Historical Biology, 30(4), 475–485. https://doi.org/10.1080/
08912963.2017.1292423
Pavanatto, A. E. B., Pretto, F. A., Kerber, L., Müller, R. T., Da-
Rosa, Á. A. S., & Dias-da-Silva, S. (2018). A new upper Triassic
cynodont-bearing fossiliferous site from southern Brazil, with
taphonomic remarks and description of a new traversodontid
taxon. Journal of South American Earth Sciences, 88, 179–196.
https://doi.org/10.1016/j.jsames.2018.08.016
Romer, A. (1970). The Chañares (Argentina) Triassic reptile Fauna
VI. A chiniquodontid cynodont with an incipient squamosal-
dentary jaw articulation. Breviora, 344, 1–18.
Schultz, C. L., Martinelli, A. G., Soares, M. B., Pinheiro, F. L.,
Kerber, L., Horn, B. L. D., Pretto, F. A., Müller, R. T., &
Melo, T. P. (2020). Triassic faunal successions of the Paran
a
Basin, southern Brazil. Journal of South American Earth Sci-
ences, 104, 102846. https://doi.org/10.1016/j.jsames.2020.102846
Soares, M. B., Martinelli, A. G., & De Oliveira, T. V. (2014). A new
prozostrodontian cynodont (Therapsida) from the Late Triassic
Riograndia assemblage zone (Santa Maria Supersequence) of
southern Brazil. Anais Da Academia Brasileira de Ciencias, 86(4),
1673–1691. https://doi.org/10.1590/0001-3765201420140455
Soares, M. B., Schultz, C. L., & Horn, B. L. D. (2011). New informa-
tion on Riograndia guaibensis Bonaparte, Ferigolo & Ribeiro,
2001 (Eucynodontia, Tritheledontidae) from the Late Triassic
9
of southern Brazil: anatomical and biostratigraphic implica-
tions. Anais Da Academia Brasileira de Ciências, 83(1),
329–354.
Stefanello, M., Kerber, L., Martinelli, A. G., & Dias-Da-Silva, S.
(2020). A new prozostrodontian cynodont (Eucynodontia, Pro-
bainognathia) from the upper Triassic of southern Brazil. Jour-
nal of Vertebrate Paleontology, 40(3), e1782415. https://doi.org/
10.1080/02724634.2020.1782415
Stefanello, M., Martinelli, A. G., Müller, R. T., Dias-da-Silva, S., &
Kerber, L. (2023). A complete skull of a stem mammal from the
late Triassic of Brazil illuminates the early evolution of prozos-
trodontian cynodonts. Journal of Mammalian Evolution., 30,
299–317. https://doi.org/10.1007/s10914-022-09648-y
Wallace, R. V. S., Martínez, R., & Rowe, T. (2019). First record of a
basal mammaliamorph from the early late Triassic Ischigual-
asto formation of Argentina. PLoS One, 14(8), e0218791.
https://doi.org/10.1371/journal.pone.0218791
Zerfass, H., Lavina, E. L., Schultz, C. L., Garcia, A. J. V.,
Faccini, U. F., & Chemale, F. (2003). Sequence stratigraphy of
continental Triassic strata of southernmost Brazil: A contribu-
tion to southwestern Gondwana palaeogeography and palaeo-
climate. Sedimentary Geology, 161(1–2), 85–105. https://doi.org/
10.1016/S0037-0738(02)00397-4
SU PP O R TI N G I N F O RMA TI O N
Additional supporting information can be found online
in the Supporting Information section at the end of this
article.
How to cite this article: Kerber, L., Pretto, F. A.,
& Müller, R. T. (2023). New information on the
mandibular anatomy of Agudotherium gassenae, a
Late Triassic non-mammaliaform
probainognathian from Brazil. The Anatomical
Record, 1–9. https://doi.org/10.1002/ar.25317