Cretaceous Research 137 (2022) 105250

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Cretaceous Research
journal homepage: www.elsevier.com/locate/CretRes

Theropods from the La Bonita site, Bajo de la Carpa Formation

(Neuque
n Group, Santonian), Río Negro, Argentina: analysis of dental
evidence
J.G. Meso a, b, *, F.A. Gianechini c, R.D. Jua

rez Valieri d, S. Apesteguía e, S.A.S. Correa a, b
a
Universidad Nacional de Río Negro, Instituto de Investigacio
n en Paleobiología y Geología, Río Negro, Argentina
b
IIPG, UNRN, Consejo Nacional de Investigaciones Científicas y Tecnolo
gicas (CONICET), Av. Roca 1242, (R8332EXZ) General Roca, Río Negro, Argentina
c
gicas de San Luis (IMIBIO-SL), CONICET-Universidad Nacional de San Luis, Ej

Instituto Multidisciplinario de Investigaciones Biolo ercito de Los Andes 950,
San Luis, D5700HHV, Argentina
d
Secretaria de Cultura de la Provincia de Río Negro, Museo Provincial Carlos Ameghino, Belgrano 2150 Cipolletti, Río Negro, 8324, Argentina
e
CONICET- Fundacio
n F
elix de Azara - Universidad Maimo
nides, Hidalgo 775, CABA, Argentina

a r t i c l e i n f o a b s t r a c t

Article history: The abundant record of theropods from Bajo de La Carpa Formation (Neuque
n Group, Santonian), known
Received 20 December 2021 from the end of the nineteenth century, come from numerous locations within the Neuque
n Basin.
Received in revised form During the excavation of the titanosaur Bonitasaura salgadoi at the La Bonita fossiliferous site, northwest
27 April 2022
of Río Negro province Argentina, were recovered three isolated teeth assignable to non-avian theropod
Accepted in revised form 7 May 2022
Available online 11 May 2022
dinosaurs. Previous studies of these dental materials suggested that MPCA-Pv-247 corresponds to an
indeterminate tetanure possibly related to Orkoraptor, a taxon of uncertain phylogenetic position at that
moment, and MPCA-Pv-249 and 251 as possible abelisauroids. Three methods were carried out, namely,
Keywords:
Megaraptoridae
a cladistic analysis performed on a dentition-based data matrix, and a discriminant and cluster analyses
Abelisauridae performed in a large dataset including measurements of non-avian theropod teeth. The results assign for
Isolated teeth the first time a confidently phylogenetic position to the described dental material. The analysis shows
La Bonita that MPCA-Pv 247 belongs to Megaraptoridae, whereas MPCA-Pv 249 and 251 were recovered as
Patagonia belonging to Abelisauridae, supporting in a reliable way the previous assignments. The results show the
presence of Megaraptoridae at La Bonita and, additionally, they represent an evidence of the first direct
association of megaraptorids and abelisaurids at the same locality of the Bajo de La Carpa Formation,
according to similar associations from other units of the Neuque
n Group.
© 2022 Elsevier Ltd. All rights reserved.

1. Introduction Gianechini et al., 2020). Particularly, the theropod fauna from these
deposits has a major role in our understanding of the evolution as a
The Bajo de la Carpa Formation (Santonian, Upper Cretaceous) is whole of the theropod lineage history in South America (see Novas
a continental unit belonging to the Neuque
n Group of the Neuque
n et al., 2013; Holtz, 2021). The non-avian theropod record includes at
Basin, which presents a wide distribution throughout northwestern least three clades, i.e., Abelisauroidea, Megaraptoridae, and Alvar-
Patagonia (Garrido, 2010, 2011). This geological unit has yielded a ezsauridae (Bonaparte, 1991; Martinelli and Vera, 2007; Ezcurra
large diversity of terrestrial tetrapods, mainly dominated by di- and Me
ndez, 2009; Filippi et al., 2016; Porfiri et al., 2018;
nosaurs (e.g., Woodward, 1896; Bonaparte, 1991; Caldwell and Gianechini et al., 2020). The first one is represented by the small
Albino, 2001; Apesteguía, 2004; Gianechini et al., 2011; Filippi abelisauroid Velocisaurus unicus (Bonaparte, 1991), an indetermi-
et al., 2016; Porfiri et al., 2018; Cruzado-Caballero et al., 2019; nate form (Ezcurra and Me
ndez, 2009), and two new taxa recently
published, Viavenator exxoni and Llukalkan aliocranianus (Filippi
et al., 2016; Gianechini et al., 2020). Concerning megaraptorids,
there are two records from the Neuque
n province to date, one of
* Corresponding author. Universidad Nacional de Río Negro, Instituto de Inves-

n en Paleobiología y Geología, Río Negro, Argentina.
tigacio
them is Tratayenia rosalesi, based on an associated but partial
E-mail addresses: [email protected] (J.G. Meso), [email protected] postcranial skeleton (Porfiri et al., 2018), and the other consists of

rez Valieri).
(R.D. Jua

https://doi.org/10.1016/j.cretres.2022.105250
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rez Valieri et al.
J.G. Meso, F.A. Gianechini, R.D. Jua Cretaceous Research 137 (2022) 105250

fragmentary postcranial remains (Me
ndez et al., 2021). Regarding bodied (i.e., CH > 20 mm) ziphodont theropod taxa, including Dilo-
the Alvarezsauria, they are known to date from two taxa provenant phosauridae, Ceratosauridae, Abelisauridae, Megalosauroidea, Allo-
from the Neuque
n and Río Negro provinces, Alvarezsaurus calvoi sauroidea, and Tyrannosauroidea. The terminologies of Smith and
and Achillesaurus manazzonei, respectively (Bonaparte, 1991; Dodson (2003) and Hendrickx et al. (2015b) were used for the
Martinelli and Vera, 2007). anatomical orientation of the theropod teeth, whereas the
In 2011, four isolated archosaur teeth were reported from the La morphometric and anatomical terms and abbreviations follow those
Bonita fossiliferous quarry, corresponding to the uppermost section defined and illustrated by Smith et al. (2005) and Hendrickx et al.
of the Bajo de la Carpa Formation (Santonian; Upper Cretaceous), (2015b) to describe and label each tooth. Also, we follow phyloge-
located 8 km NW from Cerro Policía village, in northwestern Río netic definitions by Hendrickx et al. (2015c, 2020a, b).
Negro Province, Patagonia, Argentina (Gianechini et al., 2011). This
contribution increased the knowledge of theropod and crocodyli- 2.2. Cladistic analysis
form content for the Bajo de La Carpa Formation faunal assemblage.
The aim of this paper is: i) to review and redescribe the published The inclusion of teeth in phylogenetic analyses was possible
theropod dental material MPCA-Pv 247, 249 and 251; ii) to identify after the dentition-based data matrix created by Hendrickx and
these shed crowns using the new phylogenetic and morphometric Mateus (2014), and revised and updated by Young et al. (2019)
techniques implemented for identifying isolated teeth; iii) to and Hendrickx et al. (2019, 2020a, b). One additional taxon,
discuss about the current knowledge on the middle Late Cretaceous namely, the megaraptorid Murusraptor barrosaensis (Coria and
communities of theropods of the Bajo de La Carpa Formation. Currie, 2016) was added to the dataset of Hendrickx et al. (2020a)
by first-hand examination. In this sense, our data matrix includes
1.1. Institutional abbreviations 146 scored characters across 106 genus-level operational taxo-
nomic units (see Supplementary information 1). In addition, each
MACN-Pv-RN, Museo Argentino de Ciencias Naturales “Bernar- tooth was treated as separated operational taxonomic units (see

n Paleontología de Vertebrados (Pv),
dino Rivadavia”, Coleccio Supplementary information 3: Table 1).

n Provincia de Río Negro (RN), Buenos Aires, Argentina;
Coleccio To evaluate and visualize which taxa approach morphologically
MPCA, Museo Provincial “Carlos Ameghino”, Cipolletti, Argentina. to the La Bonita teeth (i.e., in terms of dental character states) three
cladistic analyses were performed under the methodology of Young
1.2. Anatomical abbreviations et al. (2019) and Hendrickx et al. (2020a, b), using the software TNT
(Goloboff et al., 2008). Two of these analyses (1 and 2) were con-
AL, apical length; CA, crown angle; CBL, crown base length; CBR, ducted on the data matrix of dentition-based characters (146
crown base ratio; CBW, crown base width; CH, crown height; CHR, discrete characters). In turn, the analysis 1 was carried out using
crown height ratio; CTU, crown transverse undulation density; DA, positive constraints to recover a backbone topology (n.b., and at the
distoapical denticle density; DAVG, average distal denticle density; same time setting the three teeth as floating terminals) based on
DB, distobasal denticle density; DC, distocentral denticle density; the results of the phylogenetic analyses of Müller et al. (2018) for
DDT, dentine thickness distally; DLAT, dentine thickness labially; non-neotheropod saurischians, Ezcurra (2017) for non-averostran
DLIT, dentine thickness lingually; DMT, dentine thickness mesially; neotheropods, Rauhut and Carrano (2016) and Wang et al. (2017)
DSDI, denticle size density index; FABL, fore-aft basal length; LAF, for Ceratosauria, Carrano et al. (2012) and Rauhut et al. (2012,
number of flutes on the labial surface of a crown; LIF, number of 2016) for non-coelurosaurian tetanurans, Brusatte and Carr (2016)
flutes on the lingual surface of a crown; MA, mesioapical denticle for Tyrannosauroidea, Aranciaga Rolando et al. (2019) for Mega-
density; MAVG, average mesial denticle density; MB, mesio-basal raptora, and Cau et al. (2017) for neocoelurosaurs. Analysis 2 was
denticle density; MC, mesiocentral denticle density; MCE, mesial carried out without constraints. The third analysis was based on a
carina extent; MCL, mid-crown length; MCR, mid-crown ratio; reduced data matrix restricted to tooth-based characters (charac-
MCW, mid-crown width; MDE, mesiobasal denticles extent. ters 38e122 and 141e146, total number: 91 characters), and
without constraints. From this dataset all toothless taxa were
2. Material and methods removed, namely, adult specimens of Limusaurus, Chirostenotes,
Citipati, Garudimimus, and Struthiomimus. As described in previous
2.1. Descriptive and comparative methodology and terminology studies, the search strategy of each analysis used a combination of
the tree-search algorithms Wagner trees, TBR branch swapping,
The theropod teeth studied here, MPCA-Pv 247, MPCA-Pv 249, sectorial searches, Ratchet (perturbation phase stopped after 20
and MPCA-Pv 251 (deposited at the Museo Provincial “Carlos Ame- substitutions), and Tree Fusing (5 rounds), until 100 hits of the
ghino”, Cipolletti, Argentina; MPCA) come from a fossiliferous site same minimum tree length were reached (n.b., the TNT command
denominated “La Bonita” quarry. The specimens were examined first used is “xmult ¼ hits 100 rss fuse 5 ratchet 20”). The trees were
hand using a binocular microscope Nikon SMZ/800 with different subjected to a final round of TBR branch swapping (n.b., the TNT

n en Paleobiología y
magnifications, at the Instituto de Investigacio command used is “bb”).
Geología (CONICET-UNRN). The descriptions complement those
performed by Gianechini et al. (2011) and do not delve into details 2.3. Discriminant analysis
that were already explained in the aforementioned work. Eleven
crown-based measurements (i.e., CBL, CBW, CH, AL, MCL, MCW, CTU, In order to classify and predict the optimal classifications of the
MA, MC, DC, DA; Supplementary information 2) were taken on the three shed theropod teeth, and if is possible to discriminate them
best-preserved crowns with a digital caliper of 150 mm (with an from those of other theropods (i.e., family-level) based on quanti-
accuracy of 0.01 mm) following the methodology detailed by tative data, we carried out a discriminant function analysis (DFA)
Hendrickx et al. (2015b, 2020a). The new descriptions were made using the data set compiled by Hendrickx et al. (2015a) and recently
using the dental nomenclature and protocol provided by Hendrickx modified by Young et al. (2019) and Hendrickx et al. (2020a, b). This
et al. (2015b). Furthermore, each tooth was compared with teeth dataset includes fifteen measurements (i.e., CBL, CBW, CH, AL, CBR,
belonging to 156 non-avian theropod species (see Hendrickx et al., CHR, MCL, MCW, MCR, MSL, LAF, LIF, CA, MDL, DCL), taken in 1335
2020a, b, for the list of taxa), with a detailed survey of large- teeth belonging to 89 taxa (84 species and five indeterminate
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J.G. Meso, F.A. Gianechini, R.D. Jua Cretaceous Research 137 (2022) 105250

family-based taxa) separated into 20 monophyletic or paraphyletic Crown overall morphology. This is a ziphodont tooth with a labiolin-
groups (Hendrickx et al., 2020a, b). Data from 39 teeth belonging to gually compressed crown, a distal curvature, a strongly convex
two taxa (i.e., Megaraptor and Murusraptor) were added to the mesial margin and a concave distal margin in lateral view, and a well-
Hendrickx et al. (2020a, b) dataset based on first-hand measure- developed serrated distal carina. The mesial border lacks a carina and
ments of the crowns. The final dataset includes fifteen measure- denticles at its basal half, whereas the apical half is crossed by a
ments, of which only could be used eleven for this study because longitudinal rupture, which prevents discerning whether a carina
the completeness of the material (i.e., CBL, CBW, CH, AL, MCL, MCW, was present in this area. The distal carina bears 14 denticles per
MSL, LAF, LIF, MDL, DCL), taken in 1374 teeth belonging to 91 taxa 5 mm at mid-crown and 16 denticles per 5 mm close to the cervix.
(i.e., 86 species and five indeterminate family-based taxa) sepa- The labiolingual compression of the crown (CBR) is equal to 0.54,
rated into 21 monophyletic or paraphyletic groups (Supplementary whereas is not possible to calculate the exact value of the baso-apical
information 2). Since most researchers measure theropod crowns elongation of the crown (CHR), although the latter ratio is considered
differently (see Hendrickx et al., 2020a, b for further information on to be in a range between a mean to high value (1.5 < CHR  2.5
this tendency), a second analysis was performed on a dataset or > 2.5). The lingual and labial surfaces show a shallow median
restricted to measurements taken by Hendrickx et al. (2020b). depression with a triangular shape in their basal half, being the
Furthermore, because most of the isolated theropod teeth here lingual depression better defined and flanked by subtle ridges.
studied belong to relatively large-sized animals, a third analysis Because of these depressions, the cross-section is figure-eight shaped
was carried out on a dataset restricted to theropod taxa with near the level of the cervix. The crown shows arcuate marginal un-
crowns of more than 20 mm. These two datasets include 764 and dulations on labial and lingual surfaces about the distal margin and a
439 teeth belonging to 55 and 48 theropod taxa each separated into braided enamel texture. Transverse undulations, flutes, longitudinal
14 groups, respectively (Supplementary information 2). The grooves, or ridges (sensu Hendrickx et al., 2015b) are absent.
discriminant function analysis (DFA), it was performed following
Material. MPCA-Pv 249 (Fig. 3).
the protocol by Young et al. (2019), in which all variables were log-
transformed to normalize the quantitative variables. Finally, the Theropoda Marsh, 1881
DFA was run in Past version 3.19 (Hammer et al., 2001) using the Ceratosauria Marsh, 1884
discriminant analysis function, and treating each tooth as unknown Abelisauroidea Bonaparte, 1991
taxa. Abelisauridae Bonaparte and Novas, 1985
Gen. and sp. indet.
2.4. Cluster analysis
Crown overall morphology. The crown of this tooth is labio-lingually
compressed and slightly curved in anteroposterior direction, with
As a third way to classify fallen teeth and be able to visualize
both mesial and distal well-developed serrated carinae and cen-
which theropods at the genus level are grouped in a dendrogram
trally positioned along their margins. The mesial carina bears
based on the measurements of the teeth, a cluster analysis was
approximately 17 denticles per 5 mm close to the cervix, 16 den-
carried out in Past version 3.19 (Hammer et al., 2001). This analysis
ticles per 5 mm at mid-crown, and 18 denticles per 5 mm in the
was based on the different datasets of first-hand measurements by
apex of the crown. The distal carina possesses 16 denticles per
Hendrickx (see Hendrickx et al., 2020a, b), using the paired group
5 mm at the crown base, 12 denticles per 5 mm at mid-crown, and
algorithm and a neighbor-joining clustering, choosing Euclidean
13 denticles per 5 mm close to the apex. The relationship between
distances for the similarity index.
mesial denticles vs. distal denticles (DSDI) is 0.75. All denticles are
asymmetrical and hook-shaped, and the interdenticular spaces are
2.5. Geological settings
broad. Despite lacking its most basal sector at the level of the cervix,
it is possible to estimate its labiolingual compression (CBR), whose
Locality and Horizon. La Bonita locality (39 380 42 S;
range oscillates approximately in 0.54. Concerning to the ratio CHR,
68 330 06 W), located 8 km NW from Cerro Policía town, in north-
it is inferred that it could have a mean value (i.e., 1.5 < CHR  2.5). In
western Río Negro Province, Patagonia, Argentina (Fig. 1); the
lateral view, the mesial margin is convex, while the distal margin is
represented horizon in the site is the uppermost section of the Bajo
straight with its apex positioned at the same level as the distal
de la Carpa Formation, Santonian, middle Upper Cretaceous (Pe
rez
border. Both lingual and labial surfaces are strongly convex. The
et al., 2009; Garrido, 2010).
cross-section is lenticular almost at the level of the cervix. The
crown has marked wrinkles on the enamel separated by sulci that
3. Description
traverse continuously the labial and lingual surfaces. The texture is
smooth and not oriented in any preferential direction.
3.1. State of preservation and general morphology
Material. MPCA-Pv 251 (Fig. 3).
The collected teeth have most of the crown preserved and are
Theropoda Marsh, 1881
interpreted as shed teeth. The three specimens lack the basal most
Ceratosauria Marsh, 1884
sector of the crown, and a small portion of the apical tip of the
Abelisauroidea Bonaparte, 1991
crown is absent in MPCA-Pv 247 and 249. Additionally, the enamel
Abelisauridae Bonaparte and Novas, 1985
is well-preserved on the entire surface of the three crowns. The
Gen. and sp. indet.
denticles are something worn both in the mesial and the distal
carinae, and only very few of them show their original shape. Crown overall morphology. The crown has a convex mesial margin
and a straight distal one, in lateral view, with the apex positioned
Material. MPCA-Pv 247 (Fig. 2)
centrally on the crown, strongly convex lingual and labial surfaces,
Theropoda Marsh, 1881 and mesial and distal carinae with well-developed denticles along
Tetanurae Gauthier, 1986 their entire extension. Also, the mesial and distal carinae are
Megaraptora Benson, Carrano, and Brusatte, 2010 strongly displaced lingually. On the lingual face, there is a concave
Megaraptoridae Novas, Agnolín, Ezcurra, Porfiri, and Canale, 2013 surface adjacent to the mesial carina. In cross-section, almost at the
Gen. and sp. indet. more basal level of the crown, the tooth is salinon-shaped. The
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J.G. Meso, F.A. Gianechini, R.D. Jua Cretaceous Research 137 (2022) 105250

n Basin (A); location map of the Neuque

Fig. 1. Sketch map of the Neuque
n Basin showing the work area in detail (red box) (B); and geological map showing the regional occurrence
of the Bajo de la Carpa Formation at the La Bonita locality, and the fossiliferous provenance site (C). Abbreviations: LP, La Pampa province; MZ, Mendoza province; NQ, Neuque
n
province; RN, Río Negro province. (For interpretation of the references to color in this figure legend, the reader is referred to the Web version of this article.)
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J.G. Meso, F.A. Gianechini, R.D. Jua Cretaceous Research 137 (2022) 105250

Fig. 2. MPCA-Pv 247 in labial (A), lingual (B), mesial (C) and distal view. Detail of the distal denticles (B1, B2 and B3). Red scale bar equal 1 cm and blue scale bar equal 1 mm. (For
interpretation of the references to color in this figure legend, the reader is referred to the Web version of this article.)

labiolingual compression and the baso-apical elongation of the The cladistic analysis performed from the dentition-based data
crown are not possible to calculate, although it can be inferred a matrix without constraints found 100 MPTs (CI ¼ 0.242; RI ¼ 0.586;
low value for the first ratio (i.e., CBR > 0.75) and a mean value for L ¼ 1074; strict consensus tree: CI ¼ 0.237; RI ¼ 0.574; L ¼ 1097). The
the second one (i.e., 1.5 < CHR  2.5). The crown exhibits arcuate strict consensus tree of these unconstrained analyses recovered
marginal undulations on the lingual surface about the mesial and MPCA-Pv 247 strongly nested within Megaraptoridae, accurately as
distal margins. The enamel texture varies from smooth to irregular the sister taxon of the megaraptorid Murusraptor, being Orkoraptor
and transverse undulations, flutes, longitudinal grooves, or ridges the basal member of this subclade (Fig. 5). Regarding MPCA-Pv 249
are absent. The mesial carina shows 12 denticles per 5 mm at mid- and 251, they were found nested within a subclade essentially
crown and close to the apex. The distal carina bears 12 denticles per composed of abelisaurid theropods, and the inclusion the allosauroid
5 mm at mid-crown and close to the apex and possesses 13 den- Erectopus (Fig. 5). The tooth MPCA-Pv 249 is found in a small poly-
ticles per 5 mm close to the cervix. The denticles are chisel-like and tomy that also includes the abelisaurids Aucasaurus and Abelisaurus.
the interdenticular space is broad. The relationship between mesial With respect to MPCA-Pv 251, this tooth is recovered as the sister
denticles vs. distal denticles (DSDI) is 1. taxon of Skorpiovenator, and in turn, it was found as the sister taxa of
a small group comprised of Kryptops þ (Majungasaurus þ Rugops).
4. Analysis The strict consensus tree is well-resolved, with the exception of three
small polytomies containing Coelophysis, Haplocheirus, Liliensternus,
4.1. Cladistic analysis and Arcovenator (Fig. 5).
The cladistic analysis conducted from the tooth-crown-based
Four most parsimonious trees (MPT) were found (CI ¼ 0.197; data matrix without constraints found a poorly resolved strict
RI ¼ 0.462; L ¼ 1318; strict consensus tree: CI ¼ 0.197; RI ¼ 0.460; consensus tree from 100 most parsimonious trees (CI ¼ 0.242;
L ¼ 1322) when using a fully constrained search. The shed tooth RI ¼ 0.626; L ¼ 661; strict consensus tree: CI ¼ 0.202; RI ¼ 0.527;
MPCA-Pv 247 was either recovered within Megaraptoridae, as the L ¼ 793). Even though the strict consensus tree generated is poorly-
sister taxon of Murusraptor in all of the most parsimonious trees resolved and not consistent with the general consensus theropod
(Fig. 4). The two remaining teeth are found within Abelisauridae; phylogeny, the shed tooth MPCA-Pv 247 is here recovered anew as
MPCA-Pv 249 was either found as the sister taxon of “Abelisaurus” the sister taxon of Murusraptor in a subclade including all Patago-
(n.b., this taxon here refers to the specimens MC 1/5/267/689/709 nian megaraptorids (Fig. 6). Concerning MPCA-Pv 249 and MPCA-
referred to Abelisaurus comahuensis by Hendrickx and Mateus, Pv 251, both teeth were recovered in a small subclade that con-
2014; see also Meso et al., 2021a), and MPCA-Pv 251 being recov- tains the abelisaurids Chenanisaurus, Skorpiovenator, Rugops,
ered as the sister taxon of a small subclade formed by Aucasaurus Majungasaurus, Kryptops, Aucasaurus, Indosuchus, and “Abelisau-
and Skorpiovenator (Fig. 4). rus”. Specifically, MPCA-Pv 249 is deeply nested in abelisauridae as

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J.G. Meso, F.A. Gianechini, R.D. Jua Cretaceous Research 137 (2022) 105250

Fig. 3. MPCA-Pv 249 (AeD) and MPCA-Pv 251 (EeH). MPCA-Pv 249 in labial (A), lingual (B), distal (C) and mesial view (D). Detail of the mesial denticles (A1, A2 and A3) and detail of
the distal denticles (B1, B2 and B3). MPCA-Pv 251 in lingual (E), labial (F), distal (G) and mesial view (H). Detail of the mesial denticles (E1 and E2), and detail of the distal denticles
(F1 and F2). Red scale bar equal 1 cm and blue scale bar equal 1 mm. (For interpretation of the references to color in this figure legend, the reader is referred to the Web version of
this article.)

the sister taxon of “Abelisaurus”; whereas MPCA-Pv 251 is recov- account for 48.15% and 19.83% of the total variance, respectively). At
ered as the sister taxon of Skorpiovenator (Fig. 6). the taxon-level, MPCA-Pv 247 is found closely related to Dilopho-
saurus, while MPCA-Pv 249 and 251 are classified as indeterminate
4.2. Discriminant analysis abelisaurids (PC1 and PC2 account for 41.45% and 20.76% of the total
variance, respectively; Supplementary information 3: Table 2). The
The DFA carried out from the whole dataset classified the teeth reclassification rate (RR) is relatively high, being 57.78% at the
here studied as abelisaurids (clade-level analysis; PC1 and PC2 clade-level and 57.92% at the genus-level. However, the

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J.G. Meso, F.A. Gianechini, R.D. Jua Cretaceous Research 137 (2022) 105250

Fig. 4. Strict consensus tree of four most parsimonious trees (CI ¼ 0.197; RI ¼ 0.460; L ¼ 1322) recovered in the cladistic analysis made from the dentition-based data matrix with
constrained search and setting the all teeth as floating terminals.

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J.G. Meso, F.A. Gianechini, R.D. Jua Cretaceous Research 137 (2022) 105250

Fig. 5. Strict consensus tree of 100 most parsimonious trees (CI ¼ 0.237; RI ¼ 0.574; L ¼ 1097) recovered in the cladistic analysis made from the dentition-based data matrix with an
unconstrained search.

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J.G. Meso, F.A. Gianechini, R.D. Jua Cretaceous Research 137 (2022) 105250

Fig. 6. Strict consensus tree of 100 most parsimonious trees (CI ¼ 0.202; RI ¼ 0.527; L ¼ 793) recovered in the cladistic analysis made from the tooth-crown-based data matrix.

reclassification rate is slightly low in the DFA performed when the Dilophosauridae and Abelisauridae (PC1 and PC2 account for
absence of denticles is considered as inapplicable, being 57.9% at 41.95% and 15.9%; Supplementary information 3: Table 2).
the clade-level and 55.23% at the genus-level. In this last analysis, In the discriminant function analysis conducted from the whole
the three specimens are classified again as abelisaurids (clade- dataset with first-hand measurements by Hendrickx (see
level; PC1 and PC2 account for 47% and 18.9%). Conversely, at the Hendrickx et al., 2020a, b), the specimens are classified as Tyran-
clade-level, the specimens are classified also among nosauridae and Abelisauridae (clade level; PC1 49.47% and PC2

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Fig. 7. Results of the discriminant analysis performed at the clade-level from the whole dataset with personal measurements of Christophe Hendrickx on 400 teeth belonging to 46
theropod taxa and 12 groupings along the first two canonical axes of maximum discrimination in the dataset (PC1 49.47% and PC2 24.32% of the total variance, respectively).

24.32%; Fig. 7). At the taxon-level (PC1 41.51% and PC2 19.14%), the slightly higher reclassification rate (at clade-level 58.55% and at
teeth were found closely related to members of Dilophosauridae taxon-level is 63%). Furthermore, in this analysis, the specimens are
and Abelisauridae (see Supplementary information 3: Table 2). classified as belonging non-megalosaurian Megalosauroidea, Dro-
Reclassification rate is slightly better at the taxon-level (59.84%) maeosauridae, and Megalosauridae.
than at the clade-level (53.86%). In this analysis made when the
absence of denticles is considered as inapplicable the results are 4.3. Cluster analysis
same both to clade-level as taxon-level (Supplementary
information 3: Table 2). With respect to the reclassification rate is Results of the cluster analyses performed from the two datasets
also slightly higher at the taxon-level (55.12%) than at the clade- (i.e., whole dataset of Hendrickx et al., 2020a, b), consisting of first-
level (54.96%). By contrast, the teeth are assigned to the non- hand measurements and data set restricted to large-sized teeth)
megalosauran Megalosauroidea, Dromaeosauridae, and Mega- when the absence of denticles is considered as inapplicable or not,
losauridae in the DFA performed from the dataset restricted to taxa using the hierarchical clustering option, recovered the isolated
with teeth larger than two centimeters (clade level; PC1 39.71% and crowns as belonging to Megaraptoridae, Abelisauridae, non-
PC2 27%; Fig. 8). The reclassification rate is better than in the last, tyrannosaurid Tyrannosauroidea, and basalmost Theropoda
being 57.61% (clade-level) and 63% (taxon-level). When considering (Supplementary information 3: Table 3). The cluster analyses using
the absence of denticles as inapplicable, again the result showed a the neighbour joining option, recovered the crowns as members of
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J.G. Meso, F.A. Gianechini, R.D. Jua Cretaceous Research 137 (2022) 105250

Fig. 8. Results of the discriminant analysis performed at the clade-level from the whole dataset with teeth larger than two centimeters, on 725 teeth belonging to 53 theropod taxa
and 12 groupings along the first two canonical axes of maximum discrimination in the dataset (PC1 39.71% and PC2 27% of the total variance, respectively).

Tyrannosauridae, non-averostran Neotheropoda, Metriacanthosaur- Megaraptoridae, recover two synapomorphies: 1) lateral teeth with
idae, and Abelisauridae (Supplementary information 3: Table 3). labial and lingual depressions at the bases of the crowns (i.e.,
crowns with 8-shaped basal cross-section), a feature also present in
5. Discussion the ceratosaur Berberosaurus, the metriacanthosaurid Sinraptor,
tyrannosauroids such as Alioramus, Dilong, and Proceratosaurus, and
5.1. Cladistic and morphometric analyses the allosaurid Allosaurus (Hendrickx et al., 2019, 2020b); 2) a mesial
carina absent in the lateral teeth, a condition also present in the
In the three cladistic analyses is observed a strong affinity that basal tetanuran Chilesaurus, the megalosaurid Sciurumimus, the
allows assigning MPCA-Pv 247, with confidence, as a megaraptorid. tyrannosauroid Guanlong, all compsognathids, the neocoelurosaur
Concerning MPCA-Pv 249 and MPCA-Pv 251, the results agree in Ornitholestes, some ornithomimosaurs, alvarezsaurs such as Aorun,
the identification of these teeth as abelisaurids. The latter two Haplocheirus, Mononykus, and Shuvuuia, the basal oviraptorosaur
specimens vary in position within Abelisauridae, clustering with Incisivosaurus, and some paravians (see Hendrickx et al., 2019,
different abelisaurids in the three phylogenetic analyses (i.e., 2020b). The cladistic analysis performed from the dentition-based
“Abelisaurus”, Skorpiovenator, and Aucasaurus). Thus, our results data matrix without constraints found an additional feature: 3)
support the suggestions proposed by Gianechini et al. (2011). lateral teeth with a concave surface adjacent to the distal carina on
The analyses performed from the dentition-based data matrix the lingual surface (n.b., present in Australovenator and Neovenator,
with constrained search that support the tooth MPCA-Pv 247 as a but unknown for Fukuiraptor). This feature is also observable in
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several non-coelurosaurian theropods, such as Afrovenator, Dilo- Sciurumimus, the megaraptorids Megaraptor, Orkoraptor, and Mur-
phosaurus, Piatnitzkysaurus, and ceratosaurids. This is the first time usraptor, the non-tyrannosaurid tyrannosauroid Guanlong, the
that these morphological characters are found consistently as compsognathids Compsognathus, Juravenator, and Scipionyx, the
synapomorphies of Megaraptoridae in a cladistics analysis. basal neocoelurosaur Ornitholestes, the basal alvarezsaurians Hap-
Using the dataset only based on the tooth-crown characters, locheirus and Aorun, the basal oviraptorosaurs Incisivosaurus, the
other two features were recovered as synapomorphies: 4) lateral dromaeosaurids Bambiraptor, Tsaagan, and Velociraptor, and the
teeth with a depression in the labial surface, centrally positioned on troodontids Zanabazar and Troodon (see Hendrickx et al., 2019).
the crown, and restricted to the crown base or extended along the However, mesial denticles restricted to the apical sector of the
basal half of the crown, a condition present in Australovenator crown only have been observed in the basal alvarezsaurian Hap-
(White et al., 2015a) and probably in Fukuiraptor (see Azuma and locheirus sollers (IVPP V14988). The features observable in shed
Currie, 2000; Currie and Azuma, 2006); and 5) broad inter- tooth MPCA-Pv 247 such as a crown slightly curved distally, enamel
denticular space at mid-crown of the lateral teeth. An inter- wrinkles, a distal carina slightly displaced, distal chisel-like denti-
denticular space more than one-third of the denticle width is cles, and enamel texture braided are relatively common dental
present in Australovenator (White et al., 2015a; Hendrickx et al., features in many groups of theropods (see Hendrickx et al., 2019).
2019) and Neovenator (see Hendrickx et al., 2019). Nevertheless, MPCA-Pv 247 exhibits three features whose distri-
As described in previous studies, the dental synapomorphies bution is limited to non-maniraptoriform averostrans. These fea-
recovered for Abelisauridae are numerous and highly diagnostic tures are: 1 and 2) labial and lingual depressions restricted at the
(see Hendrickx et al., 2020a; Meso et al., 2021a, b; and references crown base, conferring in eight-shape cross-section outline at the
therein). Furthermore, as mentioned by Hendrickx et al. (2020a), cervix level; and, 3) lingual depression well defined, triangular in
the synapomorphies of Abelisauridae are more extensive when is shape and flanked by shallow ridges. All these features were
used the dataset only restricted to tooth-crown-based characters recovered in this study as synapomorphies for Megaraptoridae.
(namely, eleven synapomorphies recovered in this study; one more MPCA-Pv 249 presents a combination of dental features typically
synapomorphy with respect to the analysis performed from the present in the lateral dentition of Abelisauridae such as a straight
dentition-based data matrix with constrained search; and nine distal margin, short and strongly compressed crown, a distal carina
more synapomorphies with respect to the analysis performed from centrally positioned on the distal margin of the crown, symmetri-
the dentition-based data matrix without constrained search). cally convex lingual and labial surfaces, interdenticular sulci pre-
The 2D plots resulting from the PCA conducted from the three sent, and an irregular enamel surface texture (see Hendrickx et al.,
datasets show that the specimens occupy the same region of the 2020a; Meso et al., 2021a, b). With respect to MPCA-Pv 251, it also
morphospace where converge non-megalosauran Mega- shows a combination of features only seen in the mesial dentition
losauroidea, Megalosauridae, non-tyrannosaurid Tyrannosaur- of Abelisauridae, e.g., an irregular enamel surface texture, marginal
oidea, and Tyrannosauridae. Also, MPCA-Pv 249 and 251 are undulations, apicobasally elongated concave surfaces adjacent to
specifically included within the region occupied by Abelisauridae the distal carina on the lingual surface, a labial margin convex and
and Allosauridae, while MPCA-Pv 247 occupies the same mor- lingual margin biconcave conferring a salinon or J-shaped cross-
phospace that Megaraptora. section, apicobasally and proximodistally short denticles in the
The morphometric analyses (i.e., discriminant and cluster basalmost portion of the crown, a similar or lower number of
analysis) recovered mixed results (see Supplementary information denticles at the apex than at mid-crown, and strongly developed
3: Tables 2 and 3), but both indicate that MPCA-Pv 247 shows interdenticular sulci (Hendrickx et al., 2020a; Meso et al., 2021a, b).
strong affinities with megaraptorid teeth, while MPCA-Pv 249 and
251 are related with abelisaurid teeth. This can be explained by: 1) 5.3. Faunal diversity of La Bonita site and paleoecological
the limited data from the sample size as a consequence of the de- occurrence
gree of preservation of the three specimens; or, 2) in the correctly
assigned teeth of Megaraptora and Abelisauridae in the three Since both Abelisauridae and Megaraptoridae have a biochron
datasets a strong oscillation is observable (namely, 27% for the that encompass almost the complete Late Cretaceous and are
megaraptorid teeth and 54.7% for the abelisaurid teeth in the first recorded in most of the rock units containing terrestrial vertebrate
dataset; 13.3% for the megaraptorid teeth and 61.2% for the abeli- faunas in Patagonia, is expected that both lineages were compo-
saurid teeth in the second dataset; and 96% for the megaraptorid nents of the same food chains. However, it is also likely that each
teeth and 63.26% for the abelisaurid teeth in the latest dataset). In lineage had particular ecological preferences (e.g., highland pref-
this sense, Abelisauridae together with Megaraptora possess teeth erence) that signified an environmental separation between them.
and denticles of similar dimensions (e.g., AL, MCL, MCW, MDL, DDL) To this respect, the recovery of evidence of abelisaurids and meg-
to those of Ceratosauridae, Allosauridae, Metriacanthosauridae, araptorids in the same fossiliferous site is scarce. To our knowledge,
Megalosauridae, and Tyrannosauridae. only in the Futalognko site on the northern coast of the Los Bar-
reales lake, corresponding to rocks of the Portezuelo Formation,
5.2. Morphological comparison of the MPCA-Pv teeth and in the Aerosteon site on the Can ~ ado
n Amarillo area of the
Anacleto Formation are the other sites that, like in La Bonita quarry,
The specimen MPCA-Pv 251 is inferred as mesial in position preserve direct evidence of this coexistence based on associated
based on the asymmetrical labial and lingual surfaces and their materials. Another area where abelisaurids and megaraptorids
cross-section outline, in agreement with Gianechini et al. (2011). were found is Cerro Overo-La Invernada, at northern Neuque
n
Concerning MPCA-Pv 247 and 249, they are here considered as Province, where outcrops the Bajo de la Carpa Formation (Filippi
belonging to the lateral dentition based on their labiolingual and Bellardini, 2021; Me
ndez et al., 2021). However, here the ma-
compression and their symmetrical shape (see Hendrickx et al., terials assigned to these two groups come from different sites and
2015b). Unfortunately, in MPCA-Pv 247 is not possible to discern levels in a broader area, and thus the coexistence is difficult to
if a mesial carina is absent, or restricted to the apical part of the corroborate in this case. Since the sedimentary model described for
crown. An unserrated mesial carina and serrated distal carina are the La Bonita quarry corresponds to a fluvial system (Pe
rez et al.,
present in many distantly related theropod taxa with ziphodont 2009) a short transport of the teeth from nearby areas cannot be
teeth, such as the basal theropod Eodromaeus, the megalosaurid dismissed. However, the presence of these teeth around the
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J.G. Meso, F.A. Gianechini, R.D. Jua Cretaceous Research 137 (2022) 105250

skeleton of the holotype of the derived titanosaurian Bonitasaura and 251 show a combination of unequivocal dental features typi-
salgadoi and the good preservation of all teeth allow us to consider cally present in the mesial and lateral dentition of abelisaurid
this association tentatively as indicative of feeding behavior of theropods. MPCA-Pv 247 reveals the presence of a suite of dental
these theropods (either predation or scavenging). features restricted to Megaraptoridae and, at the same time, we
The assignment of MPCA-Pv 247 to Megaraptoridae is not un- recovered a combination of synapomorphies highly diagnostic in
expected, since the record of the Santonian Bajo de la Carpa For- this clade. Therefore, megaraptorids displays several dental
mation includes Tratayenia rosalesi (Porfiri et al., 2018). The anatomical characteristic traits that are supported in our cladistics
presence of this theropod clade together with abelisaurids con- analysis as synapomorphyes of this clade, corroborating that their
forms to the proposed faunal turnover event that could have teeth possess diagnostic features. The assignment of the teeth here
happened at the end of the Turonian (Lamanna et al., 2001; Coria studied to both Abelisauridae and Megaraptoridae, provides evi-
and Salgado, 2005; Jua
rez Valieri et al., 2011; Canale et al., 2015; dence of the coexistence of both lineages in the same ecosystem.
Meso et al., 2021a). In MPCA-Pv 247 and MPCA-Pv 249 are observed
oblique fractures in the mesial and distal carinae, which can be
Acknowledgments
interpreted as injuries caused by the contact between the crown
and hard pieces of the prey, as bones or ostederms, (see Meso et al.,
The authors thank go to Alberto Caselli (Director of the Instituto
2021b, and references therein). The kind of fracture observed in
de Investigacio
n en Paleobiología y Geología, General Roca,
MPCA-Pv 251 is attributed to taphonomic (biostratinomic and
Argentina), for providing the space at the Optic Microscopy Labo-
diagenetic) processes porteriors to the fall of the tooth. Previous
ratory. We also appreciate the valuable corrections and suggestions
studies suggested that the most parsimonious explanation of this
made to this work by Juan Canale and an anonymous reviewer.
accumulation would be a feeding behavior (Alonso et al., 2017;
Meso et al., 2021b). The skeletal patron of the abelisaurids and
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