Cretaceous Research 75 (2017) 81e93

Contents lists available at ScienceDirect

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

First occurrence of tyrannosaurid theropods from the Corral de

Enmedio Formation (Upper Cretaceous) Sonora, Me !xico
Claudia Ine
!s Serrano-Bran~ as a, *, Cirene Gutie
!rrez-Blando a, Rube
!n Duarte Bigurra b,
Carlos Manuel Gonza !lez-Leo!n c
a
Facultad de Ciencias, Universidad Nacional Auto !noma de M! exico, Circuito Exterior, Ciudad Universitaria, Coyoaca!n, 04510, M!exico City, Mexico
b
Posgrado en Ciencias de la Tierra, Estacio
!n Regional del Noroeste, Instituto de Geologa, Universidad Nacional Auto
!noma de M! exico, Apartado Postal 1039,
83000, Hermosillo, Sonora, Mexico
c
Estacio
!n Regional del Noroeste, Instituto de Geologa, Universidad Nacional Auto !noma de M! exico, Apartado Postal 1039, 83000, Hermosillo, Sonora,
Mexico

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

Article history: The Cabullona Basin in the state of Sonora, Mexico is becoming recognized due to its diversity of
Received 16 January 2017 southern Laramidian continental vertebrates, especially dinosaurs. In this study we describe and analyze
Received in revised form three theropod teeth (ERNO specimens) that were found isolated and surface collected in the Corral de
4 March 2017
Enmedio Formation (Cabullona Group, Upper Cretaceous). The three specimens possess similar
Accepted in revised form 18 March 2017
morphological characteristics that match the ones present in Late Cretaceous Laramidian tyrannosaurids,
Available online 20 March 2017
so they were referred to the Tyrannosauridae, probably belonging to a new unknown taxon. The
implementation of statistical and cladistic analyses corroborated their taxonomical assignment. ERNO
Keywords:
Tyrannosaurs
specimens correspond to the rst record of tyrannosaurid dinosaurs in the basal Corral de Enmedio
Dinosaurs Formation, extending the stratigraphic distribution of these dinosaurs in the Cabullona Basin. Although
Paleoecology tyrannosaurids have been previously described in the Cabullona Basin, the ERNO specimens of the Corral
Theropods de Enmedio Formation seem to be different, because they possess more labiolingually compressed teeth.
Cretaceous This new evidence could indicate a higher taxonomic diversity of the tyrannosaurid theropods that were
Mexico present in the Cabullona Basin, adding more information to the Tyrannosauridae diversication on one of
the most southern Laramidian regions during the Late Cretaceous.
2017 Elsevier Ltd. All rights reserved.

1. Introduction
Tyrannosaurids are considered the most dominant, diverse and
best-known group of predatory dinosaurs from Laramidia during
Late Cretaceous time (Loewen et al., 2013). In general, tyranno-
saurid fossils have been documented from several localities ranging
from the North Slope of Alaska (Brouwers et al., 1987; Clemens and
Nelms, 1993), south to the states of Coahuila and Sonora in Mexico
(Herna!ndez-Rivera, 1997; Serrano-Bran ~ as et al., 2014). However,
the bulk of tyrannosaurid remains has been collected from Alberta
and Montana (Currie, 2003; Henderson and Tanke, 2010; Bell and
Currie, 2014; Dalman and Lucas, 2015), and recently from Wyom-
ing, New Mexico, Utah and Texas (Carr and Williamson, 2010; Carr
* Corresponding author. Viaducto Miguel Alema !n #5 Esquina Divisio! n del Norte,
Colonia del Valle, Delegacio
! n Benito Ju!
arez, C.P. 03100, Mexico City, Mexico.
E-mail address: claudiabran399@gmail.com (C.I. Serrano-Bran ~ as).
et al., 2011; Lehman and Wick, 2012; Thomson et al., 2013; Loewen
et al., 2013; Dalman and Lucas, 2016).
To date, several skeletons representing distinct ontogenetic age
groups have been recovered; however, for some localities, isolated
tyrannosaurid teeth represent the only evidence of these theropods
in those places (Serrano-Bran ~ as et al., 2014). Although these fossils
are not as ashy as complete skeletal remains, they are important
elements for understanding the diversity, paleoecology and evo-
lution of dinosaur communities (Fiorillo and Gangloff, 2001;
Sankey et al., 2002; Samman et al., 2005; Buckley et al., 2010;
Larson and Currie, 2013; Torices et al., 2013; Torices et al. 2014;
Dalman and Lucas, 2015).
The Cabullona basin, located in northeastern Sonora, Mexico, is
becoming a great source for the investigation of southern Lar-
amidian continental vertebrates, especially dinosaurs. Particularly,
fossil theropods from this Basin have so far been documented and
described from the Packard Shale and Lomas Coloradas Formations,
belonging to the Tyrannosauridae and Ornithomimidae families

http://dx.doi.org/10.1016/j.cretres.2017.03.015
0195-6671/ 2017 Elsevier Ltd. All rights reserved.
82 C.I. Serrano-Bran
~ as et al. / Cretaceous Research 75 (2017) 81e93
(Serrano-Bran~ as et al., 2014; Serrano-Bran
~ as et al., 2016). However,
the other depositional formations of this Basin have also proven to
be highly fossiliferous, where several evidences of vertebrate,
invertebrate and plant remains are preserved throughout the li-
thologies (Gonza !lez-Leo ! n and Lawton, 1995). In this study, we
describe and analyze three isolated theropod teeth, collected for
the rst time in sediments of the Corral de Enmedio Formation
(Cabullona Group, Upper Cretaceous) in Sonora, Mexico.
2. Geological setting
The Cabullona Basin has a sedimentary ll of almost 4 km in
thickness, named as the Cabullona Group (Fig. 1a). Its depocenter is
approximately 80 km long and 30 km wide, extending into
southern Arizona where it correlates with the Fort Crittenden
Formation. The type section of the Cabullona Group is 2.5 km thick
(Taliaferro, 1933) and includes the following formations, from base
upwards: Corral de Enmedio Formation, Camas Sandstone, Packard
Shale and Lomas Coloradas Formation (Fig. 1b). These units repre-
sent several uvial and lacustrine depositional facies (Gonza !lez-
Leo
! n and Lawton, 1995).
Fig. 1. Study area (1a) and stratigraphic column of the Cabullona Group (1b) indicating the position of the Corral de Enmedio Formation; stratigraphic column of the Corral de
Enmedio Formation (1c).
2.1. Corral de Enmedio Formation
This formation is recorded only in the south-central part of the
Cabullona Basin. The base of this unit is covered, so it has an
incomplete observable thickness of 230 m (Fig. 1c). It consists of
variegated intercalations of mudstones and siltstones with several
lenticular beds of sandstones and limestones. The mudstones and
siltstones correspond to massive, highly bioturbated and laterally
continuous packages that range in color from grayish-black to
moderate red. Local bedding in these strata is vaguely dened;
however, it is marked by intervals of green, yellow, brown and gray
rocks. Sandstone strata are lenticular, thin to medium bedded,
range from ne to coarse grained, and are highly bioturbated.
Common sedimentary structures are represented by large-to small-
scale planar cross-stratications, some trough cross-stratications,
lateral accretion surfaces, convolute laminae and ripple cross-
laminations. On the other hand, thin lenticular limestone beds
are abundant in the mudstones and siltstones, which rarely have
laminated stromatolitic structures or oncolites (especially at the
upper part of the formation). The uppermost 20 m of the formation
correspond to laminated siltstones and lenticular interbeds of ne-
 C.I. Serrano-Bran
~ as et al. / Cretaceous Research 75 (2017) 81e93
Fig. 1. (continued).
grained sandstones with ripple marks and planar cross-beds; also,
some subordinate thin, laterally continuous to lenticular limestone
beds can be found (Gonza !lez-Leo
! n, 1994).
The Corral de Enmedio Formation is highly fossiliferous; several
vertebrate remains such as, turtles, shes and dinosaurs along with
gastropods, bivalves, ostracods and wood fragments are preserved
throughout all the lithologies, particularly in the mudstones and
siltstones (Gonza!lez-Leo! n, 1994; Lucas et al., 1995). A maximum age
of ca. 72 Ma was obtained for the Corral de Enmedio Formation,
based on the youngest population of detrial zircons dated from a
tobaceae sandstone sample collected in the lower part of this for-
mation (Fig. 1c; Gonza !lez-Leo !n et al., 2014).
3. Material and methods
3.1. Morphometrics
All the specimens described and discussed in this paper are
housed at the paleontological collection of the Estacio
!n Regional
del Noroeste (ERNO) in Sonora, Mexico. We followed the method-
ology proposed by Samman et al. (2005) in order to classify qual-
itatively the ERNO specimens to a bone origin. The teeth were
morphologically described and analyzed using the standard dental
terminology for directional terms of Smith and Dodson (2003),
Smith et al. (2005), and Hendrickx et al. (2015a). Tooth
83
measurements are as follows (all in millimeters): Crown height
(CH); Crown base length (CBL); Crown base width (CBW); Mesial
denticle density (ME); Mesial apical (MA), Mesial mid-crown (MC)
and Mesial basal (MB) denticle densities; Average mesial denticle
density (MAVG); Distal denticle density (DI); Distal apical (DA),
Distal mid-crown (DC) and Distal basal (DB) denticle densities;
Average distal denticle density (DAVG) (Smith, 2005; Smith et al.,
2005 and Smith et al., 2007). In order to obtain the above-
mentioned measurements, standard calipers were used. Denticles
were quantied by determining the number of denticles per 5 mm
of carina length counted as close to the base, mid-point, and apex as
possible thus generating the basal (MB and DB), mid-crown (MC
and DC), and apical densities (MA and DA). Denticle photographs
were taken with a Zeiss Stemi 508 stereoscopic microscope.
Data sets of dentition measurements provided by Currie and
Varrichio (2004), Smith et al. (2005), Coria and Currie (2006),
et al. (2010), Larson and Currie (2013), and
Longrich (2008), Osi
Hendrickx et al. (2015b) were used as the comparison standard for
our taxonomically unknown specimens (Appendix 1). The taxa
inside these sets include: Daspletosaurus (Russell, 1970), Gorgo-
saurus (Lambe, 1914; Holtz, 2001; Currie, 2003), Tyrannosaurus rex
(Leidy, 1856), Acrocanthosaurus (Stovall and Langston, 1950),
Carcharodontosaurus (Stromer, 1931), Giganotosaurus (Coria and
Salgado, 1995), Indosuchus (von Huene and Matley, 1933), Majun-
gatholus (Sues and Taquet, 1979), Baryonyx (Charig and Milner,
84 C.I. Serrano-Bran
~ as et al. / Cretaceous Research 75 (2017) 81e93

1986), Suchomimus (Sereno et al., 1998), Dilophosaurus (Welles, Table 2

1970), Dromeosaurus (Mathew and Brown, 1922), Sau- Principal components analysis eigenvalue and variation percentage for the ERNO
specimens combined with the theropod standard data set used in this study.
rornitholestes (Sues, 1978), Atrociraptor (Currie and Varricchio,
2004), Spinosaurus (Stromer, 1915), Torvosaurus (Galton and Principal component Eigenvalue Percentage of variance
Jensen, 1979), Afrovenator (Sereno et al., 1994), Duriavenator 1 0.33322 93.695
(Benson, 2008), Megalosaurus (Buckland, 1824), and Dubreuillo- 2 0.01279 3.5971
saurus (Allain, 2005). 3 0.00950 2.6735
4 0.00011 0.0321
5 0.00008 0.0023
3.2. Statistical analyses

A series of statistical analyses were applied to assess the effec- 4. Results

tiveness in correlating morphological characters with quantitative
data; therefore, ERNO specimens were compared against the
standard data set in order to predict their taxonomy (Tables 1 and 5
and Appendix 1). Following Smith et al. (2005), the ERNO isolated
crowns were tested on the basis of morphological congruence, so
teeth with likely taxonomic afnities would correlate more closely.
A principal component analysis (PCA) was run using a variance-
covariance matrix that included CBL, CBW, CH, CBR and CHR as
variables for 21 dinosaur groups (20 dinosaur genera included in
the standard data set plus the three ERNO specimens catalogued as
unknown). All data were log-transformed and the missing values
for the measurements were estimated using a mean value from
across the sample for that particular measurement. This analysis
was conducted in order to explore the morphospace overlap be-
tween different groups. Also, a discriminant function analysis (DFA)
was performed using the same tooth database, to assess the
effectiveness in correlating isolated crowns with taxonomically
identied taxa in order to seek a specic group membership. This
analysis uses pre-determined groups (e.g, taxonomic family clus-
ters) for creating a morphospace, where these groups are maxi-
mally separated. In this way, the DFA assigns a sample unit (e.g.
teeth of unknown afnities) into one of these pre-determined
groups according to the classication rule (Daz-Monroy and
Morales-Rivera, 2012). From this analysis, the classication proba-
bility of the specimens was calculated in each group. Afrovenator
was eliminated for not complying with the assumption of having
more samples than the number of variables. Statistical analyses and
plots for PCA were generated using PAST software v2.17 (Hammer
et al., 2001); whereas, for the DFA, we employed the Statistica 8
software (StatSoft, 2007).
3.3. Phylogenetic analysis
A cladistic analysis was performed in order to explore the
possible relationships of the ERNO specimens with other dinosaur
groups. We ran this analysis by including ERNO 8027, ERNO 8581
and ERNO 8582 in a discrete character-taxon data matrix of
dentition-based characters (Appendix 2), composed by 60 main
theropod taxa (terminals) and 141 characters (Hendrickx and
Mateus, 2014). Characters and character states are described in
Appendix 3. The phylogenetic analysis was conducted using TNT
software (Goloboff et al., 2008). Eoraptor was chosen as outgroup.
For institutional abbreviations, see Appendix 4.
4.1. Systematic palaeontology
Dinosauria Owen, 1842
Theropoda Marsh, 1881
Averostra Paul, 2002
Tetanurae Gauthier, 1986
Avetheropoda Paul, 1988
Tyrannosauridae Osborn, 1905
Material. Three isolated teeth (ERNO 8027, ERNO 8581 and ERNO
8582); gures 2a, 2b, 2c, 3, 4 and 5.
Locality and horizon. Corral de Enmedio Formation (ERNO 8027: 12R
614999, 3444244 UTM coordinates; ERNO 8581 and ERNO 8582:
12R 614800, 3444100 UTM coordinates), Cabullona Group; age:
Campanian.
Measurements. Table 1, Appendix 1.
Description. These ERNO teeth corresponded to ziphodont shed
crowns, since the root is completely absent from all of them and the
transverse width of each tooth base is less than 60% of the mesio-
distal length (Brusatte et al., 2010; Loewen et al., 2013). These
specimens have different degrees of preservation ranging from well
to medium preserved. According to Samman et al. (2005), ERNO
8027, ERNO 8581 and ERNO 8582 correspond to distal maxillary or
dentary teeth because they have an ovoid cross-sectional outline
(Fig. 2a6 and 2b6; Table 1). Each one of the specimens is described
as follows:
ERNO 8027
Crown morphology. This tooth is the most complete; although the
tip of the apex is broken (Fig. 2a5). Both carinae (mesial and distal)
are serrated, but only the distal one reaches the crown cervix
(Fig. 2a1 and 2a2). The mesial carina extends convexly along the
midline and deviates towards the lingual side, disappearing at the
middle part of the tooth (Fig. 2a3). The distal carina is concave and
runs close to the lingual side (Fig. 2a4).
Crown surface. The enamel surface is poorly preserved. It is mostly
cracked and abraded on both the lingual and labial sides (Fig. 3a1);
however in some areas, a braided texture can be observed
(Fig. 3a2). No signicant enamel structures (e.g. enamel bands,
longitudinal ridges or grooves) can be observed on this specimen.
Denticles. Mesial and distal denticles reduce gradually in size along
the carinae, where the smaller ones are located towards the tooth
base. In both carinae, all denticles are diagonally inclined towards
the tooth apex. The mesial carina has a MAVG of 10 denticles per

Table 1
Measurements of the ERNO specimens (isolated crowns) from the Corral de Enmedio Formation.a

Taxa Specimen number CBL CBW CH CBR CHR MA MC MB DA DC DB

Unknown ERNO 8025 32 18 82 0.56 2.56 9 9 14 10 11 14

Unknown ERNO 8581 26 14 52 0.53 2.0 9 10 13 10 19 e
Unknown ERNO 8582 24 14 37 0.58 1.54 e e e e 18 e
a
Abbreviations: Crown base length (CBL); crown base width (CBW); crown height (CH); crown base ratio (CBR); crown height ratio (CHR); mesial apical (MA); mesial mid-
crown (MC); mesial basal (MB); distal apical (DA); distal mid-crown (DC); distal basal (DB) denticle densities.
 C.I. Serrano-Bran
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Table 3
Matrix classication count table indicating the discriminant function accuration in the classication of the observed data. Rows represent the actual groups, whereas columns
correspond to the predicted groups.a

Actual taxa Percent correct Car Tyr Dil Abe Spi Dro Meg Un Total

Carcharodontidae 61.538 24 4 0 3 3 0 4 1 39
Tyrannosauridae 85.074 1 114 0 7 6 0 6 0 134
Dilophosauridae 25.000 0 0 1 3 0 0 0 0 4
Abelisauridae 34.375 0 2 0 11 6 6 7 0 32
Spinosauridae 89.583 0 3 0 2 43 0 0 0 48
Dromaeosauridae 88.000 0 0 0 5 1 44 0 0 50
Megalosauridae 53.125 2 1 0 2 4 5 17 1 32
Unknown 66.666 0 0 0 0 0 0 1 2 3
Total 74.853 27 124 1 33 63 55 35 4 342
a
Abbreviations: Carcharodontidae (Car); Tyrannosauridae (Tyr); Dilophosauridae (Dil); Abelisauridae (Abe); Spinosauridae (Spi); Dromaeosauridae (Dro); Megalosauridae
(Meg) and Unknown (Un; ERNO specimens).

Table 4 Table 5
CBW: CBL ratios for the theropod data set including ERNO 8027. Denticle averages for the theropod data set including ERNO specimens.a,b,c

Taxa Family CBR CHR Taxa ME MAVG DI DAVG

Unknown Unknown 0.562* 2.562* Unknown 9e14 12.5 10e19 14.5

Acrocanthosaurus Carcharodontosauridae 0.577 2.016 Acrocanthosaurus 13.3e16.9 15.1 13.3e16.2 14.75
Carcharodontosaurus Carcharodontosauridae 0.405 1.938 Carcharodontosaurus 8.7e10 9.35 9.8e11.4 10.6
Giganotosaurus Carcharodontosauridae 0.474 2.030 Giganotosaurus 8e10 9 7.5e9.0 8.25
Gorgosaurus Tyrannosauridae 0.634 1.967 Gorgosaurus 8.7e13.7 11.2 11.5e15 13.25
Daspletosaurus Tyrannosauridae 0.714 2.228 Daspletosaurus 11.6e13.9 12.75 10.5e14 12.25
Tyrannosaurus Tyrannosauridae 0.692 1.838 Tyrannosaurus 7e14.4 10.7 7.2e15.2 11.2
Dilophosaurus Dilophosauridae 0.589 2.035 Dilophosaurus e e 14e15 14.5
Indosuchus Abelisauridae 0.688 1.705 Indosuchus 8.5e13.8 11.15 9.30e12 10.65
Majungatholus Abelisauridae 0.575 1.855 Majungatholus 9.4e15 12.2 9.5e14 11.75
Baryonyx Spinosauridae 0.837 2.248 Baryonyx e e 35 35
Suchomimus Spinosauridae 0.754 2.652 Suchomimus e e 27e35 31
Spinosaurus Spinosauridae 0.890 2.513 Spinosaurus e e e e
Dromeosaurus Dromeosauridae 0.575 1.703 Dromeosaurus 18.8e28 21.7 15e22.5 19
Saurornitholestes Dromeosauridae 0.477 1.744 Saurornitholestes 5.18e8.64 6.60 3.58e5.44 4.42
Atrociraptor Dromeosauridae 0.510 1.614 Atrociraptor 2.7e8 5 2.3e4.7 3.37
Torvosaurus Megalosauridae 0.507 2.355 Torvosaurus e e 7e9.5 8.25
Afrovenator Megalosauridae 0.420 2.210 Afrovenator e e 10 10
Duriavenator Megalosauridae 0.486 1.910 Duriavenator e e 11e12.5 11.75
Megalosaurus Megalosauridae 0.563 2.106 Megalosaurus e e 8.75e20 14.37
Dubreuillosaurus Megalosauridae 0.498 1.746 Dubreuillosaurus e e 11.5e16 13.75
a
Unknown ERNO 8027. Abbreviations: Mesial denticle density (ME); average mesial denticle density
*CBR and CHR ratios of unknown specimens were calculated without considering (MAVG); distal denticle density (DI); average distal denticle density (DAVG).
b
ERNO 8581 and ERNO 8582, because they are incomplete. Unknown taxa ERNO specimens.
c
Denticle averages of unknown specimens were calculated without considering
5 mm (Table 1). Most of the mesial denticles are broken; however, a ERNO 8582.

few of them are still preserved. In lateral view, these denticles are
wider (labially-lingually) than longer (proximo-distally), which
confers on them a stout form (Fig. 3b). Laterally, the external
margin is mostly rounded; but, in distal view, it is acute, giving
them a chisel-like shape. Also, they have deep interdenticular
spaces and sulci between them. On the other hand, the distal carina
is more strongly developed and has a DAVG of 11.6 denticles per
5 mm (Table 1). As in mesial denticles, the distal ones are stout and
have a squared, chisel-like shape (Fig. 3c); they possess deep
interdenticular spaces, but the interdenticular sulci are a little
deeper in comparison with mesial denticles. In all denticles (mesial
and distal), the interdenticular sulci are diagonally oriented basally
away from them and run parallel to each other, extending from the
base of the interdenticular space on both labial and lingual crown
surfaces.
ERNO 8581
Crown morphology. This tooth is fractured towards the base, but the
apical half is well-preserved (Fig. 2b1, 2b2 and 2b5). A large wear-
facet is present at the apex on the labial side of the specimen
(Figs. 2b2 and 4a1). Both carinae are serrated; the distal carina is
concave and runs close to the lingual side (Fig. 2b4). The mesial
carina is convex and extends along the midline; it also deviates
towards the lingual side, disappearing at the middle part of the
tooth, as in ERNO 8027 (Fig. 2b3).
Crown surface. The enamel surface is well-preserved on both sides
(labial and lingual), showing a braided texture (Fig. 4a1 and 4a2). A
series of transverse, parallel, high-relief enamel bands (enamel
wrinkles sensu Brusatte et al., 2007) can be observed, particularly
on the labial surface (Fig. 4a1). These wrinkles are more numerous
towards the apex and originate at the base of some denticles,
especially on the distal carina. The spacing between them is ~1 mm.
Denticles. As in ERNO 8027, the mesial and distal denticles reduced
gradually in size along the carinae towards the tooth base and are
diagonally inclined towards the apex. The mesial carina in this
tooth has a MAVG of 10.6 denticles per 5 mm (Table 1). Most of the
mesial denticles are broken (Fig. 4b); however, a few of them are
preserved. These denticles are stout and the external margin is
mostly rounded; whereas, in distal view, they have a squared,
chisel-like shape. The interdenticular spaces and sulci are shallower
and a little smaller than in ERNO 8027. The distal carina has an
average serration of 14.5 denticles per 5 mm (Table 1). As in mesial
denticles, the distal ones are stout and have a chisel-like shape
(Fig. 4c); however, they possess deeper interdenticular spaces and
sulci in comparison with those of the mesial carina. All inter-
denticular sulci of the mesial and distal carinae are also diagonally
oriented towards the apex and run parallel to each other, extending
86 C.I. Serrano-Bran
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Fig. 2. ERNO teeth from the Corral de Enmedio Formation in labial (a1, b2, c1); lingual (a2, b1, c2); mesial (a3, b3, c3); distal (a4, b4, c4); apical (a5, b5, c5) and cross-sectional (a6,
b6, c6) views.
from the base of the interdenticular space on both labial and lingual
crown surfaces.
ERNO 8582
Crown morphology. The apical and basal sections of the tooth are
fractured (Fig. 2c1, 2c2 and 2c5); however, the remaining middle-
part is well-preserved. Although this specimen is broken, it
seems to belong to a smaller individual compared with ERNO 8027
and ERNO 8581 (Table 1; Fig. 2). The mesial carina is abraded, and
only part of the distal carina is preserved (Fig. 2c3 and 2c4). This
carina is serrated, concave and runs close to the lingual side
(Fig. 2c4).
Crown surface. The enamel surface of this specimen is well-
preserved, showing a braided texture (Fig. 5a and 5b). Several
transverse, parallel and high-relief spaced enamel bands (enamel
wrinkles sensu Brusatte et al., 2007) can be observed on both
lingual and labial surfaces (Fig. 5a). They also originate at the base
of some denticles, especially from the distal carina. These wrin-
kles are more numerous and closely spaciated towards the apex
(six enamel bands per 5 mm) than towards the base (three bands
per 5 mm), where they are more visible.
Denticles. The mesial carina is not preserved, and only part of the
distal carina can be described; so the average denticle densities
(MAVG and DAVG) cannot be calculated for this specimen. All
denticles present have approximately the same size and are not
inclined towards the tooth apex (Fig. 5c and 5d). Also, they are more
numerous and relatively smaller than the ones present in ERNO
8027 and ERNO 8581 (Table 1). Most of the denticle tips are frac-
tured; however, one of them is complete and has a rounded margin.
In lateral view, these denticles are a little longer (proximo-distally)
than wider (labially-lingually), which confers them a slightly rect-
angular form (Fig. 5c). In distal view, the only complete denticle has
an acute margin, which gives it a chisel-like shape. The shallow
interdenticular spaces and sulci are not inclined towards the apex;
on the contrary, they have a transverse position with respect to the
main tooth axis, and are parallel between each other (Fig. 5c
and 5d).
 C.I. Serrano-Bran
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Fig. 3. ERNO 8027 in labial view showing a cracked and abraded enamel surface (a1); scale bar 2 cm. Patch of complete enamel, belonging to the red-squared area in (a1), where a
braided texture can be observed (a1). Mid-crown denticles from mesial (b) and distal (c) carinae. Scale bar 2 mm. Abbreviations: interdenticular space (IDSP); interdenticular
sulcus (IDS).(For interpretation of the references to colour in this gure legend, the reader is referred to the web version of this article.)

Fig. 4. ERNO 8581 in labial view showing a large wear-facet present at the apex and several transverse enamel bands on the crown surface (a1); scale bar 2 cm. Braided enamel
texture (a2), belonging to the red-squared area in (a1). Apical-crown denticles from mesial (b) and distal (c) carinae. Scale bar 2 mm. Abbreviations: wear facet (WF); transverse
enamel bands (TEB); interdenticular space (IDSP); interdenticular sulcus (IDS).
88 C.I. Serrano-Bran
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Fig. 5. ERNO 8582 in lingual view showing several transverse enamel bands on the crown surface (a); scale bar 1 cm. Braided enamel texture (b). Mid-crown denticles from the
distal carina (c). Transversal position of the denticles from distal carina (d). Scale bar 2 mm. Abbreviations: transverse enamel bands (TEB); interdenticular space (IDSP);
interdenticular sulcus (IDS).
4.2. Statistical discrimination of the ERNO specimens
The principal component analysis (PCA) showed that the rst
three eigenvalues account for 99.96% of the variance; where the
rst component explained 93.69% of the observed variation among
all specimens (Table 2). Furthermore, the Jolliffe cut-off value for
eigenvalues, which is a method that shows how many principal
components (PCs) should be considered signicant, was 0.049791
for this study. This method retains the PCs associated with the
covariance matrix that have eigenvalues greater in magnitude than
the average of all the eigenvalues (Jolliffe, 1986; Mutsvangwa and
Douglas, 2007). The rst principal component had a greater
eigenvalue than the cut-off value (0.333228); whereas, the second
and third PCs had smaller eigenvalues (0.012793 and 0.009508,
respectively). In this way, the resulting PCA revealed that all three
ERNO specimens plot really close to many tyrannosaurid teeth,
along with some carcharodontosaurid teeth and one megalosaurid
tooth. So, our specimens fall within the maximum morphospace
occupation area (i.e. convex hull) of tyrannosaurids, carchar-
odontosaurids and megalosaurids (Fig. 6). Additionally, they also
fall within the 95% condence interval ellipse of these three groups.
Therefore, the PCA analysis indicates that ERNO 8027, ERNO 8581
and ERNO 8582 are more similar to the Tyrannosauridae, Carch-
arodontosauridae and Megalosauridae families.
On the other hand, the discriminant analysis (DFA) produced a
classication matrix and scores using 20 taxa grouped in 7 families,
our three specimens and four variables, delineating the dental
morphospace occupied by the ERNO specimens and the included
taxa (Table 3). This analysis showed that a total of 74.853% of all
theropod teeth included in the database were classied correctly,
indicating that the DFA returned reasonable results. Particularly,
the ERNO specimens had a correct assignment of 66.666%; where,
ERNO 8027 and ERNO 8581 were classied as unknown, while
ERNO 8582 was misclassied as a megalosaurid. Additionally, the
analysis placed ERNO 8027 and ERNO 8581 within the convex hulls
for dilophosaurids, dromaeosaurids, spinosaurids and tyranno-
saurids (in signicance order from the less signicant to the most
signicant) and within the 95% condence ellipses of these groups;
while ERNO 8582 was placed within dromaeosaurids and tyran-
nosaurids (also in signicance order).
4.3. Phylogenetic analysis
In order to explore the relationships between the ERNO speci-
mens and other dinosaurs, we performed a phylogenetic analysis
using a data matrix of dentition-based characters (Appendix 2;
Hendrickx and Mateus, 2014), which included 60 main theropod
taxa. This analysis resulted in 3 most parsimonious trees of 913
steps, with a consistency index of 0.346 and a retention index of
0.517. The strict consensus topology is moderately well resolved
and shows that ERNO 8027, ERNO 8581 and ERNO 8582 are posi-
tioned within the Tyrannosauroidea (characters 2, 26, 36, 37, 40, 46,
47, 48 and 82) and particularly, they are well-nested within the
Tyrannosauridae family (characters 71, 97 and 99; Fig. 7).
5. Discussion
5.1. Tyrannosaurid features of ERNO specimens and morphological
comparisons to the dentition of other theropods
Theropods belonging to Therizinosauria, Troodontidae, Coelo-
physidae, Compsognathidae, Noasauridae and Spinosauridae have
highly specialized dentitions from which the ERNO specimens can
be easily differentiated. Therizinosaurs and troodontids possess
leaf-shaped crown teeth with a constricted cervix; these teeth are
unserrated or bear very few serrations, where denticles can be
minute or large and pointed (Zanno, 2010; Hendrickx and Mateus,
2014). Coelophysids and compsognathids have small crowns
 C.I. Serrano-Bran
~ as et al. / Cretaceous Research 75 (2017) 81e93 89

Fig. 6. Principal component analysis (PCA) for 21 dinosaur groups (20 dinosaur genera grouped in 7 families, included in the standard data set, and three ERNO specimens
catalogued as unknown), using CBL, CBW, CH, CBR and CHR as variables. This gure particularly shows the plot of the ERNO specimens within the maximum morphospace
occupation area (i.e. convex hull) of tyrannosaurids, carcharodontosaurids and megalosaurids. Abbreviations: Carcharodontidae (blue square); Tyrannosauridae (blue triangle);
Dilophosauridae (pink square); Abelisauridae (gray cross); Spinosauridae (yellow triangle); Dromaeosauridae (green diamond); Megalosauridae (gray oval) and Unknown (black
circle; ERNO specimens).(For interpretation of the references to colour in this gure legend, the reader is referred to the web version of this article.)

(CH < 15 mm), which lack a serrated mesial carina in mesial teeth,
whereas the distal carina possesses minute denticles (Buckley and
Currie, 2014; Hendrickx and Mateus, 2014). Noasaurids possess
small teeth (CH < 15 mm) with strongly twisted mesial carinae and
a uted lingual surface (Hendrickx et al., 2015b). Dromaeosaurids
have labiolingually compressed crowns with mesial and distal
serrated carinae; however, some genera are devoid of serrated
carinae or the mesial most teeth often lack mesial serrations;
denticles are large and sometimes apically inclined (Hendrickx
et al., 2015b). Spinosaurids have subcircular, uted and deeply
veined crowns with minute or absent serrations on the mesial and
distal carinae (Charig and Milner, 1997; Sues et al., 2002; Hendrickx
and Mateus, 2014).
On the other hand, theropods from the Ceratosauridae, Mega-
losauridae, and Carcharodontosauridae families, as well as, Tyran-
nosauroidea and some basal allosauroids possess similar large,
elongated and labiolingually compressed teeth, where the carinae
bear coarse, symmetrical to asymmetrical, chisel-like denticles
often with deep and elongated interdenticular sulci. These groups
display similar crown structures such as, large transversal un-
dulations, short marginal undulations, and have an oriented
enamel texture (Hendrickx and Mateus, 2014). Particularly, cera-
tosaurids have lingually uted mesial teeth with mesial carinae
that extend to the cervix, and labially offset distal carinae. Mega-
losaurids possess teeth with centrally positioned mesial carinae
that do not reach the cervix, whereas the distal carinae can be
either centrally positioned, weakly sigmoid, bowed lingually or
slightly offset labially (e.g. Afrovenator and Dubreuillosaurus), and
terminate well-below the crown cervix. Carcharodontids have a
characteristic sigmoid distal prole, where mesial carinae face
medially or mesiolabially and can terminate well above the cervix.
Allosaurids possess thicker crowns with lingually twisted mesial
carinae that extend to the cervix in mesial teeth, and strongly
labially displaced distal carinae in lateral teeth (Hendrickx et al.,
2015b).
In the case of abelisaurids, these theropods possess low and
weakly recurved teeth, so the distal prole is either straight or
convex in lateral view; the enamel surface is irregular (non-
oriented), unlike the braided or veined (oriented) enamel texture of
most neotheropods; the mesial carina is often centrally positioned
but also can be little twisted towards the root of the tooth (e.g.
Abelisauridae isolated tooth ML 966; Hendrickx and Mateus, 2014),
whereas the distal carina is centrally positioned. Both carinae are
serrated and always reach the cervix. Some Abelisauridae taxa have
hooked denticles (e.g. Majungasaurus, Rugops and Kryptops);
however, they might not be present in all teeth along the tooth row
(e.g. Majungasaurus crenatissimus; Abelisauridae isolated teeth ML
327 and ML 966; Hendrickx and Mateus, 2014). All of the denticles
show deep interdenticular sulci between them (Hendrickx and
Mateus, 2014; Hendrickx et al., 2015b).
Finally, tyrannosaurids have labiolingually compressed crowns
with an ovoid cross-sectional base. They possess serrated offset
carinae, where the mesial carina extends convexly along the
midline and disappears at the middle part of the tooth, whereas the
distal carina is concave, runs closely to the lingual side and reaches
the crown cervix. Both carinae have stout, chisel-shaped denticles,
which decrease in size towards the base and the apex, and regularly
are inclined towards the tooth tip. The enamel texture is braided
and regularly has transverse, parallel, high-relief enamel bands
(Samman et al., 2005; Smith, 2005; Hendrickx et al., 2015b).
In general, the ERNO specimens possess larger ziphodont teeth
with an ovoid cross-section, and lack a constricted cervix (e.g.
ERNO 8027). Both carinae are offset and serrated bearing regular-
sized, chisel-shaped denticles. The mesial carina extends convexly
along the midline and deviates towards the lingual side, dis-
appearing at the middle part of the tooth (e.g. ERNO 8027 and ERNO
8581); whereas the distal carina reaches the crown cervix (e.g.
ERNO 8027) and runs closely to the lingual side. The enamel surface
of these teeth is braided, not uted or veined; and displays some
transverse, parallel, high-relief enamel bands (e.g. ERNO 8581 and
ERNO 8582). Therefore, all three ERNO specimens possess similar
morphological characteristics that match the ones known for Late
Cretaceous Laramidian tyrannosaurids.
Additionally, it is well known that the teeth from most tyran-
nosaurid genera are labiolingually nearly as thick as mesiodistally
long (e.g. Albertosaurus, Gorgosaurus and Daspletosaurus), and that
90 C.I. Serrano-Bran
~ as et al. / Cretaceous Research 75 (2017) 81e93

Fig. 7. Results of the phylogenetic tree analysis between ERNO specimens and different theropod groups using TNT software (Goloboff et al., 2008). Strict consensus tree showing
the position of ERNO 8027, ERNO 8581 and ERNO 8582 specimens within Tyrannosauridae.

Tarbosaurus, Tyrannosaurus and Zhuchengtyrannus possess the most
robust teeth of all tyrannosaurid dinosaurs (e.g. Hurum and Currie,
2000; Brochu, 2003; Hurum and Sabath, 2003; Smith, 2005; Hone
et al., 2011; Dalman and Lucas, 2015, 2016). In the case of the
ERNO specimens, although they resemble the teeth of several
Campanian albertosaurines and tyrannosaurines in labial and
lingual views, they are more labiolingually compressed than those
(Appendix 1; Table 4). On average, these tyrannosaurids have higher
CBW: CBL ratios (i.e. Gorgosaurus 0.634, Daspletosaurus 0.714,
Tyrannosaurus 0.692; Table 4) indicating that their teeth are more
robust; whereas the ERNO specimens (in this case only ERNO 8027
was considered for calculating the CBW: CBL ratio, because it was
more complete than ERNO 8581 and ERNO 8582) had a smaller ratio
(0.562; Table 4), implying the presence of more slender teeth. On the
other hand, Currie (2003) has stated before that juvenile tyranno-
saurid specimens have signicantly labiolingually compressed
teeth; however, ERNO 8027 does not correspond to a young indi-
vidual, due to its large size (Table 1; Appendix 1). Therefore, this
evidence could suggest the presence of a new unknown taxon in the
Corral de Enmedio Formation, which is different from the northern
Laramidian genera and from the other tyrannosaurids present in the
Cabullona Basin (e.g. Tyrannosaurus rex; Serrano-Bran~ as et al., 2014).
In fact, it appears that the ERNO specimens are more similar to some
indeterminate tyrannosaurid teeth found in the Claggett Formation
(Elk Basin, Montana e Wyoming) and in the San Juan Basin of New
Mexico (Dalman and Lucas, 2016), which are also more
 C.I. Serrano-Bran
~ as et al. / Cretaceous Research 75 (2017) 81e93
labiolingually compressed. However, the need for more extensive
sampling is required in order to corroborate this assumption.
5.2. Statistical taxa discrimination of ERNO specimens
The PCA results performed in this study showed that all ERNO
specimens fall within the convex hulls and the 95% condence
interval ellipses of tyrannosaurids, carcharodontosaurids and
megalosaurids, indicating a greater similarity with these groups. In
the case of the DFA, although the analysis could not recover the
ERNO specimens within a particular theropod family (i.e. ERNO
8027 and ERNO 8581 were classied as unknown taxa, while ERNO
8582 was misclassied as a megalosaurid), these teeth also fall
within the convex hulls for dilophosaurids, dromaeosaurids, spi-
nosaurids and tyrannosaurids (in signicance order). Now, as
stated before in Section 5.1, all ERNO specimens possess tyranno-
saurid features that can distinguish them from the other theropod
groups; therefore the statistical analyses (PCA and DFA) along with
the morphological comparison of the teeth, point to a tyranno-
saurid identication for the ERNO specimens. On the other hand,
the misclassication of ERNO 8582 in the DFA could be due to the
fact that this tooth is more fractured than the other specimens and
because it could belong to a juvenile (Table 1; Appendix 1). It has
been pointed out that taxa identication of isolated teeth from
juvenile individuals could be problematic, because they possess
higher values on denticular densities (as in the case of ERNO 8582;
see Table 1) and have morphologies that are different enough from
adults, resulting in misleading classications (Smith et al., 2005;
Serrano-Bran ~ as et al., 2014).
5.3. Phylogenetic position of ERNO specimens
The use of several morphological qualitative characters within a
cladistic framework together with quantitative data, have been
suggested in order to help in the identication of isolated crowns
(Hendrickx and Mateus, 2014; Hendrickx et al., 2015b; Csiki-Sava
et al., 2016). In this study, the cladistic analysis revealed that all
three ERNO specimens form a well-supported clade nested within
the Tyrannosauridae family (Fig. 7), based on the absence of a
concave surface adjacent to carinae along the crown and a higher
number of denticles basally than at the mid-crown, both on the
mesial and distal carinae (characters 71, 97 and 99). In this way,
ERNO 8027, ERNO 8581 and ERNO 8582 seem to be more related to
derived tyrannosaurids, represented here by the genus Tyranno-
saurus; however, in order to obtain a better resolution of the
phylogenetic relationships of these ERNO specimens, additional
and more complete comparative material will be required.
5.4. Biogeographic implications of the ERNO specimens
Theropod dinosaurs have been considered an ecologically
diverse component of terrestrial ecosystems since their appearance
during the Late Triassic (Benson et al., 2010). Particularly for
western North America (Laramidia), the origin and diversication
of the Tyrannosauridae family through the Late Cretaceous had a
profound impact in the dinosaur faunas, where several genera and
species belonging to this family appeared and diversied (e.g.
Gorgosaurus libratus, Daspletosaurus torosus, Teratophoneus curriei,
Bistahieversor sealeyi, Lytronax argestes and Tyrannosaurus rex),
becoming the apex predators in this region (Loewen et al., 2013;
Serrano-Bran~ as et al., 2014).
It is well-known that isolated theropod dinosaur teeth are
frequently used to extend the stratigraphic and biogeographic
ranges of taxa known elsewhere from more complete specimens
(Brusatte et al., 2007). In this case, the ERNO specimens represent
91
the rst record of tyrannosaurid dinosaurs in the Corral de Enmedio
Formation, extending the stratigraphic distribution of these thero-
pods within the Cabullona Basin in northeastern Sonora, Mexico.
Although, tyrannosaurids have been previously described in the
upper Lomas Coloradas Formation from this Basin, the specimens of
the basal Corral de Enmedio Formation are different, due to the fact
that their teeth are more labiolingually compressed, whereas the
others are by far more robust (Serrano-Bran
~ as et al., 2014). As stated
before, this evidence could suggest the presence of a new type of
tyrannosaurid in the Corral de Enmedio Formation, different from
other Laramidian tyrannosaurids (e.g. Gorgosaurus, Daspletosaurus,
Tyrannosaurus; see Appendix 1); and particularly, it could provide
more evidence for a higher taxonomic diversity of tyrannosaurid
theropods within the Cabullona Basin, adding more information on
the diversication of the Tyrannosauridae family in one of the most
southern Laramidian regions during the Late Cretaceous.
6. Conclusions
Isolated teeth represent the most abundant theropod fossils that
could be found in worldwide Mesozoic rocks. Particularly, they are
an important source of information concerning the dental
morphologic variation within tyrannosaurids. The aim of the pre-
sent work was to describe and analyze three unknown theropod
teeth found isolated and surface collected in the Corral de Enmedio
Formation (Cabullona Group, Upper Cretaceous) in northeastern
Sonora, Mexico. On the basis of the combination of several impor-
tant morphological characteristics along with statistic and phylo-
genetic analyses, we refer ERNO 8027, ERNO 8581 and ERNO 8582
as members of the Tyrannosauridae family, probably representing a
new indeterminate taxon.
Until now the taxonomic identity of the ERNO specimens cannot
be asserted beyond the Tyrannosauridae family level, due to the
lack of more complete skeletal elements. So, the need for extensive
sampling is required in order to recover more complete tyranno-
saurid remains, which would provide a better understanding of the
taxonomic diversity and paleogeographic distribution of these di-
nosaurs in the Cabullona Basin, adding new information to the
diversity of Laurasian tyrannosaurs in the Late Cretaceous of
western North America.
Acknowledgements
The authors thank the editor, Dr. Steve Brusatte and an anony-
mous reviewer for their detailed and thoughtful comments that
greatly improved the preparation of this manuscript. Also, we
thank Dr. Maria del Socorro Lozano Garca from the Instituto de
Geologa, UNAM for her support.
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Appendix A. Supplementary data
Supplementary data related to this article can be found at http://dx.doi.org/10.
1016/j.cretres.2017.03.015