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Tyrannosaurus Mcraensis
Tyrannosaurus Mcraensis
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Tyrannosaurids were the dominant predators in North America and Asia during the Late Cretaceous1–3. Evolv-
ing from small-bodied ancestors in the mid-Cretaceous, tyrannosaurids became apex predators in the latest
Cretaceous, and finally saw the appearance of Tyrannosaurus rex4,5. T. rex, characterized by a robust skeleton
and powerful, bone-crushing jaws6, was the dominant carnivore in the late Maastrichtian of western North
America3,7. Growing to 12 m long and ~ 10 tons in w eight8, T. rex was the largest terrestrial predator of its time,
and perhaps of all time.
The origins of the Tyrannosaurus lineage, Tyrannosaurini, are unknown. Tyrannosaurus rex appeared sud-
aastrichtian3,7. No close relatives have been reported from North America prior to this time.
denly in the latest M
Instead, the closest relatives of T. rex come from Mongolia. To explain this pattern, one hypothesis suggests
tyrannosaurins dispersed into Asia via Beringia, followed by back-dispersal of Tyrannosaurini into Laramidia
in the late Maastrichtian9–11. This hypothesis is based on the existence of stratigraphically earlier Asian taxa,
Tarbosaurus bataar12 and Zhuchengtyrannus magnus13, which are close relatives of T. rex9,10,14,15. Alternatively,
Tyrannosaurus has been hypothesized to represent an endemic, North American lineage14. Here, we describe
NMMNH P-3698, a giant tyrannosaurid from the late Campanian—early Maastrichtian of New Mexico (Fig. 1).
Although originally identified as T. rex16–19, this assignment has been questioned20. Restudy of the specimen,
including newly recovered skeletal elements (Figs. 2, 3) confirm NMMNH P-3698 represents a distinct taxon,
while recently published radiometric dates show that it predates T. rex by up to 7 million years. Phylogenetic
1
New Mexico Museum of Natural History and Science, 1801 Mountain Road N.W., Albuquerque, NM 87104,
USA. 2Department of Geology and Geophysics, University of Utah, Salt Lake City, UT, USA. 3Natural History
Museum of Utah, University of Utah, Salt Lake City, UT, USA. 4Department of Biological Sciences, The George
Washington University, 2023 G St. NW, Washington, DC 20052, USA. 5Department of Environmental Science
and Sustainability, Harrisburg University, 326 Market Street, Harrisburg, PA 17101, USA. 6Pennsylvania State
University, Lehigh Valley, Center Valley, PA 18034, USA. 7Department of Biological Sciences, University of Alberta,
Edmonton, AB T6G 2E9, Canada. 8Department of Biology and Biochemistry, University of Bath, Bath BA2 7AY,
UK. *email: [email protected]
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Figure 1. Locality and stratigraphy of Tyrannosaurus mcraeensis gen. et sp. nov., NMMNH P-3698. (A), type
locality in Sierra County, New Mexico; (B), stratigraphy of fossil and the Hall Lake Formation (C), recovered
skull elements. Scale = 10 cm. Map by Ron Blakey.
Figure 2. Cranial elements of Tyrannosaurus mcraeensis gen. et sp. nov. (NMMNH P-3698). Right postorbital
in (A), lateral view; (B), medial view; (C), dorsal view. Right squamosal in (D), lateral view; (E), medial view;
(F), ventral view. cor cornual boss, fos ventral fossa, fr frontal articulation, ltf lateral temporal fenestra, po
postorbital articulation, sp suborbital process, stf supratemporal fossa, qc quadrate cotyle, qj quadratojugal
process. Scale = 10 cm.
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Figure 3. Mandibular elements of Tyrannosaurus mcraeensis gen. et sp. nov. (NMMNH P-3698). Left dentary
in (A), medial view; (B), lateral view; (C), dorsal view; (D), right splenial, medial view; (E), right angular, medial
view; (F), right prearticular, medial view. ang angular contact, de shelf for dentary, mec Meckelian canal, pre
prearticular facet, a1, a5, a15 alveoli 1, 5, and 15, sp splenial, sym symphysis. Scale = 20 cm.
analysis recovers NMMNH P-3698 as the closest known relative of T. rex, suggesting giant Tyrannosaurini
evolved in southern Laramidia.
Systematic paleontology
Dinosauria—Owen, 1842.
Theropoda—Marsh, 1881.
Tetanurae—Gauthier, 1986.
Coelurosauria—von Huene, 1914.
Tyrannosauridae—Osborn, 1905.
Tyrannosaurinae—Currie, 2003.
Tyrannosaurini—Olshevsky, 1995.
Tyrannosaurus mcraeensis sp. nov.
LSID:
urn:lsid:zoobank.org:pub:F1658ACA-60DB-442E-AA04-015C050205BD.
(Tyrannosaurini is here defined as the last common ancestor of Tarbosaurus baatar and Tyrannosaurus rex
and all its descendants).
Etymology
The species name, mcraeensis, refers to the McRae Group of western New Mexico.
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Diagnosis
Large tyrannosaurin distinguished from Tyrannosaurus rex (Fig. 4; SI) by the following characters (*autapo-
morphies): postorbital with low, posteriorly positioned cornual process; postorbital with anteriorly project-
ing prefrontal/frontal articular surfaces; squamosal with ventrally projecting quadratojugal process; squamosal
with a concave medial margin; strong ridge bounding the anterior margin of the squamosal ventral pneumatic
fossa; dentary very shallow posteriorly and with a convex posteroventral margin*; splenial with anteriorly posi-
tioned apex*; splenial with shelf-like dentary overlap*, splenial with deep, posteriorly directed angular process;
prearticular weakly bowed*; small ventral prearticular-angular contact; articular T-shaped in dorsal/ventral
view; retroarticular process deep and quadrangular in posterior view.
Holotype
NMMNH P-3698, partial skull including right postorbital and squamosal (Fig. 2), left palatine, fragment of
maxilla, and lower jaws (Fig. 3) including left dentary, right splenial, prearticular, angular and articular, isolated
teeth and associated chevrons.
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Tarbosaurus12 and Tyrannosaurus rex (Fig. 4E). The angular process is deep, as in Tarbosaurus; it is shallow in
T. rex5. The anteroventral process, beneath the mylohyoid foramen, has a shelf overlapping the dentary, in T. rex
it abuts the dentary.
The anterior process of the angular (Fig. 3E) is longer and narrower than in Tyrannosaurus rex (Fig. 4F)
and more similar to Lythronax and Tarbosaurus. It has a narrow ventral prearticular contact; T. rex has a broad
anteroventral flange here. The ventral margin forms a sharp angle between the anterior and posterior processes.
The prearticular (Fig. 3F) is gently curved, versus strongly bent in Tyrannosaurus rex5 (Fig. 4D) and other
tyrannosaurids12,24. The articular is T-shaped in dorsal view, versus more triangular in T. rex. In posterior view,
the retroarticular process is deep and subrectangular; that of T. rex is wider, and semicircular (SI).
Teeth (Fig. 3; SI) resemble Tyrannosaurus rex5 in being large, robust, and labiolingually expanded. Tooth
crowns have massive apices, as in Tarbosaurus12; T. rex teeth have more pointed, spike-like apices5. The apex of
the sixth dentary tooth is worn; T. rex tooth wear is often heavy as a result of biting bone6,29,30. The labiolingual
width of anterior teeth approaches their mesiodistal diameter, as in T. rex; posterior teeth are robust, but more
laterally compressed. Both carinae bear serrations.
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Figure 5. Size, relationships and biogeography of Tyrannosaurus mcraeensis. (A), relative sizes of
Tyrannosaurus mcraeensis (NMMNH P-3698) and Tyrannosaurus rex (FMNH PR 2081 and CM 9380) ( after31);
(B), evolutionary tree based on Bayesian tip-dated phylogeny and biogeographic analysis (see SI for full results).
Phylogenetic analysis
Phylogenetic analyses consistently recover Tyrannosaurus mcraeensis as sister to T. rex (Fig. 5) using either a
parsimony-based or Bayesian tip-dated analysis (SI). The two share a reduced tooth count, deep dentary sym-
physis, posteriorly shallow lingual bar, and large size. T. mcraeensis is furthermore united with the Tarbosau-
rus + Tyrannosaurus clade by a postorbital with a broad ventral ramus, strongly convex posterior margin, and
large suborbital flange, by the strongly concave dentary alveolar margin, and by the laterally bowed mandible.
Discussion
Systematics
NMMNH P-3698 was referred to Tyrannosaurus rex17,19,21 based on size and overall resemblance to T. rex, but
this assignment has been questioned20. Newly collected material shows that T. mcraeensis differs from T. rex in
the shape of the postorbital, squamosal, dentary, prearticular, angular, and articular. The characters that diagnose
T. mcraeensis and differentiate it from T. rex are relatively subtle characters relating to the shape and articulation
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of the skull bones, but because T. rex is known from multiple individuals, it is possible to show that T. mcraeensis
lies outside of the range of individual variation seen in T. rex (Fig. 6).
Tyrannosaurus rex exhibits a high degree of variation31–33 and specimens assigned to the genus may or may not
represent multiple species31,33,34. However in each of the diagnostic characters identified here, NMMNH P-3698
is unlike any other specimen referred to T. rex; furthermore each bone has at least one diagnostic character, it
is therefore an outlier from all other specimens referred to T. rex, in every element. Neither can the differences
between T. rex and T. mcraeensis be explained as ontogenetic, given that the animal matched T. rex in size. It
is worth noting that these differences are subtle, but the differences between species are often relatively subtle.
Characters diagnosing different species of Daspletosaurus10,15 tend to be relatively subtle, as are those diagnosing
different species of Alioramus35.
Figure 6. Variation in the postorbitals (A–F), dentaries (G–K) and splenials (M–Q) of Tyrannosaurus
mcraeensis (A, G, M) and Tyrannosaurus rex (B–F, H–L, N–Q). See SI for specimen numbers. Scale bars = 10
cm.
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Morphology also suggests Tyrannosaurus mcraeensis is older than T. rex, since it consistently lacks many
derived characters characterizing T. rex. Tip-dated phylogenetic analysis recovers NMMNH P-3698 at 69 Ma,
slightly younger than implied by the radiometric dates, but still much older than any known T. rex (Fig. 7).
Radiometric dates also point towards a latest Campanian or early Maastrichtian age. U/Pb d ating22 gives an
age of 73.2 ± 0.7 Ma for a tuff 9 m above the base of the Hall Lake Formation, and a series of Late Campanian
ages for the upper 70 m of the immediately underlying Jose Creek Formation, at 75.2 ± 1.3 Ma, 74.6 ± 0.6 Ma,
74.9 ± 0.7 Ma, and 75.0 ± 1.1 Ma. Although no radiometric dates are found above NMMNH P-3698, a titanosaur
is found 108 m above the site, and can be no younger than 66 Ma, i.e. the K-Pg boundary. No hiatus, coal, or
other evidence of the K-Pg boundary is visible in the exposed section, suggesting around 150 m of Cretaceous
rock overlay the site. Depending on the estimated rate of sedimentation, Tyrannosaurus mcraeensis may there-
fore have lived between 72.7 and 70.9 Ma (SI), i.e., latest Campanian or earliest Maastrichtian, 5–7 million years
before Tyrannosaurus rex.
Overall, three independent lines of evidence—(i) the dinosaur fauna, (ii) the morphology of NMMNH
P-3698 itself, and (iii) the age constraints provided by underlying radiometric dates and overlying dinosaur
bones suggest an age somewhat older than the latest Maastrichtian, and that the animal predates Tyrannosaurus
rex. However, additional radiometric dates, palynostratigraphy, or vertebrate fossils are needed to provide more
precise constraints on the age of NMMNH P-3698.
Tyrannosaurus mcraeensis shows several derived characters T. rex does not: an upturned dentary posteroven-
tral margin, a weakly curved prearticular, and an anterior splenial apex (Fig. 4). These characters imply that T.
mcraeensis was a side-branch in tyrannosaurin evolution, and would not have been directly ancestral to Tyran-
nosaurus rex. If so, then at least two giant tyrannosaurids existed concurrently in North America, with another
species ultimately giving rise to Tyrannosaurus rex.
Figure 7. Stratigraphy and radiometric dates of the basal Hall Lake Formation and top of the underlying Jose
Creek Formation.
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the Campanian, one lineage dispersed to Asia, giving rise to Tarbosaurus and Zhuchengtyrannus; another gave
rise to Tyrannosaurus, eventually moving north to displace albertosaurines in the Late Maastrichtian.
In an alternative character-matrix51, Tyrannosaurus mcraeensis again emerges as sister to T. rex (SI). In
this topology, Tarbosaurus and Zhuchengtyrannus are successive outgroups to the T. mcraeensis–T. rex clade.
This topology is ambiguous about the origin of Tyrannosaurini, consistent with either two dispersals of Tyran-
nosaurinae into Asia, or dispersal into Asia, followed by back-dispersal into North America. The difference is
due to the poorly known Zhuchengtyrannus acting as a wildcard taxon, clustering either with Tarbosaurus14 or
as sister to the T. rex—Tarbosaurus clade9,10,15. The out-of-Asia scenario is dependent on the placement of this
uncertain taxon. Resolving these ambiguities requires a better understanding of Asian tyrannosaurs. Similarly,
placement of Daspletosaurus spp. along the line leading to T yrannosaurini15 would tend to support a northern
origin of the Tyrannosaurini. Further study of southern tyrannosaurs such as Bistahieversor and Lythronax can
potentially resolve these issues, along with revisions of tyrannosaur phylogeny, but we argue that the appearance
of T. rex-like dinosaurs in the Javelina Formation, which appear to be slightly older than T. rex itself, better fits
the southern s cenario9,50.
Tyrannosaurus mcraeensis shows that Tyrannosaurini achieved giant size near the end of the Campanian.
It furthermore suggests that they did so in southern Laramidia (Fig. 5) but were initially restricted to southern
Laramidia, and later dispersed n orth7. Chasmosaurinae show similar patterns, with giant Triceratopsini evolving
in southern Laramidia in the Campanian, then moving into the northern Great Plains in the Maastrichtian46.
Several other giant dinosaurs evolved in southern Laramidia, including large kritosaurin hadrosaurs52, the giant
lambeosaurine Magnapaulia laticaudus53, and the titanosaurian Alamosaurus sanjuanensis54, but did not migrate
north. Meanwhile, a tyrannosaurid from northernmost Laramidia, Nanuqsaurus hoglandi27, was smaller than
tyrannosaurines from farther south. This, and the presence of giant ceratopsians and hadrosaurs in the Hall Lake
Formation (SI) implies geographic patterns of body size evolution: large dinosaurs evolved in southern Laramidia.
Sea level changes have been hypothesized to drive tyrannosaurid d iversification14, and among mammals,
land area was found to be associated with body mass of top carnivores and h erbivores55. However, the patterns
raised here suggest that other processes are at work in driving tyrannosaurid size evolution. The evolution of
Tyrannosaurus-sized tyrannosaurs appears to precede end-Maastrichtian marine regression by several million
years; furthermore the giant tyrannosaurs appear to be initially restricted to a limted area of the relatively small
Laramidian landmass. Gigantism therefore appears not to be driven by increases in land area, instead giant
theropods evolved despite having relatively small geographic ranges.
Giant tyrannosaurins likely evolved to prey on the giant herbivores found in the south, although the reason
for the evolution of large herbivorous dinosaurs—possibly including latitudinal variation in mean annual tem-
perature, primary productivity, or seasonality—remains unknown. These patterns also suggest caution in relying
on local dinosaur diversity patterns to study global dinosaur diversity changes preceding the K-Pg extinction.
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Data availability
All data is available in the manuscript or Supporting Information.
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(2020).
Acknowledgements
The dentary, palatine, and prearticular of NMMNH P-3698 were collected by D. Gillette; additional remains
were collected by T. Williamson. Photos for the SI courtesy T. Miyashita.
Author contributions
S.G.D., N.R.L., M.A.L. designed the project. S.G.D., S.G.L., M.A.L., R.A.P., S.E.J., D.E.M., A.R.F., P.J.C. conducted
research and wrote the paper. S.G.D., M.A.L., N.R.L. and R.A.P. conducted the phylogenetic analysis.
Competing interests
The authors declare no competing interests.
Additional information
Supplementary Information The online version contains supplementary material available at https://doi.org/
10.1038/s41598-023-47011-0.
Correspondence and requests for materials should be addressed to N.R.L.
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