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OPEN A giant tyrannosaur

from the Campanian–Maastrichtian
of southern North America
and the evolution of tyrannosaurid
gigantism
Sebastian G. Dalman 1, Mark A. Loewen 2,3, R. Alexander Pyron 4, Steven E. Jasinski 5,
D. Edward Malinzak 6, Spencer G. Lucas 1, Anthony R. Fiorillo 1, Philip J. Currie 7 &
Nicholas R. Longrich 8*

Tyrannosaurid dinosaurs dominated as predators in the Late Cretaceous of Laurasia, culminating in

the evolution of the giant Tyrannosaurus rex, both the last and largest tyrannosaurid. Where and when
Tyrannosaurini (T. rex and kin) originated remains unclear. Competing hypotheses place tyrannosaurin
origins in Asia, or western North America (Laramidia). We report a new tyrannosaurin, Tyrannosaurus
mcraeensis, from the Campanian–Maastrichtian Hall Lake Formation of New Mexico, based on a fossil
previously referred to T. rex. T. mcraeensis predates T. rex by ~ 6–7 million years, yet rivaled it in size.
Phylogenetic analysis recovers T. mcraeensis as sister to T. rex and suggests Tyrannosaurini originated
in southern Laramidia. Evolution of giant tyrannosaurs in southern North America, alongside giant
ceratopsians, hadrosaurs, and titanosaurs suggests large-bodied dinosaurs evolved at low latitudes in
North America.

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.

Horizon and locality

Uppermost Campanian or lower Maastrichtian of the Hall Lake Formation, McRae Group, NMMNH locality
343, near Kettle Top Butte, Sierra County, New ­Mexico21 (Fig. 1A). The site lies 43 m above the base of the Hall
Lake Formation. A tuff 33 m below the tyrannosaur site has a U/Pb age of 73.2 ± 0.7 ­Ma22 (Fig. 1B).

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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.

Description and comparisons

The postorbital of Tyrannosaurus mcraeensis (Fig. 2A–C) is typical of t­ yrannosaurines5,12 in bearing a massive
cornual boss projecting above the postorbital dorsal margin. The cornual boss is C-shaped as in Tarbosaurus12
but lacks the strong anteroventral expansion or undercut margin seen in Tarbosaurus and T. rex. The cornual
boss lacks the prominent, anteriorly positioned apex above the orbit seen in T. rex, instead being more posteriorly
positioned. The postorbital’s dorsal margin is arched, as in T. rex5. In dorsal view the orbital margin and the body
of the postorbital form a wide angle, indicating a posteriorly expanded skull as in Lythronax14, Tarbosaurus12, and
Tyrannosaurus rex5. The frontal/prefrontal suture projects forward, whereas it is downturned in T. rex (Fig. 4A).
The jugal process is anteroposteriorly expanded, with a convex jugal contact, as in T. rex5 and T. bataar12. A large
suborbital process projects forward to constrict the orbit as in Bistahieversor23, Teratophoneus14, and especially
T. rex5 and Tarbosaurus12.
The squamosal (Fig. 2D–F) recalls Tyrannosaurus rex5 in being elongate in dorsal/ventral view. It is shorter
in Tarbosaurus12, and especially ­Albertosaurinae24. It differs from the T. rex holotype, where the squamosal is
straight (Fig. 4B) in having a strongly downturned end in lateral view; however some specimens of T. rex show
this feature (SI1). The quadratojugal process is downturned relative to T. rex5 or Tarbosaurus12 and strongly
curved. Ventrally, the squamosal bears a deep pneumatic recess. The recess’ anterior margin is defined by a
transverse bar as in Tarbosaurus; this bar is reduced to a low ridge in T. rex12.
The palatine (SI5) is broad, similar to Tarbosaurus bataar12 but not to the degree seen in Tyrannosaurus rex;
the palatine of Albertosaurinae is n­ arrower24. This wide palatine contributes to the formation of a broad rostrum
as in T. rex and Tarbosaurus.
The dentary (Fig. 3A–C) measures 645 mm from the tip back to the level of the coronoid process; the bone
measures 894 mm along the long axis as preserved, however the posteroventral end is broken; it may have
measured around 900 mm when complete. By comparison, corresponding measurements for the holotype of
Tyrannosaurus rex are 589 and 855 mm; RSM P2523.8, one of if not the largest known T. rex ­specimens25, meas-
ures ~ 650 from the tip of the dentary to the coronoid process. Tyrannosaurus mcraeensis therefore overlaps T.
rex in size (Fig. 5A), although the holotype is smaller than the largest known T. rex individuals. The dentary
has 13 alveoli, a low tooth count uniquely shared with T. rex5 which typically has 13 dentary alveoli, and rarely
14 or ­1526 alveoli. Zhuchengtyrannus has 15 teeth, Tarbosaurus has 14–1512,13; Daspletosaurus horneri10 and D.
torosus have 17. The first alveolus is smaller than the second, as in other tyrannosaurins. The symphysis is deep,
being about 125% the depth of the dentary at mid-length, as in T. rex. The symphysis’ anteroventral margin
rises up steeply, creating a squared-off chin, again as in T. rex. The dentary’s occlusal margin rises up steeply at
the back forming a strongly concave dorsal margin, as in other t­ yrannosaurines5,12–14. Unusually, the dentary’s
posterior end is shallow and upturned; it is deep and typically has a downturned ventral margin in T. rex and
other ­tyrannosaurids5,14,24. This condition is unique to T. mcraeensis but approached in Tarbosaurus5,12 and
Zhuchengtyrannus13. The angular process projects posteriorly, as in Tarbosaurus and Zhuchengtyrannus13 but
unlike T. rex and other tyrannosaurids, where it projects ­posteroventrally5,14,24.
In dorsal view the tip of the dentary bows outward so that the symphysis forms a broad “U”. Similar bowing
is present in Tyrannosaurus rex5,14 and Tarbosaurus, associated with transverse expansion of the rostrum. The
dentary tip is straight in Daspletosaurus, Lythronax14, Nanuqsaurus27 and more basal t­ axa28. The dentary and
toothrow curve outwards posteriorly, again suggesting transverse expansion of the skull’s postorbital region, as
in Lythronax14, Tarbosaurus, and T. rex5,12.
Medially, interdental plates are large and rectangular to triangular, as in other ­tyrannosaurins5,13; they are
smaller in A­ lbertosaurinae24 and Daspletosaurus24. The lingual bar covers the first two alveoli. It is deep anteriorly
then narrows posteriorly to half its anterior depth, a condition otherwise seen only in Tyrannosaurus rex5. The
symphysis is rugose and covered with bumps and grooves, as in other tyrannosaurines. It extends back to the
third alveolus, as in Tarbosaurus12. In Zhuchengtyrannus13 and some individuals of T. rex5 it ends under the fourth
alveolus. The symphysis lies above the dentary ventral margin, as in Tarbosaurus12 and Zhuchengtyrannus13; in
T. rex the symphysis projects below the dentary’s ventral m ­ argin25 in some, but not all individuals. The bone’s
lateral surface bears pits and scars suggestive of intraspecific combat.
The triangular splenial (Fig. 3D) resembles other ­tyrannosaurids5,12,24. The tall dorsal process resembles T.
rex12 and Tarbosaurus12; Daspletosaurus24 has a low dorsal process. The apex is triangular, like Tarbosaurus; in
T. rex the apex is usually quadrangular (Fig. 4e). The apex is anteriorly displaced compared to other tyranno-
saurids. The ventral margin is straight, and the angular process projects posteriorly, versus posteroventrally in

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Figure 4.  Tyrannosaurus mcraeensis compared with Tyrannosaurus rex. Postorbital of Tyrannosaurus

mcraeensis (A1) and Tyrannosaurus rex (A2); Squamosal of (B1) Tyrannosaurus mcraeensis and (B2)
Tyrannosaurus mcraeensis; Postorbital of (C1) Tyrannosaurus mcraeensis and (C2) Tyrannosaurus rex;
prearticular of (D1) Tyrannosaurus mcraeensis and (D2) Tyrannosaurus rex; splenial of (E1) Tyrannosaurus
mcraeensis and (E2) Tyrannosaurus rex; angular of (F1), Tyrannosaurus mcraeensis and (F2), Tyrannosaurus rex.
Diagnostic characters: 1, low cornual process behind orbit; 2, anteriorly projected frontal/prefrontal articulation;
3, ventrally projected quadratojugal process; 4, upturned posteroventral margin of dentary; 5, weakly bowed
prearticular; 6, anteriorly positioned splenial apex; 7, deep and posteriorly projected angular process; 8, shelf-
like articulation overhanging dentary; 9, small splenial facet. Scale bars = 10 cm (A, B, D–F), 20 cm (C).

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.

Age of NMMNH P‑3698

Referral of NMMNH P-3698 to Tyrannosaurus rex was also based on the assumption that the Hall Lake Forma-
tion is late Maastrichtian aged; but no geological or paleontological evidence supports this assignment. In fact,
one of the main arguments for a late Maastrichtian age was the incorrect assumption that Hall Lake dinosaurs
could be referred to the late Maastrichtian taxa Torosaurus and Tyrannosaurus rex19. The ceratopsid previously
referred to Torosaurus is now identified as a distinct genus, Sierraceratops16; as shown here, NMMNH P-3698
is distinct from T. rex.
The dinosaur fauna suggests a latest Campanian or Maastrichtian age. Sierraceratops is similar to species
known from the latest Campanian, including Coahuilaceratops magnacuerna36, and to Bravoceratops polyphemus
from the Javelina Formation of ­Texas37, which is probably either latest Campanian or early Maastrichtian. The
existence of a titanosaurian sauropod in the Hall Lake Formation has been cited evidence of a late Maastrichtian
or ‘Lancian’ a­ ge19,38. Titanosaurs appear suddenly in the latest Cretaceous of the American Southwest, suggest-
ing a distinct biogeographic event resulting from their sudden dispersal into North ­America39; sauropods are
therefore a useful biostratigraphic constraint. However, titanosaurs range from just beneath the K-Pg ­boundary40
to at least 69 ± 0.9 Ma, i.e. mid-Maastrichtian40. Sauropods are unknown from the De-Na-Zin Member of the
Kirtland Formation of New M ­ exico39, the top of which is dated to 73.5 ­Ma41, or the Cerro Del Pueblo Formation
of Mexico, which is latest Campanian, ~ 73.5–73 ­Ma42. Sauropods therefore appear to immigrate into North
America between 73 and 69 Ma. The existence of titanosaurs in the assemblage, therefore, does not reject a latest
Campanian or early Maastrichtian age, and given that the sauropod appears higher in section, NMMNH P-3698
could predate the arrival of titanosaurians in North America. Finally, a large hadrosaur femur resembles that of
the Campanian-Maastrichtian42 Edmontosaurus43 and a large edmontosaurin from the latest Campanian Cerro
Del Pueblo F ­ ormation44 in terms of its large size and strongly inturned proximal femoral shaft. The presence of
an edmontosaurin is consistent with an age range of latest Campanian to latest Maastrichtian. The dinosaur fauna
as a whole is consistent with and suggests either a Campanian to Maastrichtian age. The ceratopsid Sierraceratops
also tends to suggest a latest Campanian to early Maastrichtian age. Strikingly, no diagnostic late Maastrichtian
dinosaur species are known.

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.

Biogeography and the evolution of body size

Tyrannosaurus mcraeensis belonged to an endemic southern dinosaur community. The Hall Lake fauna (Fig. 8)
included T. mcraeensis, the giant chasmosaur Sierraceratops16, the titanosaurian cf. Alamosaurus, and a giant
­hadrosaurid19. This fauna shows no overlap with penecontemporaneous faunas in the latest Campanian/early
Maastrichtian of Canada. This northern fauna included the albertosaurine tyrannosaurid Albertosaurus, the
centrosaurine Pachyrhinosaurus, the lambeosaur Hypacrosaurus, the hadrosaurs Edmontosaurus and Saurolophus,
and the chasmosaurine Anchiceratops; sauropods are ­absent45. Discovery of T. mcraeensis, therefore, corroborates
hypotheses that Laramidian faunas were characterized by high e­ ndemicity16,46–48, with the Southwest supporting
distinct dinosaur species and clades (Fig. 8).
The presence of Tyrannosaurus mcraeensis in New Mexico and its phylogenetic position suggest that
­Tyrannosaurini14 originated as part of this southern fauna. A southern origin of Tyrannosaurini is further
supported by the presence of cf. Tyrannosaurus in the Javelina Formation of ­Texas49,50 and T. rex in the latest
Maastrichtian North Horn of U ­ tah7. Following the appearance of tyrannosaurins in southern Laramidia during

Figure 7.  Stratigraphy and radiometric dates of the basal Hall Lake Formation and top of the underlying Jose
Creek Formation.

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Figure 8.  Dinosaurs of the Campanian–Maastrichtian Hall Lake Formation. Scale = 2 m.

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.

Materials and methods

We added Tyrannosaurus mcraeensis to an existing character-taxon ­matrix14, adding 10 new characters as well as
additional taxa (SI2). Phylogenetic analysis was implemented using cladistic methods and total-evidence estima-
tion of phylogeny and divergence times using the fossilized clock model and birth–death priors under diversified
sampling as described by recent ­authors56, with tip-dated estimates of fossil ages from all sampled taxa chosen
using a “diversity” strategy to span internal branches. We optimized biogeographic history across the summary
tree by classifying species into six areas: Europe, South America, Asia, Appalachia, Northern Laramidia, and
Southern Laramidia, the latter two separated by present-day Utah and Colorado as the demarcating boundary.
We tested a standard suite of biogeographic models in R ­ ASP57, selecting the DEC + J model as the best fit, and
present the most-likely estimates for ancestral range (Fig. 5B).

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Data availability
All data is available in the manuscript or Supporting Information.

Received: 8 June 2023; Accepted: 7 November 2023

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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
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