Gregory S.

Paul and colleagues asserted in a 2022 study that Tyrannosaurus rex, as it has been
conventionally regarded, actually represented three species: the original species Tyrannosaurus
rex and two additional species, T. imperator (meaning "tyrant lizard emperor" and "tyrant lizard
queen") and T. regina. The Sue specimen serves as the holotype for the former (T. imperator),
whereas Wankel rex serves as the holotype for the latter (T. regina). The discovery of a relatively
high degree of diversity in the proportions and robusticity of the femur (and other skeletal parts)
between recorded T. rex specimens, more so than that reported in other theropods regarded as
one species, served as the primary basis for the separation into many species. In addition to the
number of short, slender incisiform teeth in the dentary, based on tooth sockets, differences in
overall body proportions indicating robust and gracile morphotypes were also employed as a line
of evidence. The paper's T. rex was distinguished by robust anatomy, a moderate femur length to
circumference ratio, and the presence of a single slender incisiform dentary tooth. T. imperator
was also considered robust, but it had two of the slender teeth and a high femur ratio. T. regina
was a gracile form with a low femur ratio. It was noted that, stratigraphically, proportional and
robustness variation became more extreme at higher sample altitudes. This was explained as the
speciation of more than one taxon, T. rex and T. regina, from a single older population, T.
imperator.[69]

When contacted by various media outlets for comment, a number of other prominent
paleontologists, including Stephen Brusatte, Thomas Carr, Thomas Holtz, David Hone, Jingmai
O'Connor, and Lindsay Zanno, disputed the study or voiced doubt about its findings.[70][71][72]
They then had their criticism written up in a technical paper.[73] Although Holtz and Zanno
agreed that it was possible for there to be more than one species of Tyrannosaurus, they believed
the new study lacked adequate evidence to back up the species it suggested. Even if
Tyrannosaurus imperator was a different species from Tyrannosaurus rex, according to Holtz, it
might actually be the same species as Nanotyrannus lancensis, in which case it would need to be
referred to as Tyrannosaurus lancensis. A curator at the Field Museum, where the T. imperator
holotype Sue is on display, O'Connor, believed that the new species had insufficient evidence to
warrant changing the exhibit placards. The distinguishing characteristics proposed to separate the
species were seen by Brusatte, Carr, and O'Connor as reflecting natural variation within a
species. Carr and O'Connor both voiced dissatisfaction with the study's failure to identify which
of the hypothesized species a number of well-preserved specimens belonged to. The study's
initial co-author, another paleontologist named Philip J. Currie, withdrew because he did not
want to be engaged in the process of identifying the new species.[70]

Critics' arguments were dismissed by Paul, who insisted that they were hesitant to accept the
possibility that Tyrannosaurus might have represented multiple species.[74] In a later work that
is still being published, Paul upheld the finding that there are three different species of
Tyrannosaurus. He emphasized that while the original study also compared the robustness of
other bones (the maxilla, dentary, humerus, ilium, and metatarsals), the criticism of the study
naming T. imperator and T. regina only focused on two of the features used to distinguish the
two new species (the number of small incisiform teeth and femur robustness). Paul further
asserted that three distinct species of Tyrannosaurus can be identified based on the design of the
knob-like lumps (referred to as "postorbital bosses") located behind the eyes. Paul further argued
that previous studies that concluded there was only one species of Tyrannosaurus (T. rex) had
simply assumed that all Tyrannosaurus skeletons belonged to a single species. He also claimed
that many new dinosaur species had been named based on less variation than he and his
colleagues did when they proposed the names T. imperator and T. regina.[75]