THE ANATOMICAL RECORD 294:119131 (2011)

The Tail of Tyrannosaurus: Reassessing

the Size and Locomotive Importance of
the M. caudofemoralis in Non-Avian
Theropods
W. SCOTT PERSONS IV* AND PHILIP J. CURRIE
Department of Biological Sciences, University of Alberta, Edmonton, Alberta, Canada

ABSTRACT
Unlike extant birds and mammals, most non-avian theropods had
large muscular tails, with muscle arrangements similar to those of
modern reptiles. Examination of ornithomimid and tyrannosaurid tails
revealed sequential diagonal scarring on the lateral faces of four or more
hemal spines that consistently correlates with the zone of the tail just an-
terior to the disappearance of the vertebral transverse processes. This se-
quential scarring is interpreted as the tapering boundary between the
insertions of the M. caudofemoralis and the M. ilioischiocaudalis. Digital
muscle reconstructions based on measurements of fossil specimens and
dissections of modern reptiles showed that the M. caudofemoralis of
many non-avian theropods was exceptionally large. These high caudofe-
moral mass estimates are consistent with the elevation of the transverse
processes of the caudal vertebra above the centrum, which creates an
enlarged hypaxial region. Dorsally elevated transverse processes are
characteristic of even primitive theropods and suggest that a large M.
caudofemoralis is a basal characteristic of the group. In the genus Tyran-
nosaurus, the mass of the M. caudofemoralis was further increased by
dorsoventrally lengthening the hemal arches. The expanded M. caudofe-
moralis of Tyrannosaurus may have evolved as compensation for the ani-
mals immense size. Because the M. caudofemoralis is the primary hind
limb retractor, large M. caudofemoralis masses and the resulting con-
tractile force and torque estimates presented here indicate a sizable
investment in locomotive muscle among theropods with a range of body
sizes and give new evidence in favor of greater athleticism, in terms of
overall cursoriality, balance, and turning agility. Anat Rec, 294:119131,
2011. V C 2010 Wiley-Liss, Inc.

Key words: dinosaurs; Tyrannosaurus; locomotion; tail

musculature; biomechanics

Abbreviations used: AMNH American Museum of Natural
History, New York, New York, USA; BHI Black Hills
Institute of Geological Research, Hill City, South Dakota, USA;
FMNH Field Museum of Natural History, Chicago, Illinois,
USA; NMNH Smithsonian Institution National Museum of
Natural History, Washington, District of Colombia, USA; TCM
The Childrens Museum of Indianapolis; TMP Royal Tyrrell
Museum of Palaeontology, Drumheller, Alberta, Canada
*Correspondence to: W. Scott Persons IV, Department of Biological
Sciences, CW 405 Biological Sciences Bldg., University of Alberta,
Edmonton, Alberta, Canada T6G 2E9. E-mail: persons@ualberta.ca
Received 11 January 2010; Accepted 9 August 2010
DOI 10.1002/ar.21290
Published online 12 November 2010 in Wiley Online Library
(wileyonlinelibrary.com).

V
C 2010 WILEY-LISS, INC.
120 PERSONS IV AND CURRIE
The associations between muscle mass, vertebrae mor-
phology, and function have been well studied in the tails
of extant mammals, particularly procyonids and prima-
tes, and tail biomechanics are better understood among
mammals than any other group of terrestrial vertebrates
(Dor, 1937; German, 1982; Lemelin, 1995; Organ et al.,
2009). However, in terms of mass and volume, most fully
terrestrial mammals have unimpressive tails. Large ter-
restrial mammals, in particular, tend to have minimalist
y-swatter tails, and our own species, with nothing but
a vestigial stub, is at the furthest extreme. As a result,
when considering the relatively large tails of most non-
avian dinosaurs, there has been a misleading tendency
to regard tails as predominantly dead weight or, at best,
as either defensive lashing weapons or (in bipedal taxa)
as mere counterbalances for the crania. Modern reptiles
represent the best modern analogs for the tails of most
dinosaurs and demonstrate that large tails may serve a
variety of consequential functions, including femoral re-
traction during the locomotive power stroke via the
M. caudofemoralis.
The M. caudofemoralis is a tail muscle that inserts
directly onto the fourth-trochanter of the femur, and
acts as a femoral retractor. Extant mammals lack the M.
caudofemoralis; instead, the gluteal muscles ll the role
of primary limb retractors. Some mammals do have a
muscle that is commonly termed the M. caudofemora-
lis, but this muscle is not homologous to the M. caudo-
femoralis of saurians and birds (Appleton, 1928; Howell,
1938). In modern Aves, knee exion is more important
to locomotion than femoral retraction (Gatesy and Biew-
ener, 1991; Carrano, 1998; Farlow et al., 2000), and the
M. caudofemoralis is greatly reduced in most birds and
altogether absent in others (Gatesy, 1990). In crocodili-
ans and the majority of non-serpente squamates, how-
ever, the M. caudofemoralis is both the primary and the
single largest retractor muscle of the hind limb (Snyder,
1962; Gatesy, 1990). Electromyographic studies of walk-
ing and running crocodiles have shown the M. caudofe-
moralis to be consistently active at all speeds, whenever
femoral retraction occurs (Gatesy, 1997).
The presence of large caudofemoral muscles in non-
avian dinosaurs was noted as early as 1833, when Louis
Dollo inferred their existence from the large femoral
fourth trochanter of the herbivorous dinosaur Iguanodon
(Dollo, 1883). In Gatesys 1990 study of the M. caudofe-
moralis and its reduction in the lineage leading to
extant birds, he argued that the muscles gradual
shrinkage resulted from a push towards overall tail
weight reduction and from the tails functional shift
from locomotion to dynamic stabilization, with both
trends relating to the evolution of ight (Gatesy, 1990).
Gatesy also argued insightfully that the M. caudofemor-
alis undoubtedly made the locomotion of most non-avian
dinosaurs (perhaps, particularly the bipedal taxa) funda-
mentally different from what can be observed in any
modern mammalian or avian analogue (Gatesy, 1990).
Despite such arguments, the M. caudofemoralis has
been undervalued in the majority of dinosaur biome-
chanical studies, and its contribution to locomotion has
not been quantitatively analyzed. Moreover, the M. cau-
dofemoralis has become a point of anatomical confusion,
with different authors using different osteological corre-
lates to infer its overall size and shape (Madsen, 1976;
Carpenter et al., 2005; Arbour, 2009; Schwarz-Wings
Fig. 1. Tyrannosaurus dorsal silhouette (A) previously used in body-
mass estimation (Modied from Paul, 1997), compared with a modern
Alligator dorsal silhouette (B) (Modied from Cong, 1998). The basal
bulge in the tail of the Alligator is primarily the result of a large M.
caudofemoralis.
et al., 2009). As a consequence of this confusion, the tails
of non-avian dinosaurs are commonly reconstructed with
improbable muscle morphology and overly conservative
masses, which appear altogether emaciated when com-
pared to the tails of modern reptiles (Fig. 1).
Here, basic caudal musculature of modern reptiles is
used for more accurately reconstructing the muscle anat-
omy of dinosaur tails. The locomotive implications of the
reconstructions are considered both qualitatively and
quantitatively for non-avian theropods (the primarily
carnivorous group of dinosaurs that gave rise to birds
and for which the majority of previous locomotive biome-
chanical research has been done).
MATERIALS AND METHODS
Anatomy
The pelvic and post pelvic musculature of a Basiliscus
vittatus (brown basilisk), Caiman crocodilus (spectacled
caiman), Chamaeleo calyptratus (veiled chameleon),
Iguana iguana (green iguana), and Tupinambis meria-
nae (Argentinean black and white tegu) were dissected.
All specimens had been preserved via deep freezing and
showed no signs of desiccation prior to dissection. Imme-
diately following in situ observations and measurements,
individual muscles were removed from each specimen
and massed (Table 1). Epaxial muscles that were contin-
uous into the pre-pelvic region were severed in line with
the midpoint of the acetabulum.
Three advanced non-avian theropods are considered in
this study: Gorgosaurus libratus (TMP 1991.36.500),
Ornithomimus edmontonicus (TMP 1995.11.001), and
Tyrannosaurus rex (cast of BHI 3033 currently on loan
to the Smithsonian National Museum of Natural His-
tory). The specimens were chosen based on the complete-
ness of the caudal series. Osteological dimensions of the
femora, pelvic girdle, and caudal series were taken, and
these measurements were later used in modeling digital
skeletons. TMP 1991.36.500 is a juvenile; however, com-
parisons between the measurements of TMP 1991.36.500
and those of less complete but more mature Gorgosaurus
specimens (TCM 2001.89.1 and AMNH 5458) indicate
that proportions of the caudal skeleton varied little
through ontogeny. Because both TMP 1991.36.500 and
TMP 1995.11.001 are panel mounts, some measurements
 THE TAIL OF TYRANNOSAURUS 121
TABLE 1. Caudal muscle mass of theropods and extant reptiles
M. M. ilio-
M. spinalis longissimus ischiocaudalis M. caudofemoralis
Caiman Crocodilus Mass (g) 1.4 9.6 9 13.8
Spectacled caiman Percent 4.1% 28.4% 26.6% 40.8%
Tupinambis merianae Mass (g) 3.9 22.5 12.2 13.3
Black and white tegu Percent 7.5% 43.4% 23.5% 25.6%
Iguana iguana Mass (g) 20.8 52.6 70 49.2
Green iguana Percent 18.8% 27.3% 36.3% 25.5%
Basiliscus vittatus Mass (g) 0.6 2.4 3 3.4
Brown basilisk Percent 6.4% 25.5% 31.9% 36.2%
Chamaeleo calyptratus Mass (g) 0.2 0.6 0.8 0.6
Veiled Chameleon Percent 9.1% 27.3% 36.4% 27.3%
Ornithomimus edmontonicus Mass (g) 860 2440 5050 9890
TMP 95.11.001 Percent 4.7% 13.4% 27.7% 54.2%
Gorgosaurus libratus Mass (g) 3900 6900 10300 17,300
TMP 91.36.500 Percent 10.2% 18.0% 26.8% 45.1%
Tyrannosaurus rex Mass (g) 65200 154200 159400 522200
BHI 3033 Percent 7.2% 17.1% 17.7% 58.0%

(such as transverse centrum width) could not be made
for every vertebra in the series, and measurements had
to be supplemented with those made on other similarly
sized specimens (including AMNH 514, AMNH 5355,
TMP 1999.33.1, and NMNH 2164). Likewise, the
mounted skeleton of BHI 3033 was so large and difcult
to reach that some measurements were supplemented
with those made indirectly from photographs. Whenever
possible, previously collected data on the original speci-
men of BHI 3033 were used to verify the accuracy of
these measurements. Measurements were also made on
published illustrations of other Tyrannosaurus speci-
mens (Paul, 1996; Brochu, 2002) and were used to sup-
plement portions of the tail that were not preserved in
BHI 3033.
Computer Modeling
Using the modeling software Rhinoceros (McNeel Rob-
ert and Associates, 2007), the hips and caudal series of
each of the three theropod specimens were digitally
sculpted based on the skeletal measurements. Three-
dimensional models of the various caudal muscles were
then sculpted overtop of the digital skeleton, according
to the osteological relations determined from the dissec-
tions. The volume of each restored muscle was then cal-
culated by the software. Because muscle is known to
have a fairly constant density of 1.06 g/cm3 (Mendez and
Keys, 1960), the muscle volume estimates could then be
multiplied by this density value to obtain estimates of
muscle mass.
To verify the accuracy of the restoration techniques
used in sculpting the various theropod caudal muscles,
digital skeletons were also made from measurements
taken on three of the dissected reptiles: the Argentinean
black and white tegu, green iguana, and spectacled cai-
man. Digital skeletons were not made for the brown bas-
ilisk and the veiled chameleon, because these specimens
were too small to measure using the same methods and
equipment. The same modeling techniques were then
used to reconstruct the musculature and to estimate
muscle volume and mass for the three extant reptiles,
and these muscle mass estimates were then compared to
the true muscle masses as measured during the
dissections.
Biomechanics
To evaluate the potential contribution of the M. caudo-
femoralis to femoral retraction, the digital muscle recon-
structions of each theropod were used in combination
with a series of standard biomechanical equations. First,
M. caudofemoralis mass was calculated using a standard
muscle density of 1.06 g/cm3 (Mendez and Keys, 1960).
Then, the physiological cross-sectional area (PCSA) of
the M. caudofemoralis was calculated according to Eq.
(1) (Sacks and Roy, 1982; Snively and Russell, 2007),
where m is the total mass (the mass of one M. caudofe-
moralis, not the mass of the total bilaterally symmetric
muscle set); r is the pennation angle of the muscle bers
within the M. caudofemoralis and, based on measure-
ments of the dissected specimens, is assumed to be
approximately equal to 1.00 (0.95), d is muscle density
(again, assumed to be 1.06 g/cm3), l is the average fasci-
cle length within the M. caudofemoralis and was calcu-
lated for each taxa based on the digital model and close
examination of the fascicles in the spectacled caiman
specimen:
PCSA m cos r=dl (1)
Because calculating physiological cross-sectional area
requires several assumptions, many authors favor the
simpler method of measuring anatomical cross-sectional
area (ACSA), which is simply the cross-sectional area
through the muscle at its greatest width (Snively and
Russell, 2007). Anatomical cross-sectional areas were
also measured from the digital models and compared to
the calculated physiological cross-sectional areas. In all
three instances, the two values varied only slightly from
one another, as would be expected, given the assumed r
value of approximately 1 (Bamman et al., 2000)
(Table 2).
Specic tension (ST), also sometimes referred to as
specic force (Brooks and Faulkner, 1994), is a ratio of
the strength of a muscle to its area and has been shown
to vary among different muscles and among different
122 PERSONS IV AND CURRIE
TABLE 2. Cross-sectional area in the M. caudofemoralis among theropod taxa
Mass (g)
Ornithomimus edmontonicus (TMP 95.11.001) 4945
Gorgosaurus libratus (TMP 91.36.500) 8650
Tyrannosaurus rex (BHI 3033) 261,100
TABLE 3. Summary of previously reported specic tension values for hind limb
muscles of extant animals
Animal Muscle(s)
Domestic cat Caudofemoralis
Human Thigh average
Quarter horse Gluteus medius
White rhinoceros Hamstring
taxa. Known ST values for hind limb muscles likely offer
the best basis for estimating the ST value appropriate
for the M. caudofemoralis of non-avian theropods. An
overview of previously reported ST values that might be
good analogs for the specic tension of the M. caudofe-
moralis of non-avian theropods is provided in Table 3.
Estimates within the range of 2535 N/cm2 would seem
reasonable. Here, ST is assumed to be 25 N/cm2 for all
the theropod taxa (certainly a conservative estimate).
The total contractile force (Ft) of the M. caudofemora-
lis was estimated by multiplying the calculated PCSA by
the assumed ST (25 N/cm2):
Ft PCSA ST (2)
Although the calculated total contractile force of the M.
caudofemoralis is useful as a comparable factor between
taxa, a more accurate measure of its true contribution to
locomotion is its capacity to generate torque. As outlined
in Snively and Russell (2007), a muscles torque-generat-
ing capacity is proportional to the force the muscle
exerts in a direction orthogonal to its moment arm (the
moment arm being measured from the joints center of
rotation to the muscles insertion). This orthogonal force
is here termed the effective force (Fe). The Fe of the M.
caudofemoralis is calculated with the following equation,
where y is the angle between the vector of line of pull of
the M. caudofemoralis and a vector orthogonal to the
moment arm (for the M. caudofemoralis, the moment
arm is the dorsal-ventral distance from the midpoint of
the femoral head to the midpoint of the femoral fourth
trochanter), calculated for a femur positioned perpendic-
ular to the ground:
Fe Ft
cos h (3)
Finally, the potential torque generation of the M. caudo-
femoralis is calculated with the equation:
sm R
Fe
sin U (4)
Here, U is the angle between the vectors of the moment
arm and the effective force (this angle is assumed to be
90 and the sinU is, therefore, equal to 1), and R is the
Average fascicle PCSA ACSA PCSA/ACSA
length (cm) (cm2) (cm2) variation
31.423 141.038 127.675 1.105
50.585 153.254 144.486 1.061
129.697 1804.242 1794.442 1.005
ST (N/cm2) Source
31.2 Brown et al. (1997)
27.330.0 Storer et al. (2003)
25.9 Marx et al. (2005)
34.4 Marx et al. (2005)
length of the moment arm (again assuming a femur
positioned perpendicular to the ground).
RESULTS
Caudal Musculature and Osteological
Correlates of Extant Reptiles
The literature is lled with inconsistent and often
redundant terminology schemes for the caudal muscles
of reptiles. Following the recent scheme established by
Arbour (2009), I will acknowledge four major paired
muscle sets: the M. spinalis, M. longissimus, M. ilio-
ischiocaudalis, and M. caudofemoralis. It should be
noted that these four muscles form a good general, but
overly simplistic, scheme, and subdivisions are clearly
visible within many of these muscles, particularly in the
posterior portions of the M. longissimus and M. ilio-
ischiocaudalis. Like those of the limb, caudal muscles
vary widely in size and relative proportions across reptil-
ian taxa; however, the overall pattern of muscle inser-
tions and positions remains relatively uniform (Figs. 2
and 3). The results presented here are based on the dis-
sections of ve taxa (one specimen each) and, combined
with the previous literature, are intended to offer only a
cursory overview of reptilian tail musculature.
On the basis of the dissections, the arrangement and
osteological insertion sites of the caudal muscles are as
follows.
M. spinalis. The M. spinalis is the most dorsal of the
caudal muscles and is present along the full length of
the tail. The M. spinalis inserts onto the dorsal tips and
the full lateral surfaces of the neural spines.
M. longissimus. Composing the primary bulk of the
epaxial tail musculature, the M. longissimus is also
present along the full length of the tail and may be dif-
cult to distinguish from the M. spinalis. This distinction
becomes more difcult posteriorly, when subdivisions in
the two muscles become increasingly independent. The
M. longissimus has two strong dorsoventrally stacked
subdivisions that are distinguishable in even the most
anterior of cross sections. These two subdivisions are of-
ten regarded as separate muscles, with the more ventral
 THE TAIL OF TYRANNOSAURUS
Fig. 2. Sequential cross-sections through the tail of Tupinambis
merianae. Tupinambis merianae was chosen to illustrate the anterior/
posterior changes in muscle arrangement and morphology, because
its tail is relatively unspecialized and its musculature is unremarkable
in most respects. The cross-sections are anatomical abstractions, as
they depict neural arches and hemal arches in the same vertical plane
(these illustrations are similar in this way to those of Cong et al., 1998;
Arbour, 2009; Schwarz-Wings et al., 2009), but these views are prefer-
able to ones that would omit either hemal or neural arches. A: Caudal
vertebra 7; the M. caudofemoralis inserts across the lateral sides of
the centrum and across the entire lateral face of the hemal arch. The
M. ilio-ischiocaudalis is thin medially, but inserts across the entire
ventral surface of the transverse processes, and only inserts onto the
ventral tip of the hemal arch. B: Caudal vertebra 13; the M. caudofe-
moralis has begun to taper and the M. ilio-ischiocaudalis inserts
across a ventral portion of the lateral face of the hemal arch. C: Cau-
dal vertebra 18; the M. caudofemoralis is no longer present. The M.
ilio-ischiocaudalis insets across the entire lateral face of the hemal
arch and the lateral sides of the centrum. The transverse processes
are absent. (Note, the posterior portion the tail is also a fat storage
location in Tupinambis merianae; however, for simplicity, fat deposits
have been not been shown).
subdivision retaining the name M. longissimus, and the
more dorsal subdivision occasionally referred to as the
M. tendinoarticularis (although this term has also been
applied to the M. spinalis). There is no clear osteological
indicator of the level at which these two M. longissimus
subdivisions meet, and the functional differences
between the two are currently unknown. Attempts
herein to subdivide the M. longissimus of theropods
would, therefore, be both highly speculative and ulti-
mately uninformative.
Anteriorly, the M. longissimus inserts onto the full
dorsal surfaces of the transverse processes and to the
lateral faces of the neural arches. Posteriorly, after the
transverse processes have terminated, the lateral faces
of the neural arches are the only osteological insertion
points of the M. longissimus. Throughout its anterior/
posterior length, the M. longissimus is separated dor-
sally from the M. spinalis by a septum that stems from
the lateral edges of the pre-zygapophyses and extends
lateral at an often strongly dorsally inclined angle.
[Note: Consistent with convention the large lateral pro-
jections of caudal vertebra are herein referred to as
transverse processes. However, as originally argued by
123
Fig. 3. Lateral view (anterior left and posterior right) of the dis-
sected tail of Tupinambis, with all muscles, except the M. caudofemor-
alis, removed. Highlighted regions correspond to cross-sections in
Fig. 2.
Romer (1956), the majority of these projections are prob-
ably homologues with the ribs of the dorsal series and
are not true transverse processes. The term caudal
ribs is likely more accurate for all but the most poste-
rior caudal vertebra]. Both the M. longissimus and M.
spinalis are continues into the dorsal region (Organ,
2006).
M. ilio-ischiocaudalis. Consistent with its nomen-
clature, the M. ilio-ischiocaudalis is composed of two
major subdivisions: the m. iliocaudalis and the m.
ischiocaudalis, with the former originating from the
ilium and the later from the ischium. The M. ilio-ischio-
caudalis is relatively thin anteriorly, where it attaches
dorsally to the lateral tips and ventral surfaces of the
transverse processes, raps around the M. caudofemora-
lis, and attaches ventrally both to the ventral tips of the
hemal spines and to its bilaterally symmetric muscular
doppelganger that raps around from the other side. Pos-
teriorly, the M. ilio-ischiocaudalis increases in relative
thickness as the thickness of the M. caudofemoralis
diminishes. After the disappearance of the M. caudofe-
moralis, the M. ilio-ischiocaudalis inserts onto the full
lateral surface of the centrum and the hemal spines.
M. caudofemoralis. Unique among the caudal
muscles, the M. caudofemoralis is not partitioned by
conical myosepta, and its overall form more closely
resembles a limb muscle rather than an axial muscle.
The primary caudofemoral tendon attaches the M. cau-
dofemoralis to the fourth trochanter of the femur, and
the auxiliary caudofemoral tendon (present in most
extant reptiles but absent in birds) also anchors the M.
caudofemoralis to the knee joint (Fig. 4).
The M. caudofemoralis is composed of the m. caudofe-
moralis brevis, which lls the brevis fossa and may also
insert across the anteriormost caudal vertebra, and the
m. caudofemoralis longus. Throughout its anterior/poste-
rior length, the m. caudofemoralis longus inserts medi-
ally onto the lateral faces of the caudal vertebra.
However, the insertion of the m. caudofemoralis longus
across the lateral faces of the hemal spines varies across
the caudal series (Fig. 2). The M. caudofemoralis does
not extend down the entire length of the tail (in none of
124 PERSONS IV AND CURRIE
Fig. 4. The anterior end of the M. caudofemoralis, its insertion onto
the fourth trochanter of the femur, and the auxiliary tendon that inserts
at the knee joint, seen in (A) Caiman (oblique dorsolateral view, ante-
rior top, and posterior bottom) and (B) Tupinambis (oblique posterolat-
eral view, anterior upper right, and posterior bottom left).
the dissected reptiles did the M. caudofemoralis extend
across even half of the tails total length). Rather, the M.
caudofemoralis extends posteriorly down the caudal se-
ries with a gradual reduction in overall size, then
reaches a point where it begins to rapidly taper, and,
shortly thereafter, terminates (Fig. 3).
Prior to the taper point, the M. caudofemoralis medial
insertion extends ventrally across the entire lateral face
of the hemal arches. As it tapers, the M. caudofemoralis
medial insertion shrinks dorsally; correspondingly, the
ventral insertion of the M. ilio-ischiocaudalis expands
dorsally, and the M. ilio-ischiocaudalis begins to insert
across an increasing proportion of the lateral faces of the
hemal arches. Past the termination of M. caudofemora-
lis, the M. ilio-ischiocaudalis inserts across the entire
lateral faces of both the hemal arches and the centrum.
As they are no longer needed to provide dorsal insertion
points for the M. ilio-ischiocaudalis, the transverse proc-
esses are usually no longer present after the M. caudofe-
moralis termination. However, because the transverse
processes also serve as lateral insertion points for the M.
longissimus, they may (as is the case in the Argentinean
black and white tegu) remain present well after the M.
caudofemoralis has terminated.
Fig. 5. Lateral view (anterior right and posterior left) of the M. cau-
dofemoralis termination point of Ornithomimus TMP 1995.11.001. Cau-
dal vertebra 14 bears the last transverse processes, which sit at
roughly mid dorsal/ventral height. Hemal arch 13 (between caudal ver-
tebra 13 and 14) is displaced slightly, with its posterior end tilted dor-
sally. Hemal arch 13 has a strong grove or scar across its lateral face.
Reproduced from Arbour, 2009.
Fig. 6. Lateral view (anterior right and posterior left) of caudal verte-
bra 1017 of Ornithomimus TMP 1995.11.001. Arrows point to the an-
terior edges of the M. ilio-ischiocaudalis/M. caudofemoralis septum
scarring, visible on hemal arches 1114 and the posterior end of
hemal arch 10. Reproduced from Arbour, 2009.
Evidence for the Posterior Taper Point and
Ventral Boundary of the M. caudofemoralis in
Non-Avian Theropods
Caudal 14 is the last transverse process-bearing verte-
bra of Ornithomimus TMP 1995.11.001 (Fig. 5). The
transverse process is located at roughly the centrums
dorsoventral midpoint. Hemal arch 13 (positioned
between caudal vertebra 13 and 14) is displaced slightly,
with its posterior end tilted dorsally. A groove or scar is
apparent running across its lateral face.
The scar seen on hemal arch 13 is part of a continuous
diagonal sequence of scars that runs anteroventral to
posterodorsal across hemal arches 1014 parallel to the
axis of the tail (Fig. 6). This scar sequence coincides
with the rapid decent of the transverse processes from
an elevated height above the centra (a full 19.8 mm
above at caudal vertebra 1) to the mid-centrum elevation
seen on caudal vertebra 14. Together, the descending
transverse processes and hemal arch scarring are here
interpreted as demarcating the boundaries of the
 THE TAIL OF TYRANNOSAURUS
Fig. 7. Lateral view (anterior right and posterior left) of caudal verte-
bra 13-8 of Gorgosaurus TMP 91.36.500. The last transverse process
is present on caudal vertebra 12. Although fainter than TMP
1995.11.001, the M. ilio-ischiocaudalis/M. caudofemoralis septum
scarring is visible on hemal arches 11-7. Arrows point to the anterior
edges of the scarring.
tapering M. caudofemoralis. Under this interpretation,
the hemal arch scar is regarded as the insertion point
for the skeletogenous septum that divided the M. caudo-
femoralis from the M. ilio-ischiocaudalis, and the ante-
roposterior ascent and eventual posterior disappearance
of this scar is, therefore, taken to mark the M. ilio-
ischiocaudalis gradual dorsal intrusion and eventual
usurpation of the M. caudofemoralis.
The M. ilio-ischiocaudalis/M. caudofemoralis septum
scarring disappears anteriorly, reaching the ventral tip
of hemal arch 10. The scar does not reappear on any of
the more anterior hemal arches, and this is interpreted
as evidence that, prior to the taper point, the M. caudo-
femoralis inserted across the entire lateral surfaces of
the hemal arches. This is in contradiction to some resto-
rations that have shown the M. caudofemoralis of dino-
saurs riding high on the caudal series (see Arbour, 2009;
for one example), but is entirely consistent with the mor-
phology seen in all of the dissected reptiles.
TMP 1995.11.001 displays the M. ilio-ischiocaudalis/
M. caudofemoralis septum scar sequence more clearly
than any of the other fossil specimens examined in this
study; however, the scar sequence is by no means a fea-
ture unique to TMP 1995.11.001 or to Ornithomimus.
The scars of TMP 1991.36.500 (Gorgosaurus), which,
although faint, are unambiguously present (Fig. 7). The
M. ilio-ischiocaudalis/M. caudofemoralis septum scars
have not gone unnoticed by previous authors. For
instance, although no explanation for the structure is
offered, Brochu (2002) notes it existence in his wonder-
fully thorough description of the Tyrannosaurus speci-
men FMNH PR2081.
Evidence for the Lateral Boundary of the
M. caudofemoralis in Non-avian Theropods
To model the M. caudofemoralis of theropods, the
muscles extent must be dened in all dimensions. The
femoral fourth trochanter is the muscles anteriormost
insertion point. The transverse processes cap its dorsal-
most limit, and the ventralmost margins of the hemal
125
Fig. 8. Comparison of anterior caudal cross-section reconstructions
for (A) Allosaurus (Modied from Madsen, 1976), with greatest width
assumed to be roughly equal to the lateral extent of the transverse
processes and (B) Caiman, reconstructed based on dissection and
showing greatest lateral extent to be at roughly mid hypaxial height.
arches indicate its ventral extent. Both the M. ilio-
ischiocaudalis/M. caudofemoralis septum scars and the
rapid descent of the transverse processes mark its poste-
rior taper point. What remains is determining the lat-
eral extent of the M. caudofemoralis.
Traditionally, the lateral width of the caudal trans-
verse processes has been used as a correlate for the girth
of the M. caudofemoralis in all saurians. However, this
method is without sound anatomical basis. As described,
and contrary to the depictions and descriptions of Cald-
well (2006), Arbour (2009), Schwarz-Wings et al. (2009),
and various others, the M. caudofemoralis does not
attach to the lateral tips or ventral surfaces of the trans-
verse processes. Rather, both are insertion points of the
M. ilio-ischiocaudalis (the near vertical caudal trans-
verse processes of some advanced abelisaurids are a pos-
sible exception).
When viewed casually in most reptile dissections the
transverse processes are easily mistaken for an M. cau-
dofemoralis insertion point, because the portion of the
M. ilio-ischiocaudalis that attaches to the transverse
processes is usually quite thin. Adding to this confusion
is the high prole and commonly cited 1923 reconstruc-
tion by Alfred Romer of the hip and hind limb muscula-
ture of Tyrannosaurus, in which the M. caudofemoralis
is clearly, but incorrectly, shown attaching to the lateral
tips of transverse processes (Romer, 1923). Romer does
comment in the same manuscript that there is a lack of
good tail muscle data available for crocodilesthe ana-
logue on which he most heavily reliedand, 4 years
later, he did correct himself when reconstructing the hip
and hind limb musculature of ornithischian dinosaurs
(Romer, 1927). Nonetheless, his Tyrannosaurus recon-
struction remains the one to which most researchers
have paid attention.
The transverse processes also do not appear to be good
indirect indicators of the M. caudofemoralis lateral
extent. The spectacled caiman, for example, had the rel-
atively largest M. caudofemoralis of any of this studys
126 PERSONS IV AND CURRIE

Fig. 9. Tyrannosaurus BHI 3033 in dorsal view with M. caudofemoralis musculature reconstructed and
in lateral view with full caudal musculature reconstructed. Skeletal image Modied from Paul, 1989.

extant reptiles, but the lateral widths of its transverse

processes, relative to other aspects of its caudal verte-
brae, were only average. In contrast, the dissected
Argentinean black and white tegu specimen had among
the relatively smallest caudofemoral muscles, but the
relatively widest transverse processes. The dissections
also revealed that the width of the M. caudofemoralis
varies relative to the length of the transverse processes
across the caudal series of the same individual, with the
caudofemoral muscles extending well beyond the lateral
tips of the transverse processes at the anterior base of
the tail, and usually extending well short of the tips at
and near the taper point.
Recognizing that the transverse processes do not bear
a M. caudofemoralis insertion and that they are not
good indicators of its lateral width is important, because
this has dramatic implications for the overall shape of
the hypaxial region of the tail. Figure 8a shows a
restored cross-section through the anterior tail base of
Allosaurus from Madsen (1976). Madsens reconstruction
follows the traditional restoration method, and assumed
a caudofemoralis that inserted onto the ventral surfaces
of the transverse processes and that did not bulge past
their ventral-most tips. As a result, the tail is widest
dorsal to its midpoint, at or just below the level of the
transverse processes, producing an inverted tear-drop
shaped cross-section. For comparison, the widest portion
of the tail of a modern caiman is at roughly mid-hypax-
ial height (Fig. 8b).

Muscle Reconstruction
In creating this studys reconstructions (Figs. 911),
the vertical cross-section of the M. caudofemoralis was
treated as elliptical in overall shapeconsistent with
the M. caudofemoralis in all the dissected specimens
and was modeled across the caudal series (Fig. 10) by
tracing an arc from below the medioventral base of the
transverse process to the ventral tip of the hemal arch.
Comparisons, based on the dissected reptile speci-
mens, between mass predictions for the M. caudofemora-
Fig. 10. Prior to the anterior tapering of the M. caudofemoralis
longissimus, its lateral margin was reconstructed at each vertebra, as
shown, by tracing a symmetric arch from below the medioventral base
of the transverse process to the ventral tip of the nearest hemal arch.
lis derived from digital reconstruction models and the
M. caudofemoralis masses actually measured, conrm
the accuracy of this restoration technique (see Table 4).
All predicted M. caudofemoralis masses are within 6% of
 THE TAIL OF TYRANNOSAURUS 127

Fig. 11. Modeling sequence for the tail of Tyrannosaurus BHI 3033. A: Skeleton model constructed based
off dimensional measurements. B: Model with M. caudofemoralis longissimus sculpted overtop of the skele-
ton model. C: Full muscle restoration with M. spinalis, M. longissimus, and M. ilio-ischiocaudalis visible.

the true mass and slightly lower than the true value.
This is probably because, at the anterior-most base of
the tail, the M. caudofemoralis has a more extensive lat-
eral bulge that diverges from the overall shape seen
more posteriorly. The mass estimates presented here
are, therefore, conservative.
The measured tail muscle masses of the dissected
extant reptiles and the estimated tail muscle masses of
the three theropods are presented with the percentages
of total tail muscle mass (Table 1). These percentages
offer one way to quantify the relative size of the differ-
ent tail muscles across the various taxa.
Among the extant taxa, the spectacled caiman (the
only crocodilian considered in this study) had the high-
est percentage of its tail muscle mass contributed by the
M. caudofemoralis. The brown basilisk had the second
128 PERSONS IV AND CURRIE

TABLE 4. True M. caudofemoralis mass versus

estimated mass, based on digital models, in the
extant reptiles
True Modeled
mass (g) mass (g) Accuracy
Caiman crocodilus 13.8 12.997 5.8%
Tupinambis merianae 13.3 12.692 4.6%
Iguana iguana 49.2 48.194 2.0%

highest caudofemoral percentage. Given the caimans

closer phylogenetic relation and the basilisks bipedal
running style, the large M. caudofemoralis found in both
of these taxa could be taken as indirect support for a
large M. caudofemoralis in bipedal non-avian theropods.
Among the three extinct taxa, Tyrannosaurus had the
largest relative (and absolute) M. caudofemoralis. How-
ever, all three theropods were estimated as having sub-
stantially larger caudofemoral muscles than any of the
modern reptiles. Given that the M. caudofemoralis is the
primary retractile muscle of the hind limb, and given
the numerous advanced locomotive adaptations present
in non-avian theropods, this high relative investment in
caudofemoral mass should, perhaps, not be surprising.
But the osteological features associated with this caudo-
femoral bulk-up do merit special note. Two such features
are readily identiable from this study: the strong dor-
sally inclined angle of the transverse processes and the
elevation of the transverse processes on the neural arch
(Fig. 12).
The caudal transverse processes of all the dissected
extant reptiles protrude at at, fully horizontal angles.
Anterior vertebrae with dorsally angled transverse proc- Fig. 12. Anterior view of caudal vertebra 1 of Ornithomimus NMNH
esses are present in many theropod taxa, and the angled 2164. Note the dorsally angled transverse processes, and the dorsal
transverse processes would have provided an expanded elevation of the transverse processes above the centrum, which cre-
ate an expanded hypaxial region.
hypaxial region that the M. caudofemoralis could have
lled. However, angled transverse processes are not
characteristic of all theropod taxa (they are absent in data can most condently be used to compare different
both Tyannosaurus and Gorgosaurus), and, when pres- theropod taxa, but it should be noted that the high tor-
ent, usually only occur on the rst three or four caudal que estimates calculated for all three theropod taxa are
vertebra. consistent with what would generally be predicted for
More important, then, would seem to be the elevation muscles of their size based on what is known about the
of the transverse processes on the neural arch, above physiology of smaller modern taxa.
the centrum. This too creates a greatly expanded hypax-
ial region (at the expense of the epaxial region), and the
M. caudofemoralis is the only muscle that is likely to DISCUSSION
have lled this space. Dorsally elevated transverse proc-
Among the three theropods examined here, an intrigu-
esses are found in even primitive theropods, including
ing comparison can be made between the estimated M.
Coelophysis and Herrerasaurus. This suggests that a
caudofemoralis masses of Tyrannosaurus BHI 3033 and
large M. caudofemoralis is a basal characteristic of the
the juvenile Gorgosaurus TMP 1991.36.500. As tyranno-
group and appears to have remained present in nearly
saurids, the taxa are closely related, and one might be
all non-avian theropod lineages (the character appears
surprised by the substantially larger tail mass percent-
to be absent in the highly derived and specialized tails
age that the M. caudofemoralis of Tyrannosaurus is pre-
of Paraves). Elevated transverse processes are not only
dicted to comprise (58.0%) compared with that of
present on the anteriormost tail vertebra of most non-
Gorgosaurus (45.1%). However, this discrepancy in rela-
avian theropods, but remain elevated down the series,
tive M. caudofemoralis mass makes sense when consid-
until rapidly descending at the taper point.
ering the absolute body mass discrepancy between the
two tyrannosaurids.
Based on mass estimates of other Gorgosaurus speci-
Biomechanics
mens (Paul, 1988), the body mass of TMP 1991.36.500
Interpreting the results of the biomechanical calcula- can be estimated as roughly 400 kg, and the mass of
tions performed for each of the three theropods (Table 5) BHI 3033 has been estimated as in the range of 3,800
is hindered by the lack of analogous data available for 4,500 kg (Stevens et al, 2008; but see Bates et al., 2009
modern animals with muscles of equivalent size. The for an alternative interpretation). The need for relatively
 THE TAIL OF TYRANNOSAURUS
TABLE 5. Summary of estimated biomechanical data for the M. caudofemoralis
of the theropods
Ornithomimus
(TMP 95.11.001)
Mass (g) 4945
PCSA (cm2) 141.038
Total contractile force (N) 2820.762
Effective muscle pull (N) 2820.480
Moment arm (m) 0.081
Torque (Nm) 228.459
larger locomotive muscles in absolutely larger taxa has
been well established (Biewener, 1989; Roberts, 1998). A
muscles strength is largely a factor of its cross-sectional
area, and locomotive muscles must be strong relative to
the mass of the body they are trying to accelerate. Sim-
ple isometric growth of any animal would result in a
three-fold increase in body mass (as mass is largely a
function of volume) and only a two-fold increase in the
strength of its muscles. Hence, the general rule that
larger animals require relatively larger muscles to
achieve the same speeds as smaller animals.
That is not to suggest this relatively enlarged M. cau-
dofemoralis estimation indicates that Tyrannosaurus
could have achieved the same degree of cursoriality as a
juvenile Gorgosaurus or other smaller tyrannosaurids.
Indeed, the relatively shorter metatarsals of Tyranno-
saurus (among other anatomical features) testify that it
could not (Holtz, 1995). Nonetheless, it seems likely that
the high relative M. caudofemoralis mass of Tyrannosau-
rus did evolve as partial compensation for its colossal
body size, and it is worth noting how this increased M.
caudofemoralis mass was achieved. The higher M. cau-
dofemoralis mass estimation for Tyrannosaurus is not
the result of relatively more dorsally angled or elevated
transverse processes, but of more ventrally elongated
hemal arches, which means that the hypaxial muscula-
ture was increased without decreasing the size of the
epaxial musculature.
The expanded masses and high contractile force and
torque estimates for all three theropods conrm previous
assertions that the M. caudofemoralis was indeed a mus-
cle of fundamental importance to non-avian theropod
locomotion. These results have implications for the
ongoing discussion of the potential locomotive abilities of
non-avian theropods. For instance, in their assessment
of the cursoriality of Tyrannosaurus, Hutchinson and
Garcia (2002) assumed a total femoral retractor muscle
mass of 297 kg for each leg of a 6,000 kg Tyrannosaurus.
Here, the mass of the M. caudofemoralis alone has been
conservatively estimated as 261 kg for each femur of a
Tyrannosaurus previously estimated to have weighed as
little as 4,500 kg, which implies that the M. caudofemor-
alis should have a mass of 348 kg in a 6,000 kg individ-
ual. Hutchinson and Garcia (2002) did not provide mass
estimates for individual limb muscles, making it difcult
to assess how this new M. caudofemoralis data affects
their total estimation. However, it can surely be
assumed that the mass of the other femoral retractors
(the M. adductor femoris, M. puboischiofemoralis exter-
nus, and M. ischiotrochantericus) in a 6,000 kg Tyranno-
saurus would weigh at least 25 kg. Under this
assumption, the 297 kg estimation appears to be off by
129
Gorgosaurus libratus Tyrannosaurus
(TMP 91.36.500) rex (BHI 3033)
8650 261,100
153.254 1804.242
3065.082 36084.831
3064.775 36081.223
0.147 0.505
449.603 18208.028
over 25% (and is conceivably off by as much as 45%).
This by no means accounts for the 80% of total body
mass that Hutchinson and Garcia (2002) assert must
have been invested in the limb retractors, in order for
Tyrannosaurus to have been capable of rapid locomotion.
Nonetheless, the new M. caudofemoralis data does sug-
gest that Tyrannosaurus should have fallen towards the
higher end of Hutchinson and Garcias (2002) advocated
speed range, and the data are consistent with the faster
locomotive estimates advocated by other authors using
different speed estimating techniques (Bakker, 1986;
Paul, 2000; Sellers and Paul, 2005).
Considering the results of this study in the context of
such biomechanical studies points out the large gaps in
our current understanding of how tail muscularity is
involved in terrestrial locomotion. The M. caudofemora-
lis has been treated herein as the only tail muscle
involved in femoral retraction, but this is likely a gross
oversimplication, and the case can be made for the par-
tial involvement of other caudal muscles in femoral re-
traction as well. Studies of walking and running
alligators demonstrate that during retraction of the right
femur, the tail consistently swings to the left, and vice
versa (Reilly and Elias, 1998). Given the electromyogra-
phy evidence showing that during retraction of the right
femur, the right M. caudofemoralis retracts, one might
instead have predicted the tail to swing towards, not
away from, the right side. Pelvic rotation is partially re-
sponsible for this tail motion, but the left caudal muscles
also retract to pull the tail leftwards and do, thereby,
add to the right M. caudofemoralis femoral pull. The
elongate zygapophyses of Tyrannosaurus would likely
have reduced the overall lateral exibility of the tail, but
the assistance of other caudal muscles in femoral retrac-
tion remains plausible, and the recruitment of muscle
sets with no direct connections to limb bones has been
well documented in extant animals, such as the intercos-
tal muscles in running dogs (Carrier, 1996) or neck
muscles in galloping horses (Gellman et al., 2002).
Elasticity is another complication likely to have
improved the tails locomotive contribution. Tails are
naturally rich in tendons and septa, which are excellent
stores of elastic energy. Elastic elements within the tails
of both large and small non-avian theropods may have
greatly improved locomotive efciency beyond what
would be estimated based on the limb musculature of
most modern birds and mammals.
In addition to considerations of absolute speed, large
caudofemoral muscles also have implications for previ-
ous estimates of theropod centers of mass and, are there-
fore relevant to Hutchinson and Garcias 2002 study and
to Hutchinsons 2004 follow-up, which found via
130 PERSONS IV AND CURRIE
Fig. 13. A fully rendered reconstruction of BHI 3033 created by Scott Hartman to illustrate the appear-
ance of a Tyrannosaurus with a tail of appropriate beeness.
numerous sensitivity analyses that repositioning the
axial center of mass more posteriorly had the potential
to signicantly decrease the estimated muscle mass
needed by Tyrannosaurus to achieve higher speeds and
to support its own bulk. Obviously, a larger M. caudofe-
moralis mass results in a more posterior position of any
center of mass estimation (although the center of mass
would also be determined by the dorsoventral angle of
the torso and by the amount of curvature in the neck).
The similar enlargement of any of the other hind limb
retractors would have a largely neutral effect on the
axial center of mass. With its center of mass positioned
closer to its hips, a theropods leg muscles would be rela-
tively less strained in supporting its weight, and the ani-
mals overall balance and turning agility would be
improved, as it would be less front-end heavy and its
rotational inertia would be reduced.
The last point that should be made is primarily an
artistic one. The current prevalent fashion among paleo-
artists is to depict the tails of most dinosaurs, but partic-
ularly theropods, as relatively unmuscular and laterally
compressed. This is true not only of depictions made
strictly for aesthetic purposes but also of those intended
to support scientic research, such as estimations of
mass (for example, Paul, 1997; Bates et al., 2009). The
less-than-robust tail depictions are consistent with the
more traditional tail muscle restoration technique
described. They are also likely the result of the recent
trend towards depicting more lightly built and more
eet-footed theropods, because skinny laterally com-
pressed tails have a more aerodynamic and supercially
more athletic appearance. In reality, skinny tails are not
more athletic. Because the M. caudofemoralis is the pri-
mary retractor muscle of the hind limb, a slim-tailed
theropod would be inherently slower than one with a
large, muscular tail.
In overall appearance, the tails of most non-avian
theropods likely resembled those of their modern croco-
dilian relatives, with relatively larger hypaxial muscles
(and relatively smaller epaxial muscles) but without (in
most cases at least) dorsal osteoderms. At the anterior
base, the tails of most non-avian theropods would have
been as broad or broader laterally as they were tall dor-
soventrally. At and near the transition point, the tails
would be laterally compressed, and towards the poste-
rior tip, the tails would, as the neural spines and hemal
arches steadily shrunk, return to being roughly round in
cross-section (Fig. 13).
ACKNOWLEDGMENTS
This research was made possible by the nancial sup-
port of the Dinosaur Research Institute. We wish to
thank John Acorn, Victoria Arbour, Robert Bakker, Rob-
ert Holmes, Pierre Lemelin, and Eric Snively for their
repeated council and for many fruitful discussions about
dinosaur tails. We also wish to extend our gratitude to
Mike Brett-Surman, Chris Norris, and Brandon Strilisky
for their indispensable assistance in navigating the enor-
mous storerooms of their respective institutions. Scott
Hartman merits our gratitude (and envy) for his stun-
ning artistic skills and talents, which were graciously
lent to this project. Special thanks are also owed to Joe
Barter for his hospitality and assistance in recording tail
measurements. Lastly, we thank Michael Caldwell for
his unhesitating willingness to sacrice many tails from
his sizable frozen reptile collection.
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