JOURNAL OF MORPHOLOGY 270:856–879 (2009)
The Theropod Furcula
Sterling J. Nesbitt,1,2* Alan H. Turner,1,2 Michelle Spaulding,1,2 Jack L. Conrad1
and Mark A. Norell1
1
Division of Paleontology, American Museum of Natural History, New York, New York 10024
2
Lamont-Doherty Earth Observatory, Columbia University, Palisades, New York 10964
ABSTRACT The furcula is a structure formed by the
midline fusion of the clavicles. This is the element which
is unique to theropods and is important for understand-
ing the link between birds and other theropods. New
specimens from basal theropods suggest that the furcula
appeared very early in theropod history. We review fur-
cula development, function, and morphology, as well as
the anatomical terminology applied to it. Furcular mor-
phology is highly variable in crown-group avians but is
rather conserved among nonavian theropods. Here we
review, or describe for the first time, the furculae in
many nonavian theropods. Furculae occur in nearly all
major clades of theropods, as shown by new theropod
specimens from the Early Cretaceous of China and a
close inspection of previously collected specimens. In-
formative phylogenetic characters pertaining to the fur-
cula occur throughout Theropoda, though care should
be take to consider taphonomic effects when describing
furcular morphology. J. Morphol. 270:856–879, 2009.
Ó 2009 Wiley-Liss, Inc.
KEY WORDS: furcula; clavicle; theropod; homology
INTRODUCTION
Since the close evolutionary relationship between
birds and dinosaurs was first suggested (Huxley,
1868, 1870), the presence of the furcula (or its
homologues) has been a key component of the
debate. As the furcula in theropod dinosaurs was
unknown at the time, its ‘‘absence’’ was considered
to be an important piece of evidence for dismissing
the putative relationship. This was especially
apparent in the work of Heilmann (1926) where he
stated: ‘‘From this it would seem a rather obvious
conclusion that it is among the Coelurosaurs that
we are to look for the bird ancestor. And yet this
would be too rash, for the very fact that the
clavicles are wanting would in itself be sufficient
to prove that these saurians could not possibly be
the ancestors of birds’’ (Heilmann, 1926:182).
Since Heilmann’s monograph, the furcula has
been recognized in several different theropod
groups (Barsbold, 1983; Bryant and Russell, 1993;
Makovicky and Currie, 1998; Norell and Makov-
icky, 1999; Tykoski et al., 2002; Yates and Vascon-
celos, 2005). However, doubts concerning the
homology of the theropod ‘‘furcula’’ and the avian
‘‘wishbone’’ have persisted because of consistent
misinterpretations of the available data. Despite
Ó 2009 WILEY-LISS, INC.
mounting evidence supporting the homology of the
furcula in birds and other theropods, the same
questions and arguments resurface with little
apparent regard or acknowledgment of rebuttals
or new discoveries. Consequently, these authors
have used their own creations as evidence in sup-
port of a nondinosaurian origin for birds.
Recent identification of furculae in many thero-
pod clades (Barsbold, 1983; Bryant and Russell,
1993; Makovicky and Currie, 1998; Norell and
Makovicky, 1999; Tykoski et al., 2002; Yates and
Vasconcelos, 2005) has sparked interest in the dis-
tribution of the element among theropod dino-
saurs. Bryant and Russell (1993) reviewed the dis-
tribution of clavicles (and furculae) in dinosaurs
and found that it was unclear whether the avian
furcula is homologous to the clavicles of other rep-
tiles, or if it was a neomorphic bone shared by some
theropod dinosaur clades and birds. Increased
morphological knowledge of nonavian furculae has
led to their discovery or identification in progres-
sively more basal theropods. Moreover, the discov-
ery and identification of sauropodomorph (Huene,
1926; Yates and Vasconcelos, 2005) and ornithi-
schian (Osborn, 1924a; Brown and Schlaikjer, 1940;
Sternberg, 1951; Chinnery and Weishampel, 1998)
clavicles has allowed comparisons within Dinosau-
ria more generally. Recently discovered well-pre-
served, articulated specimens of theropod dino-
saurs and basal birds from the Early Cretaceous of
China have allowed the identification of undoubted
furculae in several theropod clades (Fig. 1). Many
Alan H. Turner is currently at: Department of Anatomical Sciences,
Stony Brook University, Stony Brook, New York 11794-8081.
Contract grant sponsor: NSF; Grant number: DDIG DEB 0608003,
ATOL 0228693; Contract grant sponsor: American Museum of Natu-
ral History and Columbia University.
*Correspondence to: Sterling Nesbitt, Division of Paleontology,
American Museum of Natural History, Central Park West at 79th
Street, New York, NY 10024. E-mail: nesbitt@ldeo.columbia.edu
Received 7 June 2008; Revised 8 November 2008;
Accepted 4 December 2008
Published online 10 February 2009 in
Wiley InterScience (www.interscience.wiley.com)
DOI: 10.1002/jmor.10724
 THEROPOD FURCULA
Fig. 1. Relationships among major clades of theropods. Rela-
tionships from Smith et al. (2007), the latest TWiG matrix in
Turner et al. (2007a), and Clarke et al. (2006). The small fur-
cula symbol indicates which clades have at least one member
with an unambiguous furcula.
specimens retain the furcula in articulation or in
close proximity to the pectoral girdles. The reinter-
pretation of previously unrecognized furculae has
also added to our knowledge of the distribution
and evolution of the furcula (Makovicky and Cur-
rie, 1998). Here, we will examine all known occur-
rences of the furcula among nonavian theropods,
evaluate higher-level diversity patterns of the fur-
cula in Aves, and where possible comment on
interspecific variation in these groups.
We restrict our descriptions as well as our phylo-
genetic discussion to nonavian theropod taxa with
unambiguously identified furculae. Description
and inclusion of all avialans into a phylogenetic
discussion is beyond the scope of this work. How-
ever, we also comment on variation within Aves.
We use birds as equivalent to Avialae as defined
by Gauthier (1986). The crown group name Aves is
used to refer to the living diversity as suggested
by Gauthier and de Queiroz (2001).
BACKGROUND
Anatomical Terminology
The furcula is a V or U -shaped midline bone
that can be divided into three anatomical regions;
the symphysis, the rami, and the epicleideal proc-
esses (Fig. 2). For clarity, the epicleideal processes
(or omal tip) will be considered ‘‘proximal’’ and the
symphyseal region as ‘‘distal.’’ The symphysis (5
synostosis interclavicularis 5 apophysis furculae
sensu Baumal and Witmer, 1993) is located at the
midline and represents the area of fusion of the
two primary clavicles. Some taxa including many
parrots, owls, Buceros, and Alcedo never ossify the
symphysis; instead, the symphysis is either carti-
lage or fibrous tissue (Newton, 1896; Baumel and
Witmer, 1993). The epicleidium (5 extremitas
omalis claviculae 5 extremital scapularis) refers to
the dorsal expansion of the furcula ramus proxi-
mally. The epicleideal processes attach to the acro-
857
Fig. 2. Terminology used to describe furculae with examples
from (A) Oviraptor philoceratops (AMNH FR 6517) and (B)
Bambiraptor feinbergorum (AMNH FR 30554).
mion process of the scapula through a ligamentous
joint. Most epicleideal processes thin as they
approach the articulation with the shoulder.
Although highly diverse in morphology, size, and
attachment to other pectoral elements, the observed
furculae always attach the epicleideal processes to
the acromion process of the scapula either through
direct contact or a ligamentous attachment. We
apply the term ramus to describe the shaft between
the epicleidium and the symphysis. The cross section
and three-dimensional architecture of the rami vary
considerably throughout the theropod tree.
Most avialans and some nonavialan dinosaurs
bear a nearly symmetrical ventrally projecting proc-
ess at the symphysis termed the hypocleidium (5
lamina interclavicularis). The shape of the hypoclei-
dium may be expressed as a rod, blade, or knob. The
hypocleidium contacts the sternum through either
direct contact or a ligamentous attachment (Baumel
and Witmer, 1993). The morphology of this feature
varies considerably within some relatively small
avian clades (e.g., procellariiforms) and throughout
the larger theropod tree (see below). Some basal
theropods (e.g., Coelophysis rhodesiensis) possess a
small tuber that displays intraspecific variation.
Topographic Connectivity of the Furcula
The ancestral archosaur pectoral girdle comprises
of paired, dorsally placed scapulae, paired ventrally
placed coracoids, an unpaired midline interclavicle,
Journal of Morphology
858 S.J. NESBITT ET AL.
and paired clavicles. Clavicles are present in the
archosaurian outgroups Lepidosauromorpha and
Sphenodontia. Within the pseudosuchian
archosaurs, the clavicle is absent in crocodylo-
morphs (Benton and Clark, 1988). Along the orni-
thodiran lineage of archosaurs, the interclavicle,
clavicle and sternum are fused in pterosaurs (Wild,
1993), whereas the clavicles are not known in Scle-
romochlus (Benton, 1999), or other dinosaurian out-
groups such as Silesaurus opolensis (Dzik, 2003)
and Marasuchus lilloensis (Sereno and Arcucci,
1994a). Given this absence of clavicles and intercla-
vicle in dinosaurian outgroups, the homology of the
furcula to other components of the shoulder girdle
has been contentious (Bryant and Russell, 1993).
This debate is based largely on absence of evidence
and will be explored more fully latter in this article.
Ancestrally in theropods, the only contact the
furcula has with the pectoral girdle is by way of
epicleidial contact with the acromion of the scap-
ula. Ornithothoraces (Enantiornithes 1 Aves) is
the basal-most clade where the furcula and the
coracoid contact by way of the prominent acrocora-
coid process (5 biceps tubercle or coracoid tuber-
cle), a derived condition within Avialae. This two-
part contact of the furcula with the coracoid and
scapula forms the triosseal canal. This canal trans-
mits the supracoracoideus ligment. In more derived
avialans (e.g., Gansus yumenensis), the furcula
sometimes makes a third contact with anterior
margin of the carina of the sternum (pers obs).
In extant avians, the furcula, and the fascia con-
necting it to the anterolateral sternum margin,
serves as the anteriormost attachment surface for
the sternobrachialis component of the pectoralis
muscles (Baumel and Raikow, 1993).
Furcula Development
Hall (2001) recently reviewed the development
of the avian clavicle. A short overview of its histo-
genesis will be presented here. The clavicles and
the bones of the mandible are the first bones to
appear during development. Intramembranous
osteogenesis is initiated at a single center of ossifi-
cation between Hamburger and Hamilton (H. H.)
stages 26 and 29, which corresponds to 5–6.5 days
of development. Based on the work by Hall (1986),
it is clear that condensation of preclavicular mes-
enchyme occurs at H. H. stage 31–32, initial osteo-
genesis at H. H. stage 33, and a rapid transitory
appearance then disappearance of secondary carti-
lage between H. H. stage 34–35.
The fused clavicles in avians are remnants of the
ancestral tetrapod dermal skeleton. Confirmation
that they are derived from dermal or membrane
bone has come from numerous studies (Bellairs and
Jenkin, 1960; Romanoff, 1960; Hamilton, 1965; Rus-
sell and Joffe, 1985; Hall, 1986). However, Cheval-
lier (1977) and Chevallier et al. (1977) demonstrated
Journal of Morphology
that the avian clavicle arises from somatopleural
mesoderm. This mesodermal origin of avian
clavicles conflicts with the dermal skeletal homology
of the clavicle, as it would be expected to be postotic
neural crest derived (Smith and Hall, 1990, 1993).
Hall (2001) suggested two possibilities to explain
the seemingly anomalous mesoderm-derived der-
mal bone. First, the clavicle is endoskeletal (endo-
chondral) and/or nonhomologous with the reptilian
clavicle. This hypothesis is clearly rejected by
numerous developmental studies confirming the
dermal homology of the clavicle (Bellairs and Jen-
kin, 1960; Romanoff, 1960; Hamilton, 1965; Rus-
sell and Joffe, 1985; Hall, 1986). Additionally, topo-
graphic connectivity of the furcula to the other
pectoral girdle elements in avian and other thero-
pod dinosaurs is entirely consistent with and sup-
portive of homology of the avian clavicle with the
ancestral reptilian and tetrapod clavicle.
The second hypothesis suggested by Hall (2001)
is that the avian clavicle is dermal, but with meso-
dermal contribution. This is supported by the
investment of lateral braincase elements by neural
crest cells (Schneider, 1999). As a result, Hall
(2001) proposed that the dermal/endochondral
(exoskeletal/endoskeletal) composition of an ele-
ment may not be strictly divided developmentally
along the neural crest/mesoderm line, but ‘‘may
reflect regional localization and allocation of migrat-
ing cell populations.’’ Indeed, Matsuoka et al. (2005)
recently showed the existence of cryptic boundaries
between cell types. Neural crest and mesoderm cells
obey a conserved muscle scaffold. Neural crest-
derived muscles anchor the head to the shoulder
girdle and Hox-gene controlled mesoderm link trunk
muscles to the posterior neck and shoulder skeleton.
These authors provide a ‘‘muscle scaffold model’’
alternative to the traditional ‘‘ossification model’’ for
inferring neck and shoulder homologies.
The traditional ‘‘ossification model’’ predicts that
the avian clavicle would be postotic neural crest-
derived because it is a dermal ossification. How-
ever, given the work of Chevallier (1977) and Che-
vallier et al. (1977) we know the avian clavicle
derives from the mesoderm. This mesodermal cel-
lular origin of the clavicle is precisely what the
‘‘muscle scaffold model’’ predicts, given that the
sternobrachialis division of the pectoralis muscles
attaches to the furcula. The sternobrachialis is
mesodermal in origin as illustrated by its enerva-
tion from branches of cranial nerves X, XI, XII.
Thus, avian furcula development provides corrobo-
rative evidence for the novel ‘‘muscle scaffold
model’’ proposed by Matsuoka et al. (2005). Addi-
tionally, the mesodermal cellular origin of the
avian furcula is identical to the mesodermal cellu-
lar origin of the posterior ossification center of the
mammalian clavicle, thus further supporting the
avian (and theropod) furcula and mammalian clav-
icle as homologues.
 THEROPOD FURCULA
Function of the Furcula
To date, only a few studies have examined the
function of the furcula. These studies have concen-
trated on its behavior during flight (Jenkins et al.,
1988; Goslow et al., 1990; Bailey and DeMont, 1991)
and the significance of furcula shape (Hui, 2002).
X-ray footage of European starlings (Sturnus
vulgaris) during flight suggests the furcula acts as
a spring (Jenkins et al., 1988). This study was re-
stricted to the small European starling because of
size restrictions in the wind tunnel where a cine-
radiographic system could record X-ray images. In
this footage, the furcula, scapula, and coracoid
bend laterally during the downstroke and restore
to their original positions after the upstroke. The
energy returned in the upstroke is negligible (Jen-
kins et al., 1988). The authors explicitly state that
these data were obtained exclusively from the Eu-
ropean starling and that further studies on other
birds on other birds are required to confirm the
result. Jenkins et al. (1988) and Goslow et al.
(1990) observed that the structural diversity of the
furculae among avians and suggested that there is
not universal utilization of the furcula as a spring.
The greatest cross-sectional area of the basal avia-
lian furcula is aligned transversely, and would not
likely flex like that of a starling (Jenkins et al.,
1988). On the basis of this, Jenkins et al. (1988)
hypothesized that the furculae of early avians
likely functioned as a strut or a brace.
Jenkins et al. (1988) found an additional possi-
ble role for the furcula–respiration. The rami of
the furcula surround the walls of the clavicular air
sac. Jenkins et al. (1988) demonstrated a coupling
between furcular flexion and sternal ascent and re-
traction. Their data indicate that the furcula may
have acted as a spring, and the sternum may have
acted as a pump, and this might represent a sec-
ondary respiratory pumping mechanism between
the air sacs and the lungs. Furthermore, Jenkins
et al. (1988) hypothesized that this system is capa-
ble of operating independently of inhalation and
exhalation and it could serve the increased meta-
bolic demands of flight (Goslow et al., 1990).
Bailey and DeMont (1991) tested the ideas of
Jenkins et al. (1988) and Goslow et al. (1990).
They found that: 1) 16 of 17 avians examined (see
Bailey and DeMont, 1991 for list of taxa) did not
store a significant amount of strain energy in the
furcula compared with the kinectic enery of the
wings; 2) the furcula is not important in panting;
and 3) the wingbeat frequency is a 1:1 ratio with
respiration in all 17 taxa examined. A 1:1 ratio
during respiration suggests that the furcula may
function in a resonant respiratory system operat-
ing during flight. This supports the original con-
clusions of Jenkins et al. (1988).
A recent paper by Hui (2002) suggests that the
morphology of the furcula (e.g., V-shaped versus
859
U-shaped and anterior-curved versus straight) cor-
relates with the flight requirements of different
birds. The canonical discriminate analysis placed
various birds into four general categories: soaring
with no flapping, flapping with no soaring, subaqu-
eous, and partially subaqueous. Hui (2002) found
soaring birds to have more of a U-shaped and
rounded furcula when compared with flapping
birds. This study concluded that a change in the
shape and curvature of the furcula alters the force
vector that the anterior sternobrachialis applies to
the wing and additionally that furcula shape
appears to be more strongly influenced by function
than phylogeny.
Clearly, much work remains to be done before
our understanding of furcula function is anything
close to comprehensive.
Pneumaticity in the Furcula
Aves is the only extant clade whose members
possess pneumatized postcranial skeletons
(Duncker, 1989). This pneumaticity is found in the
vertebrae, sternum, pelvis, humerus, femur, and
shoulder girdle (O’Connor, 2004). Pneumaticity
also occurs in a number of nonavian dinosaurs.
Pneumaticity has long been known to occur in the
vertebrae of several theropod taxa (O’Connar and
Claessens, 2005; Wedel, 2006) and in some sauro-
podomorphs (Wedel, 2003). Recently, pneumatized
or possibly pneumatized theropod furculae have
been discovered among some dromaeosaurids
(Makovicky et al., 2005).
The furcula of Buitreraptor gonzalezorum
(Makovicky et al., 2005), a dromaeosaurid from
the Cretaceous of Argentina, is highly pneuma-
tized, with trabeculae found throughout its inte-
rior as determined by CT scans. Bambiraptor fein-
bergorum (Burnham, 2004) might have pneumatic
furcula, indicated by a possible pneumatic foramen
found on either side of the synthesis. However, the
internal structure is rather solid and not open as
with Buitreraptor. Another recently discovered
pneumatized furcula is that of the basal tetanuran
Aerosteon riocoloradensis (Sereno et al., 2008). The
furcula of this specimen has a large slit-shaped
pneumatopore on the posterior-dorsal margin of
the bone, which leads to an undivided crescent
shaped pneumatocoel that extends proximally half-
way into each epicleideal process (Sereno et al.,
2008). This is in great contrast to the pneumatiza-
tion of other nonavialian theropods, and seems to
be highly apomorphic given its hypothesized phylo-
genetic position. Furthermore, in Buitreraptor the
pneumatopores are quite small and the bone itself
is highly pneumatized whereas the opposite is true
in A. riocoloradensis.
It is quite possible that other furculae are pneu-
mantic, especially those of other paravians. How-
ever, unless the furcula is broken and the internal
Journal of Morphology
 Fig. 3. Anterior views of avian furculae showing diversity of form (all specimens from the ornithology collection of the AMNH).
(A) Anas platyrhynchos 5847. (B) Phaethon rubicaudus 1203. (C) Chauna torquata 3616. (D) Mycteria americana 3768. (E) Sagit-
tarius serpentarius 4006. (F) Falco peregrinus 3394. (G) Corvus ossifragus 1050. (H) Morus serrator 5411. (I) Argusianus agrus
4969. (J) Cochlearius cochlearius 3494. (K) Diomedea epomorpha 1437. (L) Guttera plumifera 6415. Lateral view of avian furculae
showing diversity. (M) Anas platyrhynchos 5847. (N) Mycteria americana 3768. (O) Alca torda 5976. (P) Phaethon rubicaudus
1203. (Q) Corvus ossifragus 1050. (R) Mycteria americana 3768 (reversed), (S) Guttera plumifera 6415. (T) Argusianus agrus 4969.
(U) Falco peregrinus 3394.
 THEROPOD FURCULA
Fig. 3. (Continued)
structure is exposed, we must use external cues to
discern pneumatic furcula. The presence of pneu-
matic furcula outside tetanurans remains to be
shown. Many relatively large theropods such as
Tyrannosaurus rex commonly have broken furculae
that are clearly not pneumatized (MAN pers. obs.).
Taxa that have greatly pneumatized skeletons,
such as the oviraptorid Citipati osmolskae (Clark
et al., 2001) clearly do not have pneumatic furcula.
Flattening of the furcula as a result of crushing of
specimens may disguise pneumatic features of the
furcula.
Variation Across the Extant Orders
There is a large amount of variation present in
the furculae of Aves (Fig. 3). We surveyed this var-
iation in extant clades by inspecting furculae from
the AMNH Department of Ornithology collection
and referring to the character state distributions
of Livezey and Zusi (2006). Among extant birds,
ratites and many Psittaciformes lack fused
clavicles entirely. Those taxa that do possess furcu-
lae display a wide array of differing morphologies.
The most common condition of the epicleideal proc-
ess is one where its breadth is not significantly
larger than that of the ramus. The converse is
true for some groups of birds where the epicleideal
process is much wider than the ramus. Although
this occurs sporadically within many clades, such
861
as Galliformes, it reliably characterizes Falconi-
formes (Fig. 3U) (Livezey and Zusi, 2007). A trait
of the epicleideal morphology that is more consist-
ent in its distribution is the shape of the furcula at
the transition between the ramus and epicleideal
process. Most taxa lack a sharp transition between
these two regions, but in clades such as Sphenisci-
formes, Pelicaniformes (Fig. 3U), and Galliformes
(among others) the furcula is more of an inverted
L-shaped in profile, as the angle changes sharply
between the two structures. This sharp change in
structure is accompanied by a well-developed dor-
sal projection in Anseriformes (Fig. 3M). The epi-
cleideal process can also be markedly separated
from the ramus via a depression on the lateral
side of the ramus; this morphology appears pri-
marily in some Falconiformes (Fig. 3U), but it is
not confined to this clade. Pelicaniformes (Fig. 3H)
and most Falconiformes have epicleideal processes
where the facet for articulation with the coracoid
is set away from the main body of the furcula on a
‘‘pedestal.’’ In this same anatomical region the Pel-
icaniformes (and scattered other taxa such as
some Falconiformes) display a recess between the
‘‘pedestal’’ and main body of the furcula that con-
tains pneumatic pores. The epicleideal process can
be elongated, either along the same plane as the
ramus, such as in Galliformes and many Chara-
driiformes (Fig. 3O), or it can be elongated antero-
posteriorly. The latter morphology characterizes
Journal of Morphology
862 S.J. NESBITT ET AL.
most Passeriformes (Fig. 3Q), with a subset of this
clade possessing an epicedial process that is not
only widened but also bifurcated.
The ramus of the furcula is compressed medio-
laterally in most living birds. In some Galliformes
and in some taxa from numerous other clades, this
compression is absent with the furcula roughly cir-
cular in cross section. Most ramii arc laterally to
give the furcula a U-shape in anterior view. A
V-shaped furcula, however, is found predominantly
in Galliformes (Fig. 3I) and Strigiformes. In lateral
view, the ramus of the furcula typically has a
small amount of curvature. This curvature is lack-
ing in Anseriformes and Galliformes (Fig. 3T). It is
developed to a great degree in many Charadrii-
formes (Fig. 3O) and Procellariiformes and their
relatives. This subcircular morphology is also
found in isolated taxa in other areas of the avian
tree. A deep cavity is seen in the lateral wall of
the anseriform Chauna torquata (Fig. 3S), but this
feature does not characterize Anseriformes as a
whole, nor is exclusive to this clade.
The hypocleidium in living birds has a variety of
forms. Where the furcula joins the sternum, a
‘‘double lobed’’ articular surface is present in Pele-
caniformes (Fig. 3B), but this morphology is not
unique to the clade. Most taxa, such as the Anseri-
formes (Fig. 3A), have a poorly developed hypoclei-
dium, or none at all. A moderately developed hypo-
cleidium, noticeable but equal to or less than the
depth of the furcula at the symphysis, is present
in some clades, such as the Charadriiformes and is
found in some taxa in other clades (such as Passer-
iformes). An elongated hypocleidium, much longer
than the depth of the furcula at the symphysis
point, characterizes Galliformes. Some Galli-
formes, such as Guttera plumifera (Fig. 3S) pos-
sess a hollow pocket at the symphysis, forming a
scoop-shaped hypocleidium. An elongate hypoclei-
dium is also found in a subset of some other
clades, including Passeriformes. At the symphysis,
a dorsal projection of uncertain homology with the
hypocleidium is found in Ciconiiformes (Fig. 3J)
and Pelicaniformes.
On the basis of these observations, we suggest
that the morphological variation of avian furculae
is potentially phylogenetically useful. Variation
among furculae within clades may be useful for
differentiating species-level taxa. Much work
remains before the evolution of the avian furcula
and the usefulness of the furculae in phylogenetic
analyses may be realized.
MATERIALS AND METHODS
Furculae across Theropoda from various institutions (Table 1)
were selected to compare, contrast, and search for phylogenetic
characters. Most of the furculae were examined first-hand by
one of the authors; whereas a few of the specimens were
described from a cast or photograph supplemented by a partial
Journal of Morphology
or full description of the specimen in the literature. Many of
the specimens are described here for the first time whereas pre-
viously described furculae are further discussed and compared
with other theropods.
The Clavicle and Interclavicle in
Nontheropod Tetrapods
Clavicles are found in a number of tetrapods and are thought
to have originated as part of the dermal skeleton of early verte-
brates (see above). For the purposes of this review, we will start
with close relatives of the dinosaurs, early archosauriforms and
archosaurs. In the nonarchosaurian archosauriform, Euparkeria
capensis (SAM 5867) the right and left clavicles articulate with
an interclavicle (Ewer, 1965) as in many other diapsids. The
clavicles meet at the midline on their anteromedial edge, are
clearly separated by the interclavicle posteriorly, and the epiclei-
deal processes articulate to the acromion process of the scapula.
Additionally, the interclavicle attaches to both coracoids along
their medial edges. Basal members of the crocodile-line archo-
saurs (pseudosuchians), sister group to the ornithodirans, retain
this primitive pattern (e.g., Prestosuchus chiniquensis BSP 34).
The morphology of the clavicles and the presence or absence
of the interclavicle is poorly understood in the early ornithodir-
ans. Traditionally, pterosaurs have been considered to lack or
have had greatly reduced clavicles and interclavicles (Gauthier,
1986; Sereno, 1991). Recently, Wild (1993) has demonstrated
that the clavicles and the interclavicle are present but com-
pletely incorporated with the sternum into a single sternal ele-
ment in the basal pterosaur Eudimorphodon ranzii. This sug-
gests that clavicles and interclavicles are plesiomorphically
present in Ornithodira. Unfortunately, most of the pectoral gir-
dle is not preserved in known specimens of the basal dinosauro-
morphs, Lagerpeton chanarensis and Marasuchus lilloensis
(Sereno and Arcucci, 1994a,b). Thus, it is unclear if they retained
the interclavicle. Similarly, the nondinosaurian dinosauriform
Silesaurus opolensis (Dzik, 2003) fails to preserve unambiguous
clavicles or an interclavicle. Therefore, clavicles are unknown in
basal dinosauromorphs and the ancestral state for Dinosauria
remains ambiguous.
Ornithischians lack an interclavicle but retain two sepa-
rated clavicles (Osborn, 1924a; Brown and Schlaikjer, 1940;
Sternberg, 1951; Chinnery and Weishampel, 1998). Some early
sauropodomorphs retain clavicles (Huene, 1926; Galton, 2001).
Recently, Yates and Vasconcelos (2005) reported the presence
of articulated clavicles in the early sauropodomorph Masso-
spondylus carinatus. They found that the left and right
clavicles meet along their medial margin, but do not fuse. The
two reduced clavicles form a V-shape and articulate to the
acromion process of the scapula at their proximal ends (Yates
and Vasconcelos, 2005). The condition in Massospondylus
serves as the immediate outgroup condition for Theropoda and
polarizes character states pertinent to the origin and evolution
of the furcula.
Furcula in Theropod Dinosaurs
Coelophysoidea
Coelophysis bauri (Cope, 1887). Specimens: AMNH FR 30647
(Fig. 4A), AMNH FR 7223, NMMNH P-42353, NMMNH P-
42577, NMMNH P-42576, NMMNH P-46615, GR unnumbered.
Coelophysis bauri is represented by dozens of skeletons from
the famous Coelophysis Quarry in the Late Triassic of northern
New Mexico (Colbert, 1989, 1995). The presence of clavicles (or
a furcula) in C. bauri has been debated. Bryant and Russell
(1993) suggested that C. bauri had neither clavicles nor an
interclavicle because these structures had not been found in
any members of the ‘‘large sample.’’ However, at that time very
few Coelophysis skeletons had actually been fully prepared.
Padian (1997) reported that C. bauri has a clavicle without ref-
erence to a particular specimen. Recent preparation at the
AMNH, GR (Downs, 2000), and the NMMNH (Rinehart et al.,
 THEROPOD FURCULA 863
TABLE 1. Institutional abbreviations

AMNH American Museum of Natural History, New York, NY, U.S.A.

BSP Bayerische Staatssammlung für Paläontologie und historische Geologie, München, Germany
CAGS Chinese Academy of Geological Sciences, Beijing, China
CDPC China Dinosaur Park of Changzhou, Jiangsu, China
CMI The Children’s Museum, Indianapolis, ID, U.S.A.
CMN Canadian Museum of Nature, Ottawa, Canada
DINO Dinosaur National Monument, UT, U.S.A.
FMNH Field Museum of Natural History, Chicago, IL, U.S.A.
GR Ruth Hall Museum of Paleontology at Ghost Ranch, NM, U.S.A.
HYMV Heyuan Museum, Guangdong Province, China
IGM Mongolian Institute of Geology, Ulaan Bataar, Mongolia
IVPP Institute of Vertebrate Paleontology and Paleoanthropolgy Beijing, China
QG National Museum of Natural History, Bulawayo, Zimbabwe
LH1 Long Hao Geologic Paleontological Research Center, Department of Land and Resources, Hohhot, China
LH2 Unidad de Paleontologı́a, Departamento de Biologı́a, Facultad de Ciencias, Universidad Autónoma de Madrid, Spain
MACN-CH Museo Argentino de Ciencias Naturales ‘‘B. Rivadavia,’’ Coleccion Chubut, Argentina
MCF-PVPH Museo Carmen Funes, Paleontologı́a de Vertebrados, Plaza Huincul, Neuquén, Argentina
MCNA Museo de Ciencias Naturales y Antropológicas (J. C. Moyano) de Mendoza, Mendoza, Argentina
MNA Museum of Northern Arizona, Flagstaff, AZ, U.S.A.
MNN Musese National du Niger, Niamey, Republic of Niger
MOR Museum of the Rockies, Bozeman, MT, U.S.A.
MPCA Museo Carlos Ameghino, Cipolletti, Rio Negro Province, Argentina
NGMC National Geological Museum of China, Beijing, China
NMMNH New Mexico Museum of Natural History and Science, Albuquerque, NM, U.S.A.
SAM South African Museum, Cape Town, South Africa
SMM Sternberg Memorial Museum, Hays, KS, U.S.A.
TA Timescale Adventures Research and Interpretive Center, Bynum, MT, U.S.A.
TMP Royal Tyrrell Museum of Paleontology, Drumheller, Canada
UCMP University of California Museum of Paleontology, Berkeley, CA, U.S.A.
UCRC University of Chicago Research Collection, Chicago, IL, U.S.A.
UMNH Utah Museum of Natural History, Salt Lake City, UT, U.S.A.
UUVP University of Utah, Vertebrate Paleontology Collections, UT, U.S.A.
WDC-CSG Wyoming Dinosaur Center Thermopolis, WY, U.S.A.
YPM Yale Peabody Museum, New Haven, CT, U.S.A.
ZPAL Institute of Paleobiology of the Polish Academy of Sciences in Warsaw, Poland

2007) has revealed furculae disarticulated and articulated to
the pectoral girdle of C. bauri. This is the oldest known occur-
rence of the element (Rinehart et al., 2007).
The furcula of Coelophysis bauri is U-shaped, nearly round in
cross section, and the interclavicular angle varies from 115 to
1408. The epicleideal facets are unexpanded and have small lon-
gitudinal grooves. Breaks in the rami reveal dense, compact
bone. Rinehart et al. (2007) report that breaks near the end of
the epicedial process reveal a hollow cavity and the furcula and
scapulocoracoid grow in near isometry with respect to the hu-
merus. Some specimens of furculae of C. bauri have a small
ventrally directed process at the symphysis (NMMNH P-42577,
NMMNH P-42576, NMMNH P-44554) whereas other specimens
of C. bauri (NMMNH P-42353, NMMNH P-46615, AMNH
30647) do not. The small process is rounded in cross section,
terminates in a point ventrally, and is like those of C. rhode-
siensis and Allosaurus. It is unclear if this small process is ho-
mologous given the phylogenetic hypothesis presented below.
Coelophysis rhodesiensis (Raath, 1969). Specimens: QG 193
(CT6/EIII), QG 193 (CT6/E), QG 244 (CT6/F) (Fig. 4C), QG 244
(CT6/G).
Five furculae were found among disarticulated skeletons of
Coelophysis rhodesiensis (Tykoski et al., 2002). The morphol-
ogy of all the specimens is similar to those of C. bauri. The
furculae bow anteriorly, the interclavicular angle varies from
115 to 1408, the epicleideal facets are unexpanded, and the
rami are nearly round in cross section (Tykoski et al., 2002).
The shape of the furcula is U-shaped, more similar to coeluro-
saurs than to ‘‘Syntarsus’’ kayentakatae, Suchomimus, and
Allosaurus. Some specimens have a small tuber where the
hypocleidium would be located, whereas other specimens lack
any projection.
‘‘Syntarsus’’ kayentakatae (Rowe, 1989). Specimen: MNA
V2623 (Fig. 4B).
Recently, a nearly complete furcula was discovered articu-
lated with the right scapula in the only known specimen of
‘‘Syntarsus’’ kayentakatae. The right ramus of the furcula is
broken and the most proximal segment is missing (Tykoski
et al., 2002). It bows anteriorly in dorsal view, is V-shaped, and
has an interclavicular angle of 1408. It also has a small tuber
on the ventral portion of the symphysis (Tykoski et al., 2002),
as seen in some specimens of Coelophysis rhodesiensis, C. bauri,
and Segisaurus halli. The epicleideal processes are unexpanded
and flattened (Tykoski et al., 2002). Breaks in the nearly circu-
lar rami reveal dense bone.
Segisaurus halli (Camp, 1936). Specimen: UCMP 32101
(Fig. 4D).
In the original description of Segisaurus halli, Camp (1936)
reported the presence of a clavicle in articulation with the pecto-
ral girdle. Clavicles were unknown in other dinosaurs at the
time. This convinced Camp that S. halli was an unusual theropod
(Carrano et al., 2005). Identification of this element as a clavicle
followed by Bryant and Russell (1993) and Makovicky and Currie
(1998) resulted in the conclusion that the presence of a furcula
was a tetanuran synampomorphy. Recent repreparation and
study of S. halli demonstrated that Camp’s clavicle is actually a
broken furcula (Senter and Hutchinson, 2001; Tykoski et al.,
2002; Rauhut, 2003a; Carrano et al., 2005). The furcula of Segi-
saurus halli bears a small tuber at the symphysis, the intercla-
vicular angle is estimated at 1408, and the structure as a whole
may be asymmetric (Carrano et al., 2005). Additionally, the left
ramus has a striated, posterodorsally facing epicleideal process.
Abeliosauridae
Carnotaurus sastrei (Bonaparte, 1985). Specimen: MACN-
CH894.
Bonaparte et al. (1990) reported a small portion of a clavicle
associated with the pectoral girdle. Because the fragment is out

Journal of Morphology
864 S.J. NESBITT ET AL.

Fig. 4. The furculae of non coelurosaurian theropods. (A) Coelophysis bauri (AMNH FR 30647) in anterior view. (B) ‘‘Syntarsus’’
kayentakatae (MNA V2623) in anterior view. (C) Coelophysis rhodesiensis (top CT6.EIII; bottom CT6/G) from Tykoski et al. (2002).
(D) Segisaurus halli (UCMP 32101) in anterolateral view. (E) Suchomimus tenerensis (MNN GAD513) in anterior view. (F) Allo-
saurus (DINO 11541) in anterior view from Chure and Madsen (1996). Relationships based on Smith et al. (2007). Scales 5 1 cm
in A-D. Scales 5 5 cm in E and F.

of articulation and bears no identifiable features of a clavicle or Allosauroidea

furcula, it is not considered further.
Spinosauroidea
Suchomimus tenerensis (Sereno et al., 1998). Specimen:
MNN GDF500 (Fig. 4E).
The furcula of Suchomimus tenerensis represents the only
spinosaurid furcula recovered to date (Lipkin et al., 2007). The
complete and well preserved furcula is V-shaped with an inter-
clavicular of 1118, a D-shaped cross section where the anterior
surface is convex and the posterior face is flat. The ventral por-
tion of the epicleideal processes bear grooves and taper to a
point (Lipkin et al., 2007). The symphysis bears an asymmetri-
cal ventrally directed tuber (Lipkin et al., 2007). The ventral
tuber is similar to that of Allosaurus fragilis and coelophysoids.
As noted by Lipkin and Sereno (2002) the furcula, though less
robust, resembles the furcula of A. fragilis.
Allosaurus (Marsh, 1877). Specimens: Allosaurus sp. DINO
11541 (Fig. 2 of Chure and Madsen, 1996); Allosaurus fragilis
UUVP 6102, UUVP 6100, UUVP 6101, UUVP 387, UUVP 5754,
UUVP 11690, UUVP 5753, UUVP 6132 (Figs. B-I of Chure and
Madsen, 1996). (Fig. 4F).
Chure and Madsen (1996) identified an articulated furcula in
an undescribed new taxon of Allosaurus. The well-preserved
specimen is V-shaped, has an interclavicular angle of 1308, and
has a small tuber on the ventral side of the symphysis. It is
rather similar to those of coelophysoids. Chure and Madsen
(1996) also discovered eight additional furculae from the Cleve-
land-Lloyd Dinosaur Quarry that they assign to Allosaurus fragi-
lis. All eight furculae are rather similar to DINO 11541; however,
they show the following variation: 1) The small tuber on the ven-
tral portion of the symphysis in some specimens is absent in
others; 2) the epicleideal articular facets may be raised or unde-

Journal of Morphology
 THEROPOD FURCULA 865

Fig. 5. The furculae of tyrannosaurids and other basal coelurosaurs. (A) Tyrannosaurus rex (MOR 1125) in anterior view.
(B) Albertosaurus sarcophagus (TMP 86.64.1) in anterior view. (C) Daspltosaurus torosus (CMN 8056) in anterior view. (D) Gorgo-
saurus libratus (TMP 91.36.500) in anterior view. (E) Gorgosaurus libratus (TMP 94.12.602) in anterior view. (F) Huaxiagnathus
orientalis (CAGS-IG02-301) in anterior view (from Hwang et al., 2004). Relationships based on Currie et al. (2003) and the latest
TWiG matrix in Turner et al. (2007a). The relationships of UUVP 11689 is not known. Scale 5 5 cm in A-F. Scale 5 1 cm in G.

fined, and striated or smooth; and 3) grooves are present along
the rami in one of the specimens on the posterior side (Chure and
Madsen, 1996). Additionally, the cross sections of the rami vary
from round to nearly flat within the same furcula (the furcula is
coded as having a nearly round cross section because most of the
cross sections are rounded). The variation found here serves to
illuminate potential complications in the construction of phyloge-
netic characters. Therefore, characters listed below have been
constructed to take the variability into account.
Aerosteon riocoloradensis (Sereno et al., 2008). Specimen:
MCNA-PV-3137 (Fig. 11 of Sereno et al., 2008).
Recently, Sereno et al. (2008) described the unique, well-pre-
served furcula from the allosauriod Aerosteon riocoloradensis.
The furcula is quite similar to that of Allosaurus, although the
furcula of A. riocoloradensis is more anteroposteriorly com-
pressed (Fig. 11 of Sereno et al., 2008). The general V-shape,
1208 interclavicular angle, and tapering epicleideal processes
rank among the usual features of a basal tetanuran whereas a
deep pneumatocoel at the symphasis is unique among nonorni-
thuran theropods (Sereno et al., 2008).
Carcharodontosauridae
Mapusaurus roseae (Coria and Currie, 2006). Specimen:
MCF-PVPH-108.116 (Fig. 21 of Coria and Currie, 2006).
Coria and Currie (2006) report a potential furcula found
among the seven partial skeletons of Mapusaurus roseae. The
specimen was not found articulated and the authors suggest
that the bone may be a may be a furcula rather than a fused
gastralium based on 1) sigmoidal curvature of the shaft, 2) sym-
metry of the bone and 3) the lack of a joint of the line of fusion
in the middle of the element. Overall, the general shape (V-
shaped, tapering epicleideal processes) of the element agrees
with the furculae of closely related taxa such as Allosaurus.
The symphysis of the element is somewhat distally expanded
like that of a fused gastralia; however, this could be an autapo-
morphy of Mapusaurus. Because that specimen was not found
articulated and is difficult to differentiate from fused gastralia,
the specimen cannot be unambiguously assigned to a furcula.
This specimen is not considered further.
Compsognathidae
Huaxiagnathus orientalis (Hwang et al., 2004). Specimen:
CAGS-IG02-301 (Fig. 5F).
A small furcula was discovered in near articulation in the only
known specimen of Huaxiagnathus orientalis. It is round in cross
section, does not bear a hypocleidium, has an interclavicular
angle of 1308, and is U-shaped (Hwang et al., 2004). Hwang et al.
(2004) noted that the specimen is similar to tyrannosaurids. The
epicleideal processes, however, do not arc laterally as they do in
tyrannosaurids (Makovicky and Currie, 1998).
Coelurosauria
Scipionyx samniticus (Dal Sasso and Signore, 1998). Speci-
men: Soprintendenza Archeologica, Salerno unnumbered.
A well-preserved furcula was found in near articulation in
the holotype of Scipionyx samniticus. The furcula was not

Journal of Morphology
866 S.J. NESBITT ET AL.
described, but bears the following characteristics: U-shaped,
estimated interclavicular angle of 1408, and a small ventrally
directed at the symphysis. It is unclear what the epicleideal
processes ends look like, but it appears that the processes are
flexed slightly laterally like that of tyrannosaurids.
Protarchaeopteryx robusta (Ji and Ji, 1997). Specimen:
NGMC 2125.
A small articulated furcula is present in Protachaeopteryx
robusta. It is described as U-shaped and has an interclavicular
angle near 608 (Ji et al., 1998). We have not observed this speci-
men firsthand and the illustration shows little detail.
Tyrannosauridae
Tyrannosaurus rex (Osborn, 1905). Specimens: UCRC V1
(Fig. 3 of Lipkin et al., 2007), MOR 980 (Fig. 4 of Lipkin et al.,
2007), MOR 1125 (Fig. 5A).
Brochu (2003) identified a small piece of bone as a potential
furcula in his detailed description of Tyrannosaurus rex (FMNH
PR2081). The potential furcula is V-shaped and lacks a small
tuber on the ventral surface of the symphysis. The bone is simi-
lar to the furculae of other tyrannosaurids, but cannot be
clearly differentiated from a gastral segment (Brochu, 2003).
Larson and Rigby (2005) argue, without much justification, that
the ‘‘furcula’’ from FMNH PR2081 represents a pathologic gas-
tralium. Larson and Rigby (2005) then identified another bone
as the furcula of FMNH PR2081 (Larson and Rigby, 2005; Fig
12.5). Brochu (2003), however, had already identified this bone
as a segment of the last rib of the trunk vertebrae. It is highly
asymmetrical and does not bear an articular surface on the pro-
posed epicleideal processes. Larson and Rigby (2005) also iden-
tify a furcula in another specimen of T. rex (CMI 2001.90.1).
This element is identical to the element identified as the furcula
in FMNH PR2081 (Larson and Rigby, 2005).
Lipkin et al. (2007) report an unambiguous furcula in articu-
lation for Tyrannosaurus rex (UCRC V1) from the Lance Forma-
tion of Wyoming. The U-shaped furcula has an interclavicular
angle of 718 (Lipkin et al., 2007). The inside of the furcula is
solid and there is no hypocleidial process (Lipkin et al., 2007).
The articulated UCRC V1 allowed Lipkin et al. (2007) to assign
two more furculae (MOR 980, MOR 1125) to T. rex. None of
these specimens possess a hypocleidium and all have intercla-
vicular angles ranging from 718 to 1138 (Lipkin et al., 2007).
The furculae of MOR 980 and MOR 1125 possess expanded
and slightly lateral arcing epicleideal process like those furculae
of Albertosaurus sarcophagus, Gorgosaurus libratus, and Das-
pletosaurus. Moreover, the presence of expanded and slightly
lateral arcing epicleideal process is apparently apomorphic in
tyrannosaurids. Additionally, the furculae of Albertosaurus,
Gorgosaurus, Daspltosaurus, and Tyrannosaurus rex (MOR 980,
MOR 1125, UCRC V1) are all nearly symmetrical. The absence
of these characters in the proposed furculae of T. rex (FMNH
PR2081 and CMI 2001.90.1) by Larson and Rigby (2005) sug-
gest that Brochu’s (2003) original identification of this element
as the last dorsal rib is correct.
Albertosaurus sarcophagus (Osborn, 1905). Specimen: TMP
86.64.1 (Fig. 5B).
Gorgosaurus libratus (Lambe, 1917). Specimens: TMP
94.12.602 (Fig. 5E), TMP 91.36.500 (Fig. 5D), CMI 2001.89.01.
Daspletosaurus torosus (Russell, 1970). Specimens: CMN
11315 (Fig. 1G,H of Makovicky and Currie, 1998), CMN 8056
(Fig. 5C).
Dapletosaurus. Specimen: TA.1997.002.710 (Fig. 16.8 of Cur-
rie et al., 2005)
Makovicky and Currie (1998) identified and described the fur-
culae of the Late Cretaceous tyrannosaurids Albertosaurus sar-
cophagus, Gorgosaurus libratus, and Daspletosaurus. These
three closely related taxa have similar furculae and will be con-
sidered together. In all of the taxa, the furcula is U-shaped and
the epicleideal processes arc laterally (Makovicky and Currie,
1998). Breaks in the rami reveal dense bone. The interclavicular
angle ranges from 95 to 1128 and the epicleideal process expands
dorsoventrally (Makovicky and Currie, 1998). This expansion
may be apomorphic for tyrannosaurids. Small ventrally directed
Journal of Morphology
tubera are present at the symphysis. Makovicky and Currie
(1998) suggested that the greater interclavicular angle and less
pronounced curvature of the furcula of Daspletosaurus could be
used to differentiate it from Gorgosaurus. This difference in Das-
pletosaurus is supported by an isolated furcula found among Des-
pletosaurus elements cranial and postcranial elements (Currie
et al., 2005). All of the furculae are round in cross section.
Specimen: UUVP 11689 (Fig. 22 of Madsen, 1976a).
Chure and Madsen (1996) discuss UUVP 11689, a furcula
found isolated within the Cleveland-Lloyd Quarry, but do not
assign the specimen to a specific taxon. Marshosaurus bicentesi-
mus, an enigmatic theropod (Madsen, 1976b), Ceratosaurus, a
neoceratosaur (Gilmore, 1920; Madsen and Welles, 2000), and
Stokesosaurus clevelandi, a tyrannosauroid (Rauhut, 2003b) all
occur with abundant Allosaurus skeletons in the quarry (Foster,
2007). UUVP 11689 is U-shaped, and has an expansion of the epi-
cleideal processes that is consistent with tyrannosauroids. There-
fore, the furcula may belong to S. clevelandi. However, more ma-
terial of S. clevelandi must be found to test this hypothesis. As of
2-26-08, this specimen could not be located at UUVP.
Therizinosauroidea
Beipiaosaurus inexpectus (Xu et al., 1999a). Specimen: IVPP
V11559 (Fig. 6E).
Beipiaosaurus inexpectus, from the Early Cretaceous of China
is the first therizinosauroid to preserve a furcula. The articu-
lated furcula has been only partially recovered, but an impres-
sion of the missing portions remains (Xu et al., 1999a). The dor-
soventrally expanded furcula has a very wide interclavicular
angle, nearly 1608. The impression also indicates a hypoclei-
dium was not present and the furcula was oval in cross section
(Xu et al., 1999a).
Falcarius utahensis (Kirkland et al., 2005). Specimen: UMNH-
VP 14671 (Fig. 6F).
A nearly complete furcula was found among the remains of
the basal therizinosaurid Falcarius utahensis (Kirkland et al.,
2005). The well preserved furcula is missing the epicleideal
processes. The ramus and symphysis is oval (long axis oriented
dorsoventrally) in cross section. A small nub is present on the
ventral side of the symphysis. This is similar to that of Allosau-
rus, Suchomimus tenerensis, and coelophysoids.
Neimongosaurus yangi (Zhang et al., 2001). Specimen: LH1
V0001 (Fig. 6G).
The well preserved furcula of Neimongosaurus yangi is robust
and the rami diverge at an angle at approximately 1358 (Zhang
et al., 2001). There is no hypocleidium (Zhang et al., 2001), and
the epicleideal processes terminate in blunt points. The sym-
physis area is dorsoventrally expanded relative to the rami.
The high interclavicular angle, blunt epicleideal processes, and
robust rami are similar to that of Beipiaosaurus inexpectus, a
character combination unique to these two therizinosauroids.
Given that B. inexpectus is a basal therizinosauroid and N.
yangi is one of the most derived taxa according to Zanno (2006),
the characters that these two taxa share suggest that they may
be found in most therizinosaurids. As with the oviraptorosaur-
ids, the furcula of a clade containing N. yangi and B. inexpectus
seem to be easily recongnizable.
Oviraptorsauria
Caudipteryx zoui (Ji et al., 1998). Specimen: NGMC 97-4-A.
The furcula of Caudipteryx zoui was briefly described as U-
shaped (Ji et al., 1998). The two additionally specimens of C.
zoui (BPM 1001 and IVPP V12430; Zhou et al., 2000) provide
little additional information on the furcula as the elements are
preserved only in lateral profile.
Oviraptor philoceratops (Osborn, 1924b). Specimen: AMNH
FR 6517 (Fig. 6B).
The holotype of Oviraptor philoceratops preserves a nearly
complete furcula in anterior view. Osborn (1924b) originally
described this element as an interclavicle. Just 2 years later,
Heilmann (1926) rejected the close relationship of birds and
theropods because of the absence of clavicles in dinosaurs
(Makovicky and Currie, 1998). A reinterpretation by Barsbold
 THEROPOD FURCULA 867

Fig. 6. Furculae of oviraptorosaurids and therizinisauroids. (A) Citipati osmolskae (IGM 100/978) in anterior view (top) and ven-
tral view (bottom). (B) Oviraptor philoceratops (AMNH FR 6517) in anterior view. (C) Khaan mckennai (IGM 100/1127) in anterior
view. (D) Ingenia yanshini (IGM 100/32) in anterior view. (E) Beipiaosaurus inexpectus (IVPP V11559) in anterior or posterior
view. (F) Falcarius utahensis (UMNH-VP 14671) in anterior view. (G) Neimongosaurus yangi (LH1 V0001) in anterior view. Ovirap-
torsaurid relationships from Xu et al. (2007), therizinosaurids relationships taken from Zanno (2006), and larger clade relationships
from the latest TWiG matrix in Turner et al. (2007a). Scales 5 1 cm.

(1983) concluded that Osborn’s interclavicle is actually a fur-
cula. The furcula is nearly complete with a U-shaped outline
and is thick at the symphysis. It has an interclavicular angle of
approximately 908, is rounded in cross section, and preserves
an elongate spike-like hypocleidium (Barsbold, 1983). In con-
trast to more basal theropod furculae, the furcula of O. philo-
ceratops possesses a symmetrically developed and elongate
hypocleidium. These two characters also are present in other
oviraptorids. The hypocleidium tapers ventrally and bears a dis-
tinct mid-line keel, a character not present in other oviraptor-
saurs. Additionally, the rami expand laterally between the epi-
cleideal process and the hypocleidium. As in other oviraptor-
saurs, the epicleidial processes are very long and share a large
overlap with the acromion process of the scapula.
Citipati osmolskae (Clark et al., 2001). Specimen: IGM 100/
978 (Fig. 6A).
Citipati osmolskae is known from a nearly complete skeleton
that includes an articulated pectoral region (Clark et al., 2001).
The complete furcula has the following suite of characters, oval
in cross section, U-shaped, elongated tapering epicleideal proc-
esses, a bulge of the rami between the symphyseal region and
the epicleideal process, and an elongated hypocleidium. The
elongated hypocleidium projects posteroventrally and tapers to
a point. In contrast with Oviraptor philoceratops, there is no
midline keel on the anterior surface of the hypocleidium. The
distinct bulge of the rami between the hypocleidium area and
the epicleideal process is shared by other oviraptorsaurs. The
distal tip of the hypocleidium articulates with the sternal plates
as has been inferred for Heyuannia huangi (Lü et al., 2005)
and O. philoceratops (Barsbold, 1983).
The epicleideal processes taper to point and articulate on the
medial sides of the scapulae. This is in contrast with the furcu-
lae of nonoviraptorsaurs, which lie on the anterior surface of
the acromion process of the scapula.
Khaan mckennai (Clark et al., 2001). Specimens: IGM 100/
1127 (Fig. 6C), IGM 100/1002.
Well-preserved nonarticulated furculae are present in the hol-
otype (IGM 100/1127) and paratype (IGM 100/1002) of Khaan
mckennai. Both share the following characteristics; U-shaped,
circular in cross section, an interclavicular angle near 908, and
are laterally expansion between the epicleideal process and the
hypocleidium. The holotype preserves a posteriorly directed
hypocleidium whereas the furcula of the paratype is not fully
prepared and as such the hypocleidium is obscured. The hypo-
cleidium is spike-like but does not bear the keel present in
Oviraptor philoceratops.

Journal of Morphology
868 S.J. NESBITT ET AL.

Fig. 7. The furculae of deinonychosaurs. (A) The articulated pectoral girdle in anterior view of Sinornithoides youngi (IVPP
V9612) with arrows highlighting the epicleideal processes. (B) Mei long (IVPP V12733) in anterior view. (C) Buitreraptor gonzalezo-
rum (MPCA 245) in anterior view. (D) Sinornithosaurus millenii (NGMC 91) in anterior view. The arrow indicates the symphasis
of the furcula. (E) Microraptor zhaoianus (IVPP unnumbered). In anterior view. (F) Bambiraptor feinbergorum (AMNH FR 30554)
in anterior view. (G) Velociraptor mongoliensis IGM 100/976 in anterior view from Norell et al. (1997). Relationships after Turner
et al. (2007ab). Scales 5 1 cm.

Heyuannia huangi (Lü, 2002). Specimen: HYMV1-2 (Fig. 13.1 Troodontidae

of Lü et al., 2005).
A well-preserved furcula remains in articulation in the holo-
type of Heyuannia huangi. The furcula is U-shaped, oval in
cross section, has an interclavicular angle of 908, and bears a
posteroventrally directed hypocleidium (Lü et al., 2005). The
hypocleidium is smooth and ends in a point as in Khaan mcken-
nai and Oviraptor philoceratops. The ramus between the epi-
cleideal process and the hypocleidium is laterally expanded as
with all the other oviraptorsaurs.
Ingenia yanshini (Barsbold, 1981). Specimen: IGM 100/32
(Fig. 6D).
The holotype skeleton of Ingenia yanshini preserves a furcula
that has been prepared three dimensionally. The gently U-
shaped furcula bears an oval-shaped cross section. The postero-
ventrally directed hypocleidium tapers at its termination and is
similar to that of other oviraptorsaurs. As with other oviraptor-
saurs, the ramus expands laterally. The epicleideal processes
taper to a point and it appears that the furcula articulates with
the medial side of the scapula as with other oviraptorsaurs.
Mei long (Xu and Norell, 2004). Specimen: IVPP V12733
(Fig. 7B).
A furcula is present in articulation in the well-preserved com-
plete skeleton of Mei long. It is described as robust, flattened in
cross section with a small hypocleidium and expanded epicleideal
processes (Xu and Norell, 2004). The furcula is clearly U-shaped,
but the epicleideal processes seem not to be expanded much more
than those of dromaeosaurids (Norell and Makovicky, 1999) or
Archaeopteryx lithographica (Mayr et al., 2007). The ventral
expansion (described as a hypocleidium) does not have the typical
features present in birds. It is not clear if the there is a ventral
ridge as with Buitreraptor gonzalezorum (MPCA 245).
Sinornithoides youngi (Russell and Dong, 1993). Specimen:
IVPP V9612 (Fig. 7A).
Russell and Dong (1993) described and figured a small por-
tion of a clavicle in articulation in the holotype skeleton of
Sinornithoides youngi (Fig. 7A). They concluded that this frag-
ment represented a clavicle and not a furcula. Bryant and Rus-
sell (1993) accepted this observation in their review and con-

Journal of Morphology
 THEROPOD FURCULA
cluded that troodontids did not have furculae. Tykoski et al.
(2002) questioned this conclusion and suggested that the speci-
men seems to be broken, yet they still concluded that it is not
clear whether S. youngi had a pair of clavicles or a furcula. If a
paired clavicle were present, it would be the only documented
case within Theropoda (Tykoski et al., 2002).
Reexamination of IVPP V9612 in the winter of 2007 indicates
that a complete furcula was present and that the specimen was
broken, probably during preparation or excavation (Fig. 7A). In
contrast to the furcula of Mei long, the furcula of Sinornithoides
youngi is very delicate, with thin rami.
Dromaeosauridae
Velociraptor mongoliensis (Osborn, 1924b). Specimen: IGM
100/976 (Fig. 7G), IGM 100/982.
Two of the Velociraptor mongoliensis specimens (IGM 100/
976, IGM 100/982) preserve furculae. The well-preserved fur-
cula of IGM 100/976 remains articulated (Norell et al., 1997).
The furcula is U-shaped, has an interclavicular angle of 1058,
and preserves a small tuber on the ventral side of the symphy-
sis. The furcular rami are round in cross section. The epiclei-
deal processes are unexpanded and continuous with the rami.
Only the right ramus and epicleideal process of the furcula of
IGM 100/982 is preserved. The morphology is identical to that
of IGM 100/976. The furcula of V. mongoliensis differs from all
of its close relatives and resembles the furculae of coelophysoids
and Allosaurus. The only major difference between these taxa
is that the furcula of V. mongoliensis is proportionally much
larger relative to its body size.
Sinornithosaurus millenii (Xu et al., 1999b). Specimen: IVPP
V12811 (Fig. 2 of Xu et al., 1999b), NGMC 91 (Fig. 7D).
Half of a well-preserved furcula is visible in the holotype of
Sinornithosaurus millenii. The furcula is U-shaped, has an esti-
mated interclavicular angle of 808, and has an anteroposter-
iorly compressed cross section (Xu et al., 1999b). The epicleideal
processes are unexpanded and continuous with the rami as
with Bambiraptor feinbergi (AMNH FR 30554) and Archaeop-
teryx lithographica (Mayr et al., 2007). The furculae of B. fein-
bergorum and S. millenii share nearly identical morphology.
Bambiraptor feinbergorum (Burnham et al., 2000). Specimen:
AMNH FR 30554 (Fig. 7F).
A well-preserved disarticulated furcula was discovered in the
nearly complete skeleton(s) of Bambiraptor feinbergorum. The
furcula is quite similar to that of Archaeopteryx lithographica
and other early avialans, but is quite different from that of
Velociraptor mongoliensis. It is U-shaped, has an anteroposter-
iorly compressed cross section, bears no tuber or hypocleidium,
and has an interclavicular angle near 808 (AMNH FR 30554;
Burnham, 2004). Two small foramina equidistant from the sym-
physis of the furcula may be autapomorphic for B. feinbergo-
rum. The epicleideal processes are unexpanded, continuous
with the rami, and have poorly defined articular surfaces.
Buitreraptor gonzalezorum (Makovicky et al., 2005). Speci-
men: MPCA 245 (Fig. 7C).
Recently, a well-preserved furcula was found among the
remains of Buitreraptor gonzalezorum. The furcula has posteri-
orly arcing rami and no hypocleideum (MPCA 245; Makovicky
et al., 2005). A small ridge is present at the ventral margin near
the intersection of the rami. The inside is hollow with extremely
thin trabeculae. These character states suggest that the furcula
may have been pneumatized (see above). Additionally, the furcula
is U-shaped, has an interclavicular angle of 808, the cross sec-
tion is circular, and it is slightly asymmetrical. The epicleideal
processes are unexpanded and continuous with the rami. The fur-
cula preserves a three-dimensional shape typical of Aves. This
suggests that other paravian furculae may have a similar shape
but are crushed or distorted during fossilization (see Discussion).
Microraptor zhaoianus (Xu et al., 2000). Specimens: CAGS
20-8-001 (Fig. 19a of Hwang et al., 2002), CAGS 20-7-004 (Fig.
19b of Hwang et al., 2002), IVPP unnumbered (Fig. 7E).
Two specimens of Microraptor zhaoianus preserve the fur-
cula in anterior view. Like other dromaeosaurids, the furculae
have the following characteristics; U-shaped, anteroposteriorly
869
compressed, and rounded epicleideal processes. The intercla-
vicular angle varies slightly among the specimens; CAGS 20-
8-001 has an interclavicular angle of 918 and CAGS 20-7-004
has an interclavicular angle of 828 (Hwang et al., 2002). The
furcula of M. zhaoianus is very similar to that of other para-
vians.
Avialae
Archaeopteryx lithographica (von Meyer, 1861). Specimens:
BMNH 37001 (Fig. 8A), WDC-CSG-100 (Fig. 1 of Mayr et al.,
2007).
Two specimens of Archaeopteryx lithographica (BMNH 37001,
WDC-CSG-100) preserve furculae in anterior view. The furcula
is robust, anteroposteriorly compressed, U-shaped, and lacks a
hypocleidium (Ostrom, 1976). The furculae are very similar to
those of other early avialans.
Sapeornis chaoyangensis (Zhou and Zhang, 2002a). Speci-
mens: IVPP V13275 (Fig. 8C), IVPP V13276 (Fig. 8B), IVPP
V12675 (Fig. 6 of Zhou and Zhang, 2002a), IVPP 13276 (Fig. 6
Zhou and Zhang, 2002a).
Well-preserved furculae are well represented in the many
specimens of Sapeornis chaoyangensis. The rather robust rami
and epicleideal processes are anteroposteriorly compressed, thus
creating an oval cross section. The U-shaped furcula bears a ven-
trally tapering hypocleidal process much like that of enantiorni-
thine birds (Zhou and Zhang, 2003). The symmetrical and elon-
gated hypocleidal process terminates in a blunt end in IVPP
V13276. Zhou and Zhang (2003) report an interclavicular angle
of 1058 and note that the furcula of S. chaoyangensis (without the
hypocleidium) closely resembles the furculae of Archaeopteryx
lithographica, Jeholornis prima, and Confuciusornis sanctus.
Jeholornis prima (Zhou and Zhang, 2002b). Specimen: IVPP
V13274 (Fig. 8D).
The furcula of this avialan is preserved in the holotype in
ventral view. The left side of the furcula is well preserved
whereas the right half is partially weathered away and covered
by the displaced scapula. The epicleideal process on the right
side is, however, exposed and well preserved. Like that of other
basal avialans the furcula of Jeholornis prima is robust and U-
shaped. It is compressed anteroposteriorly and, although poorly
preserved distally in the holotype, a hypocleidium is lacking.
The furcula rami are generally parallel-sided but narrow
slightly at the epicleideal facets.
Jixiangornis orientalis (Ji et al., 2002). Specimen: CDPC-02-
04-001 (Fig. 8E).
The furcula of this basal avialan is well preserved but exposed
only in ventral or posterior view. Like Archaeopteryx lithograph-
ica, Jeholornis orientalis, Confuciusornis sanctus, and derived
paravians such as Mei long and Sinornithosaurus milleni, the fur-
cula of J. orientalis is large, robust, and strongly U-shaped. Also
like these taxa, J. orientalis posseses a furcula that appears to be
anteroposteriorly compressed and lacks a hypocledium. A shallow
sulcus runs alongside the dorsal edge of the rami of the furcula on
the ventral surface. Additionally, the furcula rami expand slightly
proximally a trait otherwise only seen in oviraptorsaurs.
Confuciusornis sanctus (Hou et al., 1995). Specimen: GMV-
2133 (Fig. 8H), GMV-2132, GMV-2131.
There are many well-preserved samples of furculae in Confu-
ciusornis. They are all U-shaped, robust, have an interclavicu-
lar angle of about 858. They are anteroposteriorly compressed
in cross section and lack a hypocleidium. However, some of the
specimens have a tiny tuber on the ventral edge where the
hypocleidium would be located (Chiappe et al., 1999). The epi-
cleideal processes are rounded and slightly offset from the
ramus.
Changchengornis hengdaoziensis (Ji et al., 1999). Specimen:
GMV-21290 (Fig. 54 of Chiappe et al., 1999).
A well-preserved furcula is present in the holotype and only
known specimen of Changchengornis hengdaoziensis. The furcula
is robust, has an interclavicular angle of about 858, U-shaped,
oval in cross section, and lacks a hypocleidium (Chiappe et al.,
1999). A small tubercle is located on the anterior side of the
Journal of Morphology
870 S.J. NESBITT ET AL.

Fig. 8. Furculae of basal avialians. (A) Archaeopteryx lithographica (BMNH 37001) in anterior view. (B) Sapeornis chaoyangen-
sis (IVPP V13396) in anterior or posterior view. (C) Sapeornis chaoyangensis (IVPP unnumbered) in anterior view. (D) Jeholornis
(IVPP V13274) in anterior or posterior view. Arrow highlights the epicleideal process. (E) Jixiangornis orientalis (CDPC-02-04-001)
in anterior view. Arrow highlights the epicleideal process. (F) Iberomesornis romerali (cast of LH2-022R) in anterior or posterior
view. (G) Concornis lacustris (LH2-2814) in anterior view. (H) Confuciusornis sanctus (GMV-2133) in anterior view. Relationships
after Turner et al. (2007ab) and Clarke et al. (2006). Scales 5 1 cm in A–E, H. Scale 5 1 mm in F. Scale 5 5 mm in G.

symphysis (Chiappe et al., 1999). Chiappe et al. (1999) reported
that small furrows are present on the ramus on both the ante-
rior and posterior sides. These furrows may be the result of
crushing. The furcula is similar to that of Archaeopteryx lithog-
raphica and Confuciusornis sanctus.
Iberomesornis romerali (Sanz and Bonaparte, 1992). Speci-
men: LH2-022R (Fig. 8F).
The furcula in the only known specimen of this taxon is well
preserved and exposed in anterior view. The proximal half of
the epicleideal processes are partially abraded, but their com-
plete shape is not lost. The interclavicular angle is roughly 608,
which translates into an intermediate shape between a strictly
U-shaped or V-shaped furcula, although like most enantorni-
thines it is closer to a V-shape. The epicleideal process is best
preserved on the left ramus. It is flattened mediolaterally to
form its contact facet with the coracoid. The furcular rami are
slightly dorsoventrally convex and thin toward the hypoclei-
dium. The hypocleidium is well-developed, triangular in ante-
rior profile, and extends ventrally roughly one-third the length
of the furcula body.
Concornis lacustris (Sanz and Buscalioni, 1992). Specimen:
LH2-2814 (Fig. 8G).
The furcula of Concornis lacustris is elongate, with narrow
rami as in Iberomesornis romerali. Also as in Iberomesornis
romerali, the interclavicular angle is roughly 608, thereby form-
ing a V-shaped anterior profile. The lateral margins of the furcula
are slightly indented or excavated as noted by Sanz and Busca-
lioni (1992). This feature is also present in the South American
enantiornithine Neuquenornis volans and has been identified as
an enantiornithine synapomorphy (Chiappe and Calvo, 1994).
Proximally, the epicleideal processes are mediolaterally com-
pressed and abbreviated. As in I. romerali, the hypocleidium is
long and tapering. The anterior surface of the hypocleidium bears
a weak keel (Sanz and Buscalioni, 1992; Sanz et al., 1995).

Journal of Morphology
 THEROPOD FURCULA 871

Fig. 9. The furculae of nonavian ornithurines. (A) Yanornis martini (IVPP V13358) in anterior or posterior view. (B) Yixianornis
graubai (IVPP V13631) in anterior or posterior view. From Clarke et al. (2006). (C) Hongshanornis longicresta (IVPP V14533) in
anterior or posterior view. (D) Gansus yumenensis (CAGS-CM-003) in anterior view. (E) Ichthyornis dispar (SMM 2503, YPM 1755)
in anterior or posterior view. From Clarke (2004). Relationships after You et al. (2006) and Clarke et al. (2006). Scales 5 1 cm in
A–D.

Yixianornis graubai (Zhou and Zhang 2001). Specimen: IVPP Ornithurae

V13631 (Fig. 9B).
The furcula of Yixianornis graubaui is damaged at the sym-
physis. Photographs of the furcula prior to its damage (Clarke
et al., 2006; Fig. 6) shows that this portion of the furcula lacked
a hypocleidium and had a distinctively straight, squared-off
edge. The furcular rami are generally narrow, in contrast to the
broad rami common among more basal avialans. In Yixianornis,
the furcular rami narrow slightly proximally. The epicleideal
processes are short, mediolaterally narrow, and taper to a point.
It is unclear if, or to what extent, this process contacted the
acromion of the scapula. The imperfectly U-shaped furcula has
an estimated interclavicular angle of 758.
Yanornis martini (Zhou and Zhang, 2001). Specimen: IVPP
V12444, IVPP V13358 (Fig. 9A).
The furcula in Yanornis is exposed in ventral view but the
epicleideal processes are obscured by the coracoids. In overall
appearance, the furcula is similar to those of basal avialans. It
is robust and U-shaped with broad rami that do not taper proxi-
mally. A hypocleidium is lacking.
Hongshanornis longicresta (Zhou and Zhang, 2005). Speci-
men: IVPP V14533 (Fig. 9C).
The furcula of Hongshanornis longicresta differs from those of
basal avialans in having narrow, thin furcular rami. In overall
form, the furcula is U-shaped. In contrast to that of more basal
avialans, the furcula of Hongshanornis is elongate proximodis-
tally. In this sense Hongshanornis resembles more derived avia-
lans like Gansus yumenensis or Hesperornis regalis. A short hypo-
cleidium is present and appears to have contacted the sternum.
Details of the epicleideal process are obscured by rock matrix.
Ichthyornis dispar (Marsh, 1872). Specimens: SMM 2503,
YPM 1755 (Fig. 9E).
The furcula of Ichthyornis dispar is incompletely known
(Clarke, 2004). As a result, an interclavicular angle is roughly
estimated to be greater than 708 given the angle at the symphy-
sis. The rami are circular in cross section (Clarke, 2004) and
have a nearly uniform diameter as does the symphysis. The epi-
cleidium tapers to a blunt point and it appears that a hypoclei-
dium is not present (see Clarke, 2004). The furcula of Ichthyor-
nis dispar is very similar to that of Gansus yumenensis.
Gansus yumenensis (Hou and Liu, 1984). Specimens: CAGS-
CM-003 (Fig. 9D), CAGS-CM-004.
The furcula is both well preserved and well prepared in two
specimens of Gansus yumenensis. The furcula is U-shaped and
thin relative (You et al., 2006) to more basal theropod taxa. You
et al. (2006) reported an interclavicular angle of 388; however,
given the difficulty of the measuring the interclavicular angle
(see below), an interclavicular angle of 388 is much too low in
this comparative context. The epicleideal processes are only
slightly narrower than the shaft of the rami and lack distinctly
differentiated facets. The thin furcular rami have a circular
cross section and the symphysis lacks a hypocleideal process.
The rami and symphysis have a nearly uniform diameter.
Hesperornis regalis (Marsh, 1872). Specimens: YPM 1206.
The clavicles of Hesperornis regalis are unfused, failing to form
a true furcula. Marsh (1880) noted the similarities between the
clavicles of Hesperornis and the corresponding elements in em-
bryonic birds. Proximally each clavicle is very slender, whereas
distally the clavicular ramus expands slightly forming an articu-

Journal of Morphology
872 S.J. NESBITT ET AL.
lar surface for a midline contact with the complementary clavicle.
When the two clavicles are articulated they form a U-shaped
structure superficially similar to other basal ornithurines.
Iaceornis marshi (Clarke, 2004). Specimens: YPM 1734.
The known portion of the furcula for Iaceornis marshi is very
fragmentary, consisting of only a small portion of the proximal
end. This taxon is discussed here because it preserves a distinct
autapomorphy. The proximal end (the omal tip) as described by
Clarke (2004) is elongated and pointed.
DISCUSSION
The Early Evolution of the Furcula
Gauthier (1986) and Rauhut (2003a) utilized the
presence or absence of the furcula in their phylo-
genetic analyses of theropods, but did not identify
discrete characters of the furculae. Recently, large
phylogenetic analyses of avian relationships by
Livezey and Zusi (2006, 2007) and analyses of ba-
sal avialans by Clarke et al. (2006) have used fur-
cula synapomorphies to support avialan and, in
some cases, more basal theropod relationships.
The wealth of articulated specimens of Cretaceous
theropods now allows further parsing of characters
and character states (see Appendicies A and B1).
Many of the characters presented here are a first
attempt at identifying useful phylogenetic informa-
tion derived from the furcula, whereas other previ-
ously used characters are scored for more taxa.
The evolutionary transformation of the furcula
from separate clavicles is nicely illustrated in
archosaurs and their close relatives. Euparkeria
capensis and early pseudosuchians retain both the
interclavicle and clavicles. The interclavicle is lost
at the dinosaur node or at an unknown node
among early dinosauromorphs. From there, the or-
igin of the furcula is well understood with the
addition of the work on basal sauropodomorphs of
Yates and Vasconcelos (2005) and the discovery of
furculae in early coelophysoid theropods (Tykoski
et al., 2002; Rinehart et al., 2007). Some orni-
thischians have two small clavicles that do not
contact each other (Osborn, 1924a; Brown and
Schlaikjer, 1940; Sternberg, 1951; Chinnery and
Weishampel, 1998). In contrast, saurischians
retain clavicles with the clavicles contacting at the
midline. Coelophysoids, the earliest group of
undoubted theropods, have fully formed furculae.
Thus, the critical gap that added ambiguity to Bry-
ant and Russell’s (1993) analysis of the homology
of furcula has been filled by coelophysoids. The
present analysis demonstrates that the bird fur-
cula is homologous to the clavicles of other rep-
tiles. In addition, the furcula is probably a synapo-
morphy for Theropoda and evolved at least by the
Norian of the Late Triassic (character 2[1]).
The coelophysoids ‘‘Syntarsus’’ kayentakatae,
Coelophysis bauri, Coelophysis rhodesiensis, and
Segisaurus all possess furculae with a gently
curved symphyseal area and divergent epicleideal
processes. All these taxa share a U-shaped furcula
Journal of Morphology
(character [4]) that is more similar to coelurosaurs
than to basal tetanurans. It is interesting that
Jenkins et al. (1988), followed by Bailey and
DeMont (1991), hypothesized that the furcula is
related to the breathing system in avians, part of
a pneumatic system unique to theropods. It is
clear that the first theropods evolved both pneu-
matic features and a furcula at nearly the same
time given that both features are in C. bauri.
The early tetanurans Allosaurus and Suchomi-
mus tenerensis have similar furculae. Unfortu-
nately, these two furculae are the only representa-
tives among basal tetanurans and therefore serve
as the exemplars for a diversity of clades including
neoceratosaurs, spinosaurids, and carcharodonto-
saurids. Allosaurus and S. tenerensis share the fol-
lowing characters: V-shaped, round in cross sec-
tion, and dense with no modifications to the epi-
cleideal processes (character 4 [0] and character 6
[0]). This morphology is very similar to that of the
unfused clavicles of the early sauropodomorph
Massospondylus carinatus.
All coelurosaurs, with the exception of Velocirap-
tor mongoliensis, have furculae that are U-shaped
(character 4[1]). The basal-most coelurosaurs with
furculae have U-shaped furculae. These basal coe-
lurosaurs include the tyrannosaurids Albertosau-
rus sarcophagus, Daspletosaurus torosus, Tyranno-
saurus rex, and Gorgosaurus libratus and the
compsognathid Huaxiagnathus orientalis. The
tyrannosaurids A. sarcophagus, D. torosus, T. rex,
and G. libratus share one synapomorphy in the
furculae, the expansion of the epicleideal processes
(character 5[1]).
No unambiguously recognizable clavicle or fur-
cula has been found in any ornithomimid even
though well-preserved completely articulated speci-
mens are known from many taxa [e.g., Struthiomi-
mus altus (Nicholls and Russell, 1985), Gallimimus
bullatus (Osmolska et al., 1972), and the many skel-
etons of Sinornithomimus dongi (Kobayashi and
Lü, 2003)]. It is unclear if the furcula was never pre-
served, if it was not ossified, or if it did not form at
all. The absence of a furcula would be interpreted as
a secondary loss following the phylogeny presented
here (Fig. 1). The same absence is also apparent in
alvarezsaurids. However, it is also unclear whether
the apparent absence of furculae in alvarezsaurids
is a phylogenetic loss or a taphonomic/diagenetic
one. Unlike ornithomimids, alvarezsaurids are
known from a few largely incomplete specimens
making the absence of a furcula possibly a result of
taphonomic processes.
Members of the basal maniraptoran clades Ther-
izinosauridea and Oviraptorsauria have apomor-
phic furculae that are diagnosable to each clade.
The furculae of B. inexpectus and N. yangi are ro-
bust, the epicleideal processes are blunt, and both
have an unusually large interclavicular angle, a
combination of characters not present in other
 THEROPOD FURCULA
theropods. The absence of these characters in the
basal-most known therizinosaroid Falcarius uta-
hensis, suggests that the character states shared
by Beipaiosaurus inexpectus and Neimongosaurus
yangi evolved within the group and may be found
in nearly all therizinosauriods. Therizinosauroids
have been found as the sister taxon to Oviraptor-
sauria in many theropod phylogenetic analyses
(Theropod Working Group [ 5 TWiG], Turner
et al., 2007a). The specializations of the oviraptor-
saurid furculae (see below) are not present in ei-
ther therizinosauroid with known furculae, sug-
gesting that these characters are not shared
between the two.
Oviraptorosaur furculae are easily recognizable
from three identifying features, the presence of
long tapering epicleideal processes (character 7[1]),
the presence of a bulge on the ramus between the
epicleideal processes and the symphysis (character
8[1]), and some have a distinct hypocleidium (char-
acter 9[1]). These three features are found in all of
the oviraptorsaurs observed here (Khaan mcken-
nai, Oviraptor philoceratops, Citipati osmolskae,
Heyuannia huangi), but it is not clear whether the
more basal oviraptorosaurs have the same fea-
tures. Oviraptorid furculae also have a unique
articulation with the scapula; the epicleideal proc-
esses articulate on the anterior edge of the scap-
ula, rather than just lying on the anterior surface
of the acromion process of the scapula. In some
oviraptorid taxa such as O. philoceratops and C.
osmolskae, the anterior edge of the scapula sits in
a shallow groove of the epicleideal processes.
All oviraptorsaurs share a hypocleidium that is
nearly symmetrical and tapers to point ventrally
contrasting the small tuber present in many thero-
pods. In addition, this analysis indicates that the
hypocleidium in extant birds is not homologous
with those of oviraptorsaurs. The presence of an
anterior keel on the hypocleidium (character 9[1])
separates Oviraptor philoceratops from Citipati
osmolskae and may be useful in differentiating the
two taxa without cranial material.
Paravians, with the exception of Velociraptor
mongoliensis, share two furcula synapomoprhies
an anteroposterior compression of the symphysis
and rami (character 3[1]) and a nearly symmetri-
cal furcula (character 11[1]). Dromaeosaurids
have rather simple furculae. Most dromaeosaur-
ids have furculae that are U-shaped and thin rel-
ative to those of troodontids and avialans. V. mon-
goliensis has a unique furcula among dromaeo-
saurids and paravians. It is similar to that of
Allosaurus and other earlier theropods in being
V-shaped with a small tuber instead of a distinct
hypocleidium. Mei long and avialans share one
character, thickening of the symphysis and rami
(character 12[1]).
Basal avialans do not share any unique features
relative to other theropods. Within Avialae apo-
873
morphic characters are present in the furculae and
do appear to diagnose clades. Enantiornithines
have a furcula that is laterally excavated (char-
acter 13[1]) (Chiappe and Calvo, 1994). One of the
characters supporting a sister-group relationship
between the basal ornithurines Yixianornis gra-
baui and Songlingornis lingensis is the presence of
a truncated or squared base at the furcula sym-
physis (character 14[1]) (Clarke et al., 2006).
Iaceornis marshi, a poorly known North American
ornithurine, autapomorphically possesses epiclei-
deal processes that are long and pointed (character
6[1]). This feature is also seen in oviraptorsaurs
and anseriforms.
The early evolution of the furcula shows that
most features found in the furcula of extant birds
are found in all theropods. Only small changes
separate the furculae of early theropods such as
Coelophysis bauri from those of more derived
forms such as Archaeopteryx lithographica. It is
now clear that all major theropod clades have fur-
culae plesiomorphically. Clades and taxa in which
furculae are not found can now be interpreted as a
result of preservational bias or a secondary loss.
Variation Among Nonavian Theropods
Interspecific variation in furculae has been well
documented in the more basal theropods Allosau-
rus (Chure and Madsen, 1996), Coelophysis rhode-
siensis (Tykoski et al., 2002), and Coelophysis
bauri (Rinehart et al., 2007), but it has not been
documented in coelurosaurids with the exception
of tyrannosaurids (Makovicky and Currie, 1998).
This variation occurs in the interclavicular angle,
the shape of the epicleideal process, and the pres-
ence or absence of a tuber on the ventral side of
the symphysis. Further variation also occurs
within a single element. For example, differences
in the cross section of the ramus, the length of
each ramus, and shape of epicleideal processes
seem to be variable in one specimen. The furculae
of the tyrannosaur Gorgosaurus libratus clearly
show interspecific variation in the overall shape
(Makovicky and Currie, 1998). It is unclear
whether there is as much or more variation in
maniraptorans. Microraptor zhaoianus and Tyran-
nosaurus rex are the only taxa with more than one
specimen with a furcula. The only variation in the
furcula of these taxa seems to be small differences
in the interclavicular angle. Moreover, symmetry
is an absence of variation; a character state (11[1])
present in paravians. Variation in the furcula of
avians with powered flight is very minimal. How-
ever, birds that are secondarily flightless (e.g.,
Dromaius and Struthio) have very variable and
asymmetrical furcula/clavicles. The absence of var-
iation and presence of symmetry may be useful as
a proxy for determining a change in function, in
this case powered flight.
Journal of Morphology
874 S.J. NESBITT ET AL.
Taphonomy
Taphonomic processes may lead to misinterpre-
tations in the morphology of furculae in theropods.
Furculae are three-dimensional structures. Com-
pression may hide some features or create arti-
facts. For example, in lateral view, the epicleideal
processes arch both dorsally and posteriorly. If the
epicleideal processes are crushed, they will project
laterally. This seems to be the force behind the sig-
moidal shape of one specimen of Coelophysis rho-
desiensis with ‘‘laterally’’ projecting epicleideal,
and possibly asymmetries. Lastly, anteroposterior
crushing may mimic the flattened condition of the
furculae seen in avians and some nonavian thero-
pods. Some furculae like that of Buitreraptor gon-
zalezorum are highly pneumatic and could easily
collapse under pressure. Characters were scored
as question marks when morphology was unclear.
CONCLUSIONS
There no longer remains doubt that the furcula
of birds is homologous to the clavicles of tetrapods.
Both phylogenetic and developmental data
strongly support this conclusion. The clavicles fuse
into a single element at the base of Theropoda.
The origin and evolution of character states of
theropod furculae can be traced easily in the
theropod fossil record given the plethora of new
specimens with undoubted furculae. The furculae
of early avialans are nearly identical to closely
related clades such as Dromeosauridae and Troo-
dontidae. Only the early ornithurines possess a
furcula typical of extant avian clades. The furcula
has diagnostic features that can be utilized in
clade diagnoses.
ACKNOWLEDGMENTS
The authors thank P. Sweet (AMNH), C. Forster
and D. Krause (UA), X. Xu and Z. Zhou (IVPP),
P. Sereno (UCRC), T. Rowe (TMM), P. Makovicky
(FMNH), S. Apesteguı́a (MPCA), and R. Barsbold
(IGM) for access to specimens. Particularly they
like to thank Chistine Lipkin for providing them
with an advance copy of her paper on Suchomimus,
which was in press at the time of this manuscript’s
writing. They also like to thank L. Zanno for access
and permission to illustrate undescribed material of
Falcarius. R. Irmis, M. Lamanna, J. Gauthier,
L.Chiappe, J. O’Connor, J. Horner, P. Currie,
M. Loewen, P. Barrett, A. McDonald, and J. Sanz
were very helpful in providing images for this arti-
cle. M. Ellison skillfully prepared Figure 3. They
thank F.W. Harrison for a careful review and an
anonymous reviewer. C. Forester and N. Clay pro-
vided helpful comments on an earlier draft.
Journal of Morphology
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APPENDIX A: FURCULA CHARACTERS AND
CHARACTER DISCUSSION
Fourteen characters are presented below. These
characters describe the diversity of the furcula
among nonavian theropods. Many of the character
states in this set of characters have reversals
throughout the theropod tree. Additionally, charac-
ter states found in a variety of taxa are also auta-
pomorphic for others.
01. Interclavicle: present (0) or absent (1) (Gauthier,
1986; Sereno, 1991; Juul, 1994; Benton, 1999).
0 The interclavicle is present in archosaurs
plesiomorphically (Sereno, 1991). Ornithi-
schians and saurischians lack an interclavicle.
However, the pectoral girdles in the successive
sister taxa of the Dinosauria (Silesaurus, Mar-
asuchus, Lagerpeton) do not have the pectoral
region completely preserved. As a result, the
optimization of this character is not clear at
the base of Dinosauria.
02. Clavicles: unfused (0) of fused (1).
0 This character essentially describes the pres-
ence or absence of the furcula. The clavicles of
877
some nonsauropod sauropodomorphs (e.g.,
Massospondylus) contact each other at the
midline, but do not fuse (Yates and Vasconce-
los, 2005). The clavicles in ornithischians and
sauropods that retain them do not meet. It is
unclear if the presence of the furcula is the
result of one ossification or two (one for each
clavicle). There is one ossification for the fur-
cula in most birds including Gallus gallus
(Hall, 2001). However, we have observed the
presence of two clavicles in some paleo-
gnaths, demonstrating the presence of at
least two ossification centers. This character
has been employed by various datasets
exploring theropod relationships (e.g., Clarke,
2004).
03. Cross section of the furcula, shape: nearly cir-
cular (0) or anteroposteriorly compressed (1)
near the symphysis.
0 The furculae from more basal members of the
Theropoda (e.g., Coelophysis) are rounded in
cross section. Oviraptorid furculae are more
oval in cross section, but still differ from the
anteroposteriorly compressed furculae in most
paravians whereas the furcula of Suchomimus
is D-shaped in cross section (Lipkin et al.,
2007). The furcula of Aerosteon riocoloradensis
is anteroposteriorly compressed (Sereno et al.,
2008).
04. Furcula, general shape of the furcula: V-
shaped (0) or U-shaped (1).
0 The unfused articulated clavicles in Masso-
spondylus are V-shaped. This shape is found
in most of the early theropods with furculae,
including ‘‘Syntarsus’’ kayentakatakae, Sucho-
mimus, and Allosaurus. The ‘‘V’’ shape is a
result of nearly straight rami diverging from
the symphysis whereas U shaped furculae
have curved rami diverging from the symphy-
sis. The furculae of all coelurosaurs except
Velociraptor and some of those of Coelophysis
rhodesiensis and C. bauri are U -shaped.
05. Epicleideal processes: unexpanded (0) or ex-
panded (1).
0 The furculae of most theropods have unex-
panded epicleideal processes. Expanded epi-
cleideal processes are present in tyrannosaur-
ids as identified by Makovicky and Currie
(1998). Some variation in the size of the epi-
cleideal processes is present in Allosaurus as
noted by Chure and Madsen (1996). The slight
expansion of the epicleideal processes in Allo-
saurus is notably smaller than that of tyranno-
saurids.
06. Epicleideal processes: rounded, flat, or blunt
(0) or taper to a long point (1).
0 The epicleideal processes of most furculae
are rounded or slightly pointed (e.g., Coelophy-
sis, Allosaurus, Tyrannosaurus rex). This con-
trasts with the elongated tapering processes of
Journal of Morphology
878 S.J. NESBITT ET AL.
the furculae of oviraptorsaurs. The shape of
the ends of the epicleideal processes is related
to the way the furcula articulates with the
acromion process of the scapula. In taxa pos-
sessing rounded epicleideal processes, the fur-
cula attaches to the scapula by way of a liga-
ment. Here, the epicleideal process lies on the
anterior side of the scapula. In contrast, the
epicleideal processes of oviraptorsaurs articu-
late on the anterior edge of the scapula. This
articulation is much more expanded than that
of other taxa. This is convergently present in
anseriforms and Iaceornis marshi, and is an
autapomorphy of the latter taxon as identified
by Clarke (2004).
07. Rami, lateral expansion of the between the
hypocleidium and the epicleideal process:
absent (0) or present (1).
0 Most taxa display a condition in which the
ramus of the furcula is the same width as the
epicleideal processes and the hypocleidium
region (state 0). Oviraptorsaurs have an
expanded ramus (state 1).
08. Furcula, hypocleideum: absent or a small tu-
bercle (0), or present as a distinct and elongate
process (1).
0 As described in the terminology section, a
hypocleidium is a symmetrical (or nearly so),
ventrally projecting process at the symphysis
of the furcula. The small tubera present in
early theropods are not considered homologous
because the shape is highly variable within a
taxon (e.g., Allosaurus and Coelophysis) and
the presence or absence seems to be related to
interspecific variation (e.g., Allosaurus and
Coelophysis). All oviraptorsaurs and some avi-
ans are coded as having a hypocleidium.
09. Hypocleidium: rounded (0) or keeled (1).
0 The hypocleidium of oviraptorsaurs is either
smooth or keeled. A keeled hypocledium is pres-
ent in Oviraptor and ‘‘Big Momma’’ (IGM 100/
979).
10. Interclavicular angle: between 1008 and 1308,
(0), less than 1008 (1), or greater than 1308.
0 A similar character (character 268 of Senter,
2007) was recently utilized. The interclavicular
angle of early theropods is greater than 1008.
Maniraptorans usually have an interclavicular
angle of less than 1008, with the exception of
Beipiaosaurus and Neimongosaurus which
interclavicular angles greater than 1308. Para-
vians have furculae wherein the interclavicu-
lar angle varies from 708 to 908.
0 The limited documentation of the methods
that previous authors have employed to obtain
interclavicular angles (e.g., Larson and Rigby,
2005) limits further comparisons to the inter-
clavicular angles of furculae. The interclavicu-
Journal of Morphology
lar angle can be easily measured from the
symphysis to the epicleideal processes if the
furcula is V-shaped (straight rami diverging
from the symphysis). However, most theropods
have U-shaped furculae and therefore, the
exact interclavicular angle of curved rami can-
not be precisely measured; the interclavicular
angle depends on where the measurement is
taken along the curved rami and the absence
of distinct landmarks near the symphysis does
not allow measurements to be taken from a
common point. This is even more difficult
because the epicleideal processes of ornithur-
ine furculae arc medially at their tips. Fur-
thermore, apparent or unapparent crushing
and other taphonomic deformations do not
allow precise measurements.
0 Here, we measured the interclavicular angle
from the epicleideal processes to the symphysis.
The measurements were then rounded to the
nearest multiple of ten for scoring purposes.
This character is designed to separate furculae
with high interclavicular angles from those with
lower interclavicular angles, not to quantify the
difference between furculae with similar inter-
clavicular angles.
11. Furcula shape in anterior view: asymmetrical
(0) or symmetrical/nearly symmetrical (1).
0 The furculae of most nonparavian theropods
are markedly asymmetrical (e.g., Allosaurus,
Citipati). The furculae of paravians, with the
exception of Buitreraptor, are nearly symmetri-
cal. It is unclear if the asymmetry of the fur-
cula of Buitreraptor was the result of taphon-
omy or represents morphological asymmetry.
12. Rami: thin diameter (0) or thick diameter (1).
0 This refers to the thickness of diameter of
the rami and hypocleidium region. With the
exception of Mei, Archaeopteryx, Confuciusor-
nis, and Changchengornis all nonavian thero-
pods have relatively thin furculae. The furcu-
lae of Mei, Archaeopteryx, Confuciusornis, and
Changchengornis all have thick, robust furcu-
lae. Ornithurines have thin rami, much thin-
ner than those of basal theropods.
13. Furcula: laterally excavated: absent (0) or
present (1).
0 Furculae almost universally lack lateral exca-
vations. Presence of a laterally excavated fur-
cula an enantiornithine synapomorphy, that
was first described by Chiappe and Calvo (1994).
14. Furcula: ventral margin of symphysis
(5apophysis): curved, angling (0) or with a
truncated or squared base (1).
0 Presence of a truncated ventral apophyseal
margin is a synapomorphy of a clade including
Yixianornis 1 Songlingornis as identified by
Clarke et al. (2006).
 THEROPOD FURCULA 879
APPENDIX B1. Character scorings for taxa

1 2 3 4 5 6 7 8 9 10 11 12 13 14

Euparkeria capensis 0 0 — — — — — — — — — — — —
Pseudosuchia 0 0 — — — — — — — — — — — —
Ornithischia 1 0 — — — — — — — — — — — —
Sauropodomorpha 1 0 0 0 — — — — — — — — — —
Coelophysis bauri 1 1 0 0/1 0 0 0 0 — 0 0 1 0 0
Coelophysis rhodesiensis 1 1 0 0/1 0 0 0 0 — 0 0 1 0 0
‘‘Syntarsus’’ kayentakatae 1 1 0 0 0 0 0 0 — 0 0 1 0 0
Segisaurus halli 1 1 0 0 0 0 0 0 — 0 0 1 0 ?
Allosaurus fragilis 1 1 0 0 0/1 0 0 0 — 0 0 0 0 0
Aerosteon riocoloradensis 1 1 1 0 0 0 0 0 — 0 0 0 0 0
Suchomimus tenerensis 1 1 0 0 0 0 0 0 — 0 0 0 0 0
Huaxiagnathus orientalis 1 1 0 ? 0 0 0 ? — ? ? 1 ? ?
Scipionyx samniticus 1 1 ? ? ? 0 0 ? — ? ? ? ? ?
Protarchaeopteryx robusta 1 1 0 ? ? 0 0 0 — ? ? ? ? ?
Tyrannosaurus rex 1 1 0 1 1 0 0 0 — 1 0 0 0 0
Albertosaurus sarcophagus 1 1 0 1 1 0 0 0 — 1 0 0 0 0
Gorgosaurus libratus 1 1 0 1 1 0 0 0 — 1 0 0 0 0
Daspltosaurus torosus 1 1 0 1 1 0 0 0 — 1 0 0 0 0
UUVP 11689 1 1 0 1 1 0 0 0 — 1 0 0 0 0
Beipiaosaurus inexpectus 1 1 1 1 0 0 0 0 — 2 ? 0 0 0
Falcarius utahensis 1 1 1 1 0 0 0 0 — 1 0 0 0 0
Neimongosaurus yangi 1 1 1 1 0 0 0 0 — 2 0 0 0 0
Caudipteryx zoui 1 1 1 1 0 ? ? 0 — 1 ? 0 0 0
Oviraptor philoceratops 1 1 1 1 0 1 1 1 1 1 0 0 0 0
Citipati osmolskae 1 1 1 1 0 1 1 1 1 1 0 0 0 0
Khaan mckennai 1 1 1 1 0 1 1 1 0 1 0 0 0 0
Heyuannia huangi 1 1 1 1 0 1 1 1 0 1 0 0 0 0
Ingenia yanshini 1 1 1 1 0 1 1 1 0 1 0 0 0 0
Mei long 1 1 1 1 0 0 0 0 — 1 1 0 0 0
Sinornithoides youngi 1 1 0 1 0 0 0 ? ? 1 ? 1 ? 0
Velociraptor mongoliensis 1 1 0 0 0 0 0 0 — 1 1 0 0 0
Sinornithosaurus millenii 1 1 1 1 0 0 0 0 — 1 1 0 0 0
Bambiraptor feinbergorum 1 1 1 1 0 0 0 0 — 1 1 0 0 0
Buitreraptor gonzalezorum 1 1 1 1 0 0 0 0 — 1 0 0 0 0
Microraptor zhaoianus 1 1 1 1 0 0 0 0 — 1 1 0 0 0
Archaeopteryx lithographica 1 1 1 1 0 0 0 0 — 1 1 0 0 0
Sapeornis chaoyangensis 1 1 1 1 0 0 0 0 0 1 1 0 0 0
Jeholornis prima 1 1 1 1 0 0 0 0 — 1 1 0 0 0
Jixiangornis orientalis 1 1 1 1 0 0 0 0 — 1 1 0 0 0
Confuciusornis sanctus 1 1 1 1 0 0 0 0 — 1 1 0 0 0
Changchengornis hengdaoziensis 1 1 1 1 0 0 0 0 — 1 1 0 0 0
Iberomesornis romerali 1 1 1 1 0 0 0 1 1 1 1 0 1 0
Concornis lacustris 1 1 1 1 0 0 0 1 1 1 1 0 1 0
Yixianornis graubai 1 1 0 1 0 0 0 0 — 1 1 0 0 1
Yanornis martini 1 1 1 1 0 0 0 0 — 1 1 1 0 0
Hongshanornis longicresta 1 1 0 1 0 0 0 0 — 1 1 1 0 0
Ichthyornis dispar 1 1 0 1 0 0 0 0 — 1 1 1 0 0
Gansus yumenensis 1 1 0 1 0 0 0 0 — 1 1 1 0 0
Hesperornis regalis 1 1 0 1 0 0 0 0 — 1 1 1 0 0
Iaceornis marshi 1 1 0 1 0 1 0 0 — 1 1 1 0 0

Journal of Morphology