Journal of Vertebrate Paleontology 10(2):255-265, June 1990

0 1990 by the Society of Vertebrate Paleontology

OSTEOLOGY OF ARCHAEORNITHOMIMUS ASIATICUS

(UPPER CRETACEOUS, IREN DABASU FORMATION,
PEOPLE'S REPUBLIC OF CHINA)

DAVID SMITH1and PETER GALTON2

'Department of Geosciences, University of Arizona,
Tucson, Arizona 8572 1
2Department of Biology, University of Bridgeport,
Bridgeport, Connecticut 0660 1

ABSTRACT-The remains of the ornithomimid dinosaur Archaeornithomimus asiaticus from the

Upper Cretaceous of the People's Republic of China are described and comparisons made with the
bones of Elaphrosaurus, Gallimimus, Struthiomimus, Ornithomimus, Dromiceiomimus, Deinocheirus,
and Allosaurus. The diagnosis of the Ornithomimidae of Russell (1972) is expanded to include the
characters of Elaphrosavrus and Archaeornithomimus. Some of the remains from the Arundel For-
mation of Maryland that have been referred to Archaeornithomimus afinis are probably ornithomimid
but are generically and specifically indeterminate.

INTRODUCTION
The Omithomimidae are a family of highly cursorial
theropod dinosaurs whose remains are found in rocks
of Late Jurassic to Late Cretaceous age. Fossils of these
animals have been found in Africa, the People's Re-
public of China, and North America. They had small
heads, long necks, and slender, elongate limbs. The
morphology of the Upper Cretaceous ornithomimids
is remarkably uniform, and they have frequently been
compared to the modem ratite birds, especially the
ostrich.
Gilmore (1933) described a new species of the Late
Cretaceous genus Ornithomimus Marsh, 1890, Or-
nithomimus asiaticus, based on material that was found
by the American Museum of Natural History expe-
dition to Mongolia in 1923. The material, consisting
largely of disassociated material from different indi-
viduals, was found in the Iren Dabasu Formation of
what is now the People's Republic of China. No cranial
material was found. The cotypes are some associated
limb bones.
The Iren Dabasu Formation (Cenomanian (?): Weis-
hampel and Homer, 1986) is a 30 m cross-bedded
sandstone deposit with clay layers and intermittent
limestone lenses. Morris (1936) interpreted it as a ter-
restrial deposit with shallow ephemeral lakes. The ver-
tebrate fossils are found in the top 10 m. Three major
sites were excavated: AMNH Quarries 141, 145, and
149 (see Gilmore, 1933:fig. 1) (Weishampel and Hor-
ner, 1986), and these quarries were within 33 m of
each other. In addition to Archaeornithomimus asiatic-
us,-_the Iren Dabasu fauna includes pelecypods, -che-
lonians, fish, and crocodilians. Other dinosaurs include
a n o s a u r , the carnosaurian the?opod Alectrosau-
rus olseni Gilmore, 1933, the earliest known lambeo-
?<<
saurine hadrosaur Bactrosaurus iohnsoni Neishamvel
and Homer, 1986), and the earliest know; Asian had-
rosaurine Gilmoreosaurus mongoliensis (Brett-Sur-L
man, 1979). ---===
Russell (1972) defined the Ornithomimidae based
on the characters of the Upper Cretaceous North
American genera Ornithomimus, Struthiomimus, and
Dromiceiomimus. He also made Ornithomimus asiati-
cus the type species of a new genus, Archaeornithomi-
mus. Material from the Arundel Formation of Mary-
land, described as Ornithomimus afinis by Gilmore
(1920), was referred to Archaeornithomimus by Russell
(1972) as A. afinis. Nopsca (1928) classified the Upper
Jurassic genus Elaphrosaurus in the Ornithomimidae
but did not give any reasons for this assignment. Rus-
sell (1972) and Galton (1982) listed the ornithomimid
characters of Elaphrosaurus.
In the original article on the Iren Dabasu dinosaurs,
Gilmore (1933) provided a brief description of Or-
nithomimus asiaticus with only a few illustrations. He
considered it to be sufficiently similar to rely upon the
description of Stvuthiomimus altus by Osborn (19 17).
When Russell (1972) made Ornithomimus asiaticus
the type species of Archaeornithomimus, no additional
description was provided.
A well-illustrated description of Archaeornithomi-
mus asiaticus Gilmore, 1933, is presented. The defi-
nition of the Ornithomimidae of Russell (1972) is ex-
panded to include the characters of this genus, plus
Harpymimus (Barsbold and Perle, 1984) and Elaphro-
saurus; and the Arundel material is re-examined to
determine its taxonomic position.
Abbreviations -AMNH , American Museum of
Natural Histof, New York; GI, ~ e a o g i c aInstitute,
l
Academy of Sciences of the Mongolian People's Re-
 256 JOURNAL O F VERTEBRATE PAL,EONTOLOGY, VOL. 10, NO. 2, 1990
public, Ulan Bator; HMN, Humboldt Museum fur
Naturkunde, East Berlin; NMC, National Museum of
Canada, Ottawa; QG, Queen Victoria Museum, Salis-
bury, Zimbabwe; ROM, Royal Ontario Museum, To-
ronto; UCMP, University of California Museum of
Paleontology, Berkeley; UCMZ, University of Cal-
gary, Museum of Zoology, Calgary; USNM, National
Museum ofNatural History, Washington, D. C.; YPM,
Peabody Museum of Natural History, Yale University,
New Haven; ZPal, Palaeozoological Institute, Polish
Academy of Sciences, Warsaw.
SYSTEMATIC PALEONTOLOGY
Suborder THEROPODAMarsh, 1881
Family ORNITHOMIMIDAE
Marsh, 1890
i boot; [6] proximal end of metatarsal I11 constricted,
ARCHAEORNITHOMIMUS Russell, 1972
1
1I
Type Species-Ornithomimus asiaticus Gilmore,
1931.
~i
Diagnosis -As for Archaeornithomimus asiaticus
Gilmore, 1933, given below.
1 II~
1I ARCHAEORNITHOMIMUS ASIATICUS (Gilmore, 193 3)
Ornithomimus asiaticus Gilmore, 1933:27.
Archaeornithomimus asiaticus (Gilmore, 1933) Rus-
sell, 1972:378.
Lectotype -AMNH 6565, partial pes consisting of
distal tarsals 111 and IV, metatarsals 11, 111, and IV,
and phalanx 1 of digit IV (Fig. 4A-H).
Paralectotype -AMNH 6569, partial manus, juve-
nile, consisting of radius, ulna, metacarpals I, 11, and
111; and phalanges 1 of digit I, 2 of digit I, 2 of digit
11, and 2 of digit I11 (Fig. 21-R).
Type Occurrence-Upper 10 m of the Iren Dabasu
Formation (Upper Cretaceous, Cenomanian (?)) near
the Iren Dabasu telegraph station, People's Republic
of China.
1
1~~1
Referred Specimens- All from same locality as lec-
totype: AMNH 2 1786, cervical vertebra 5? (Fig. 1E-
1 Ill I); AMNH 21787, cervical vertebra 8? (Fig. lC, D);
1 1111
AMNH 2 1788, cervical vertebra 10, dorsal vertebrae
,11 1-4 (Fig. 1A); AMNH 2 1789, dorsal vertebrae 5-10?
I
(Fig. 1B); AMNH 21790, ilium, sacral vertebrae 1-5
I (Fig. 3C-F), proximal caudal vertebrae 1-5 (Fig. 1J);
~l
AMNH 2 179 1, proximal caudal vertebrae 3-1 I?;
/1!I 111
AMNH 21802, proximal caudal vertebrae 11-15(?);
AMNH 2 1889, spines from proximal caudal vertebra;
AMNH 21792, distal caudal vertebrae (Fig. lK, L);
AMNH 2 1793, 7 distal caudal vertebrae; AMNH
I
, I
I
2 1794,8 distal caudal vertebrae (all vertebrae formerly
AMNH 6576); AMNH 21795, scapulocoracoid and
humerus (Fig. 2A, B) (formerly AMNH 6566); AMNH
21796, humerus, radius, and ulna (Fig. 2C, D) (for-
I
merly AMNH 6567); AMNH 6569, radius, ulna, meta-
carpals 1-111, phalanges (Fig. 21-R);- A-M-N-H-2 1-889,
metaczgpal.T (Fig. 2s) (formerly-AMNH 6570): AMNH
2 1888, metacarpal I1 (Fig. 2T) (formerly AMNH 6570);
AMNH 21798, ischium (Fig. 3A) (formerly AMNH
6558); AMNH 2 1799, pubis (Fig. 3B) (formerly AMNH
6570); AMNH 21800, femur (Fig. 3G-L) (formerly
AMNH 6570); AMNH 2 1801, tibia, astragalus (Fig.
3M-R) (formerly AMNH 6576); AMNH 2 1797, as-
tragalus (formerlyAMNH 6570); AMNH 2 1803, pedal
ungual (Fig. 41); AMNH 21884,21885,21886,21887,
manual unguals (Fig. 2U-X) (all formerly AMNH
6570). Specimen numbers AMNH 6558, 6565, 6569,
and 6576 from Quarry 140; AMNH 6570 from Quarry
141. For measurements see Tables 1 and 2.
Diagnosis -[1] Five sacral vertebrae; [2] short, rel-
atively robust limb bones; [3] rounded anterior margin
of the coracoid with well-developed biceps tubercle;
[4] elongate metacarpal I short compared to metacar-
pals I1 and 111; [5] distal expansion of ischium ante-
riorly expanded; relatively flat ventral surface of pubic
not visible from anterior view when metatarsals artic-
ulated.
Character [ I ] separates Archaeornithomimus from
North American ornithomimids and Elaphrosaurus.
Character [2] separates Archaeornithornirnus from all
ornithomimids except Elaphrosaurus. Character [4]
separates Archaeornithomimus from North American
ornithomimids. Character [5] separates Archaeor-
nithomimus from other ornithomimids. Character [6]
separates Archaeornithomimus from ornithomimids
except for Elaphrosaurus and Harpymimus.
A
-
DESCRIPTION
Axial Skeleton
Cervical Vertebrae -One cervical vertebra (Fig. 1E-
I) is similar to the fifth Gallimimus (Osmolska et al.,
1972). It has a small neural spine that is bordered
anteriorly and posteriorly by deep excavations. The
diaphysis is borne by a broad transverse process that
is concave ventrally. The process is supported on the
posterior end by a thin sheet of bone that is bounded
dorsally by an excavation that extends from the end
of the postzygapophysis. The zygapophyses are level
with the top of the neural canal. The facet on the pre-
zygapophysis is convex laterally and slightly S-shaped
transversely. The postzygapophysis is incomplete and
more slender than the prezygapophysis. The amphi-
coelous centrum is long and low, with parapophyses
on the anterior end. Each parapophysis is borne on a
prominent ventrolaterally directed ridge on the comer
of the centrum. The ends of the centrum are oblique
to the long axis.
Another cervical vertebra (Fig. lC, D) is similar to
the eighth of Gallimimus (Osmolska et al., 1972). It
has a small neural spine bounded and undercut ante-
riorly and posteriorly by a deep excavation. The trans-
verse process is flat and broad and is supported by
anterior and posterior ventral struts. The posterior one
is bounded by deep excavations and is more slender
than the anterior one. Neither diapophysis is pre-
served. There is another excavation behind the trans-
verse process at the base of the postzygapophysis. The
 SMITH AND GALTON-OSTEOLOGY OF ARCHAEORNITHOMIMUS ASIATICUS 257

FIGURE 1. Archaeornithomimus asiaticus (Gilmore) from the Iren Dabasu Formation (Upper Cretaceous) of China. A,
AMNH 21 788, tenth cervical and first four dorsal vertebrae in lateral view; B, AMNH 21 789, dorsal vertebrae 5-1 0 in lateral
view; C-D, AMNH 21787, cervical vertebra 8(?) in lateral (C) and dorsal (D) views; E-I, AMNH 21786, cervical vertebra
5(?) in posterior (E), anterior (F), dorsal (G),lateral (H), and ventral (I) views; J, AMNH 21790, proximal caudal vertebrae
1-5 in lateral view; K-L, AMNH 21 792, distal caudal vertebrae in lateral (K) and dorsal (I ,) Abbreviations: cf, coracoid
views.
foramen; dp, diapophysis; n, neural spine; po, postzygapophysis; pr, prezygapophysis; r, rib. Scale bar = 5 cm.

prezygapophyseal facet is flat and dorsomedially di-
rected. The centrum is shorter than that of cervical 5
and the parapophysis is higher up on the centrum. The
ends are opisthocoelous and perpendicular to the long
axis.
Cervical 10 has a small neural spine bounded by
excavations as in 5 and 8 (Fig. 1A). The incomplete
transverse process is broad and supported by obliquely
inclined slender struts that are bounded by deep ex-
cavations. Neither diaphysis is preserved. The prezyg-
apophysis is borne by the transverse process and the
facet is flat and oriented dorsomediaIly. The incom-
hss lzteia: .*j;~-w&--T~L-
j;gapoP~4-~is -
centrum is shorter than that of 5 and 8. The parapophy-
sis is higher than in 8 and the ends are opisthocoelous
and perpendicular to the long axis.
Dorsal Vertebrae-Passing posteriorly along the col-
umn, the dorsal vertebrae increase in size (Fig. lA, B).
The neural spines are broad, rectangular, and over the
posterior half of the centra. The transverse processes
of the proximal dorsal vertebrae are supported by two
narrow struts, but the anterior strut merges with the
process by dorsal 6 (Fig. 1B). Oval diapophyses are
preserved on some of the processes. The processes of
the first dorsal vertebrae are flat, but become increas-
ingly oblique and oriented posteriorly further along the
c-r\l..mn
VVIWX-A-L-LZ7
A& +h;-
1 Llllll.
chon+ nf hr\n~
LIIIWFVl-xIV
~r\nnar+
-VVI.AzLVV C V

apophysis to the transverse process. A strut runs from

t h~e m t ~ v x r ~ - -
llilV
YLY"Ja
 Ij 258 JOURNAL OF VERTEBRATE PALEONTOLOGY, VOL. 10, NO. 2, 1990
1111 1

1 1 I
TABLE 1. Measurements (mm) of the appendicular skeleton of Archaeornithomimus asiaticus. Phalanges are labeled so that
1
I
1 the first number indicates the digit and the second is the phalanx.

Proximal width

I
II~II
1 /1

l~ 11 1
~
I
Specimen number
AMNH 21795
AMNH 21795
AMNH 21795
AMNH 21796
AMNH 21796
AMNH 21796
Element
Scapula
Coracoid
Humerus
Radius
UIna
Humerus
Length
132.0
110.6
256.0
205.0
2 17.0
272.0
Width (min)
14.8
64.5
17.0
11.4
12.0
21.3
(max)
5.83
2.53
57.9
23.6
29.3
59.5
Distal width (max)
2.44

38.4
21.7
23.0
49.6

I1 I
ill
AMNH 6569
AMNH 6569
AMNH 6569
Radius
Ulna
Metacarpal I
140.0
148.0
46.9
7.5
7.9
5.6
17.0
19.2
13.0
14.7
18.0
12.1
i /I AMNH 6569 Metacarpal I1 54.5 5.7 15.6 11.8
AMNH 6569 Metacarpal I11 50.3 5.1 13.1 10.3
il'l AMNH 6569 Phalanx I, 1 57.2 6.3 11.4 10.2
AMNH 6569 Phalanx I,2 16.6 7.8 8.5 11.4

11 lIlI1
1
AMNH 6569
AMNH 6569
AMNH 6569
AMNH 21790
AMNH 21799
Phalanx II,2
Phalanx 1142
Phalanx III,3
Ilium
Pubis
43.5
26.7
57.0
114.0
303
280
5.8
8.1
-
73.5
14.2
11.4
10.0
14.6
-
-
54.4
99.5
131
9.7
9.9
-
-
56.9
I l ~ l /I AMNH 21798 Ischium
I 11 AMNH 21800 Femur 314 31.5 64.7 65.3
I
111 1 AMNH 2180 1
AMNH 6565
Tibia
Tarsal I11
40 1
33.9
24.0
29.4
81.3
-
67.7
-
;1 l1 AMNH 6565 Tarsal IV 32.6 28.5 - -
AMNH 6565 Metatarsal I1 257 18.1 28.9 51.8
AMNH 6565 Metatarsal I11 286 10.5 36.7 33.7
AMNH 6565 Metatarsal IV 259 18.2 45.5 33.3
AMNH 6565 Phalanx I,4 70.8 16.4 31.1 -5.4
AMNH 21797 Astragalus 60

the prezygapophysis obliquely to the postzygapophy-
sis. In dorsals 4 and 5, a thin sheet of bone runs from
the circular parapophysis to the transverse process.
From dorsal 6 caudally, the sheet merges with the dor-
sal sheet to form a triangular depression. The postzyg-
apophyses are borne on the base of the neural spine.
The centra are high with opisthocoelous ends.
Sacral Vertebrae-There are four sacral vertebrae
fused by sacral ribs to the ilium (Fig. 3C-E), and there
is room for a fifth (Fig. 3F) that is preserved separately.
Sacral centra 1 and 2 are the longest and 3 is the short-
est. The ribs are borne high on the centra. The first rib
~11 I is located on the anterior end of the centrum. The
second is jointly held by centra 1 and 2. The third rib
is solely borne by the anterior end of centrum 3, and
the fourth is shared by centra 3 and 4. The last rib is
in the middle of centrum 5. The ends of the sacral ribs
are expanded and rugose. There are pleurocoels on the
first four centra, but they are most highly developed
on sacrals 2 and 3. There are shallow grooves on the
borne by the neural spine. There is a small thin ridge
between the postzygapophysis and the base of the sa-
cral rib. The opisthocoelous centrum is not as con-
stricted at midlength as the centra of sacrals 1-4 are.
The rugose posterior end is round, but there is a shal-
low concavity on the -smooth anterior end.
Caudal Vertebrae-The most proximal caudal ver-
tebrae (Fig. 1J) have longer centra than the last sacral
vertebra, but these, along with the neural spines and
transverse processes, get progressively smaller passing
' posteriorly. The proximal neural spines are tall, broad,
and thin. The level of the transverse processes is above
the neural canal. The prezygapophyses extend in front
of the centrum, but are relatively short. The postzyg-
apophyses are borne by the neural spine and do not
project behind the centrum. Each has a laterally di-
rected concave facet with an oval outline. The centra
are amphicoelous with a midlength costriction, and
each has a shallow ventral groove.
The distal caudal .vertebrae (Fig. lK, L) have rela- I

I
I 1 ventral side of centra 1, 2, and 4. tively long centra and small neural spines that are even-
I
I
Sacral 5 (Fig. 3F) is distorted, but it is the best pre- tually reduced to slight ridges. There are no transverse
Y
served of the sacral vertebrae. It has a short centrum processes. The long, horizontal prezygapophyses ex-
I
~l
l
and a very high, thin, broad neural- spine. TKe stout tend almost halfiway over the preceding vertebra, but 1
--- - mlLf.Ib is s72ppc-teG2bG7y7e-k,;~+ , h ; ~Tt ~ - "P -~-S- ~
9 P + ->.IL-
+he
A 2A
U
;Jg o~n+
U V V I I A .
onn+q
W X . ~ X
An nnt
-VVII". .
h z x rY
nUrl;S+;no+
I C I Y V .
f ~ o aCU.t a T
IUVV I --I
~ P
- ,!
I
from the neural spine. The zygapophyses are small and postzygapophyses are short rods that extend back from
i
+
SMITH AND GALTON-OSTEOLOGY OF ARCHAEORNITHOMIMUS ASIATICUS 259

, TABLE 2. Measurements (mm) of the axial skeleton of Ar-

? chaeornithomimus asiaticus. Distal caudal vertebrae are ar-
bitrarily numbered. Abbreviations: d, distorted; e, estimate.
\ Specimen number Vertebra Centrum length
Q
1 AMNH 21786 C5 69.5
AMNH 21787 C8 64.0
AMNH 21788 C10 56.9

i AMNH 21788
AMNH 21788
AMNH 21788
AMNH 21788
Dl
D2
D3
D4
53.5
47.7
45.7
45.0

6,
I AMNH 21788
AMNH 21788
D5
D6
48.5
49.8
AMNH 21788 D7
AMNH 21788 D8
AMNH 21788 D9
AMNH 21789 D5
AMNH 21789 D6
AMNH 21789 D7
AMNH 21789 D8
AMNH 21789 D9
AMNH 21789 Dl0
AMNH 21790 S1
AMNH 21790 S2
AMNH 21790 S3
AMNH 21790 S4
AMNH 2 1790 S5
AMNH 2 1790 Ca 1
AMNH 2 1790 Ca2
AMNH 21790 Ca3
--s3
AMNH 21790 Ca4
AMNH 21790 Ca5 FIGURE 2. Archaeornithomimus asiaticus (Gilmore) from
AMNH 21791 Ca3 the Iren Dabasu Formation (Upper Cretaceous) of China. A-
AMNH 21791 Ca4 B, AMNH 21795, right coracoid in lateral view (A), and
AMNH 2179 1 Ca5 right scapula in lateral view (B); C-D, right humerus, AMNH
AMNH 2179 1 Ca6 21 796, in lateral (C) and anterior (D) views; E-F, ulna in
AMNH 2179 1 Ca7 anterior (E) and medial (I?) views; G-H, radius in anterior
AMNH 2 1791 Ca8 (G)and medial (H) views; I-R, AMNH 6569, paralectotype
AMNH 21791 Ca9 articulated metacarpals in proximal, palmar, and distal views
AMNH 21791 CalO (I);metacarpal I in dorsal (J)and palmar (K) views; meta-
AMNH 21791 Call carpal I1 in dorsal view (L);metacarpal I11 in palmar view
AMNH 21802 Ca12 (M);phalanx I, 1 in lateral (N) and palmar (0)views; phalanx
AMNH 21802 Ca13 I,2 in dorsal view (P); phalanx II,2 in dorsal view (Q); pha-
AMNH 2 1802 Ca14 lanx III,2 in dorsal view (R); S, AMNH 21889, adult meta-
AMNH 21802 Ca15 carpal I in proximal, palmar, and distal views; T, AMNH
AMNH 21792 Ca 1 2 1888, adult metacarpal I1 in proximal, dorsal, and distal
AMNH 21792 Ca2 views; U-X, AMNH 21884, 21885, 21886, 21887, ungual
AMNH 21792 Ca3 phalanges in lateral and proximal views. Abbreviations: bt,
AMNH 2 1794 Ca 1 biceps tubercle; cf, coracoid foramen; dp, deltopectoral crest;
AMNH 21794 Ca2 it, internal tuberosity; op, olecranon process; rc, radial con-
AMNH 21794 Ca3 dyle; uc, ulnar condyle. Scale bar = 5 cm.
AMNH 21794 Ca4
AMNH 21794 Ca5
AMNH 21794 Ca6
AMNH 2 1794 Ca7 42.4
AMNH 21794 Ca8 41.7 the neural spine. Haemal arches are preserved inter-
AMNH 21793 Cal 44.9 vertebrally on some of the vertebrae.
AMNH 21793 Ca2 45.7
AMNH 21793 Ca3 44.8 Appendicular Skeleton
AMNH 21793 Ca4 44.5
AMNH 21793 Ca5 41.5 Scapula-The preserved ventral half of the scapula
AMNH 21793 Ca6 42.5 (Fig. 2B) is concave longitudinally and convex later-
L&-?TE 2 !7 83 C27 20 < z3,r. The s2.,2;k, sectis=. vet-
trally, the bone is expanded and thick. An oblique ridge
260 JOURNAL OF VERTEBRATE PALEONTOLOGY, VOL. 10, NO. 2, 1990
is present on the lateral side adjacent to the glenoid
fossa.
Coracoid-The coracoid (Fig. 2A) is plow-shaped
with a concave medial surface. The thickest region is
below the glenoid fossa and the adjacent part of the
scapular suture, and the bone becomes thinner passing
anteriorly. There is a prominent biceps tubercle set in
from the ventral edge. The supracoracoid foramen is
below the middle of the scapular suture.
Humerus -The shaft of the right humerus (Fig. 2C,
D) is long and slender with an oval cross section. It is
curved medially and twisted counterclockwise. The
head is more expanded than the distal end. There is a
thin, poorly developed deltopectoral crest on the upper
third of the bone. Distally, the radial and ulnar con-
dyles are subequal and separated by a shallow groove.
Radius-The radius (Fig. 2G, H) is shorter and
slightly more slender than the ulna. It is straight with
a circular cross section. The ends are not expanded.
The proximal articulation has an oval outline. Distally,
the articulation is obliquely inclined.
Ulna -The ulna (Fig. 2E, F)is somewhat longer than
the radius. The shaft is slender but is distorted: it should
be straight as it is in another specimen. It has a tri-
angular cross section proximally and becomes more
rounded distally. Both the proximal and radial artic-
ulations are concave. Distally, the ulna is only slightly
expanded anteroposteriorly and has a subrectangular
outline. The distal radial articulation is continuous with
the distal articulation.
Metacarpals -The associated metacarpals are from
a juvenile (Fig. 21-M). Isolated adult metacarpals I and
I1 are also illustrated (Fig. 2S, T). Metacarpal I is the
shortest present. It has a subtriangular cross section
and neither end is expanded. The outline of the prox-
imal articulation is subrectangular. There is a sharp
medial edge and the lateral surface of the shaft is con-
cave where it contacts metacarpal 11. Distally, the lat-
eral condyle is larger than the medial one.
Metacarpal I1 is about the same thickness as meta-
carpal I. The shaft is straight with a subrectangular
cross section. The proximal end is more expanded than
the distal one. The distal condyles are equally devel-
oped, but the end has an irregular outline. The lateral
and medial fossae are both shallow.
Metacarpal I11 has a thin shaft with a rounded cross
section. There is a prominent concave scar on the me-
dial side where it contacts metacarpal 11. Both ends
are expanded. The proximal articular surface is irreg-
ular and the distal end has a subtriangular outline.
Phalanges -Not all of the phalanges are preserved.
The three metacarpals are associated with phalanx 1
of digit I, 2 of digit I, 2 of digit 11, and 2 of digit I11
(Fig. 2N-R). Gilmore (1933) measured phalanx 3 of
digit 111, but this element is now missing from the
collection. Phalanx I,1 is long and slender. The shaft
is predominantly oval in cross section, but i t has a
flattened palmar surface. The ends are not expanded.
Zlistaiiy, the conciyies are ~e~i-ci-e~ei~p~-iran-Qm-cm-
pletely separated by a distinct intercondylar sulcus.
The lateral and medial fossae are distinct.
Phalanx I,2 is very short with a trenchant palmar
surface that is continuouswith the intercondylar groove.
Neither end is expanded. The distal condyles are well
developed and the lateral and medial fossae are indis-
tinct.
Phalanx II,2 is long and slender and has slightly
expanded ends. Proximally, the palmar surface is
slightly concave, but distally the shaft becomes oval
in cross section. The condyles, the intercondylar groove,
and the fossae are all well developed.
Phalanx III,2 is short and relatively thick. The pal-
mar surface is trenchant throughout the length of the
shaft. The proximal end is expanded. The condyles
and intercondylar groove are well developed, but the
lateral and medial fossae are shallow.
The ungual phalanges (Fig. 2U-X) vary from sharply
recurved to almost straight. The recurved unguals are
more laterally compressed than the straighter ones.
Flexor tubercles and lateral grooves are present on all
of the manus unguals. By comparison with the figures
in Nicholls and Russell (1985), the ungual in Figure
2W can probably be referred to digit I, the ungual in
Figure 2 to digit 11, and the unguals in Figures 2V and
X to digit 111.
Pelvis-The ilium (Fig. 3C) is long and low. Later-
ally the body is concave and the anterior process is
shorter and more rounded than the posterii36ne. The
outline of the dorsal margin is convex anteriorly and
more concave posteriorly. The ventral margin is in-
complete. Posteriorly, the blade flares away from the
vertebral column. The acetabular margins are thin but
become more robust near the peduncles. The semicir-
cular ischiadic peduncle is more robust than the sub-
triangular pubic peduncle. There is a broad, medially
directed brevis shelf that extends from the ischiadic
peduncle to the posterior end.
The pubis (Fig. 3B) has a long shaft with an antero-
posteriorly expanded proximal end. The contacts for
the ilium and ischium are poorly defined. The acetabu-
lar surface is short and almost flat. A thin sheet that
starts about one-fourth ofthe way down the shaft forms
the apron. The boot is robust and extends more pos-
teriorly than anteriorly. The ventral surface of the boot
is slightly convex.
The ischium (Fig. 3A) has a narrow, flattened shaft
that becomes more rounded distally. The contacts with
the ilium and pubis are well defined with a semicircular
acetabular edge in between-The obturator process is a
wedge-shaped flange about one-fifth of the way down
the shaft. Below the obturator process, the edge of the
shaft is sharp anteriorly and more rounded posteriorly.
The distal end is expanded anteroposteriorly.
Femur-The femur (Fig. 3G-L) is almost certainly
shorter than the tibia, but these bones have not been
found associated. The shaft is relatively robust and has
an almost square cross section. The proximal end is
- expanded to form a he&-an-d-aweii-deveioped iesser
 SMITH AND GALTON- OSTEOLOGY OF ARCHAEORNITHOMIMUS ASIATICUS 26 1

t FIGURE 3. Arclzaeornithomimus asiaticus (Gilmore) from the Iren Dabasu Formation (Upper Cretaceous) of China. A,
AMNH 21798, ischium, in lateral view; B, AMNH 21799, pubis, in lateral view; C-F, AMNH 21790, ilium and sacral
vertebrae 1-4 in lateral (C), ventral (D), and anterior (E) views; F, sacral vertebra 5 in lateral view; G-L, AMNH 21 800, right
femur in posterior (G),lateral (H), anterior (I),medial (J),proximal (K), and distal (L) views; M-R, AMNH 2 1801, left tibia
1 with astragalus in proximal (M), distal (N), posterior (0), lateral (P),anterior (Q), and medial (R) views. Abbreviations: as,
astragalus; cn, cnemial crest; fc, fibular crest; gt, greater trochanter; h, head; It, lesser trochanter; op, obturator process; sr,
sacral ribs. Scale bar = 5 cm.

trochanter. The top of this trochanter is separated from
L
I
the greater trochanter by a notch 10 mm deep. The
neck is almost perpendicular to the shaft and is min-
imally constricted. Distally, the condyles are subequal,
the condylar fossae are shallow, and the intercondylar
i sulcus is only present posteriorly.
Tibia-The tibia (Fig. 3M-R) has a long shaft with
an oval cross section. The proximal end is rugose and
expanded anteroposteriorly. There is a large but thin
cnemial crest extending one-fifth of the shaft length. A
smaller fibular crest extends about one-sixth of the
shaft length. The distal end is transversely expanded
-I to become flat and wedge shaped-urn-der the astraealus.
Tarsus -The condyles of the astragalus are variable
in their relative development in difkrent individuals.
The astragalus wraps around the distal end of the tibia
but is not fused to it (Fig. 30). There is a well-devel-
oped intercondylar sulcus. The ascending process is
incomplete in all of the specimens available. The an-
terior surface of the process is transversely concave.
The calcaneum is not present in the collection but
would have been a separate bone.
Two distal tarsals were found associated with three
metatarsals (Fig. 4H). They are identified as tarsals I11
and IV following Gilmore (1933). They are compressed
and similar to those of the dromaeosaurid theropod
De!'vouycb~&(Ostro- 19-69_>s_nk_z-e gently convex
proximally and concave distally. As in Deinonychus,
~I
I I
262 JOURNAL OF VERTEBRATE PALEONTOLOGY, VOL. 10, NO. 2, 1990
I 1 1
& phalanx (Fig. 41). Phalanx IVY1 is robust and neither
0 tral comers, there are long tubercles that end before
I l l
1 curved pedal unguals as a diagnostic character for Ar-
FIGURE 4. Archaeornithomirnus asiaticus (Gilmore) from
the Iren Dabasu Formation (Upper Cretaceous) of China. A-
H, AMNH 6565 (lectotype), partial right pes, metatarsal 11
in posterior (A) and medial (B) views; metatarsal I11 in pos-
tenor (C) and lateral (D) views; metatarsal IV in posterior
(E) and lateral (F) views; articulated metatarsals in anterior
1 1 and distal views (G),in proximal views with tarsals 111 and
IV removed and added (H); I, AMNH 2 1803, isolated pedal
ungual in proximal and lateral views. Scale bar = 2.5 cm in
A-H, 5 cm in I.
there is a transverse ridge on the distal side of tarsal
111. Tarsal IV is similar to 111, except that it is slightly
thinner and lacks the transverse ridge.
Metatarsals-Metatarsal I1 (Fig. 4A, B) is long and
straight. The shaft has an oval cross section. The prox-
imal end is expanded with a subtriangular outline. It
has a flat tarsal articulation. A concave facet on the
medial side suggests the presence of a metatarsal I.
Medially, a concave contact surface is present for meta-
tarsal 111. Distally, the larger bulbous lateral condyle
is separated from the wedge-shaped medial condyle by
a deep posterior groove.
Metatarsal I11 (Fig. 4C, D) is the longest and most
slender of the three metatarsals. The proximal half of
the shaft is constricted and has a semicircular cross
section. The proximal end is expanded anteroposte-
riorly to become wedge shaped. The distal end is more
robust and has a triangular cross section. The distal
condyles are well developed and equal in size with
pronounced fossae.
Metatarsal IV (Fig. 4E, F) has a robust shaft com-
pared to metatarsal I11 and is approximately the same
length and thickness as metatarsal 11. It has an oval
cross section and the bone becomes somewhat more
slender distally. The proximal end is expanded with
concave facets for contact -with metatarsal I11 and the
1~
missing metatarsal V. Distally, the condyles are not
expanded. The posterior intercondylar groove is shal-
I i
low and-poody developed. There is a well-developed
medial fossa.
Phalanges-The only phalanges preserved are the
first phalanx ofdigit IV (Fig. 4G) and an isolated ungual
end is expanded. The top is shorter than the bottom,
so the concave proximal articulation is obliquely in-
clined. There is a pair of tubercles on the proximal end
that converge a short distance down the shaft. Distally,
the wedge-shaped condyles are well developed and sep-
arated by a deep intercondylar sulcus.
The single, isolated pedal ungual is not recurved but
straight. There is a round concave base. On both ven-
the base is reached. All of these characters are shared
with the late Cretaceous ornithomimids. Gilmore
(1933) and Russell (1972) cited the possession of re-
chaeornithomimus, but this conclusion was based on
manus unguals- The recurved unguals the Iren
Dabasu r or mat ion that not referable to the manus of
Archaeornithomimus should be referred to the thero-
pod cf. Alectrosaurus also described by Gilmore (1933)
from the Iren Dabasu Formation of the People's Re-
public of China. The recurved pedal ungual from
~ ~( ~ i ~ l 1920)
~ ~should
~ ~ ~belreferred
, ~to The- ~
ropoda, incertae sedis.
COMPARISONS
The bones ofArchaeornithomimusare compared with
descriptions of those of Elaphrosaurus (Jamensch, 1925;
Galton, 1982; HMN) and Allosaurus (Gilmore, 1920;
Madsen, 1976) from the Upper Jurassic of North
America and East Africa, Gallimimus (Osmolska et
al., 1972) and Deinocheirus (Osmolska and Roniewicz,
1970) from the Upper Cretaceous of Mongolia, and
Ornithomimus (Osborn, 1917; Sternberg, 1933; Rus-
sell, 1972; Nicholls and Russell, 1980), Struthiomimus
(Osborn, 1917; Russell, 1972; Nicholls and Russell,
1980, 1985), and Dromiceiomimus (Parks, 1933; Rus-
sell, 1972) from the Upper Cretaceous of North Amer-
ica. Skeletons of Allosaurus (ROM 509 I), Struthiomi-
mus (ROM 840), and Ornithomimus (ROM 851), as
well as a cast of Gallimimus (GI No. DPS 100/11) at
the Royal Ontario Museum, were also compared to
- Archaeornithomimus. All except Allosaurus have been
referred to the Ornithomimidae. In addition, compar-
isons are made with isolated bones of Struthiomimus
(AMNH 5335, 5339, 5385, 5375) and Dromiceiomi-
rnus (AMNH 5201).
There are five sacral vertebrae in Archaeornithomi-
rnus and Gallimimus (Osmolska et al., 1972), and six
in Elaphrosaurus and the North American genera Stru-
thiomimus, Ornithomimus, and Dromiceiomimus (Ja-
nensch, 1925; Russell, 1972). The prezygapophyses of
the distal caudal vertebrae ofArchaeornithomimusand
the other ornithomimids are proportionally more slen-
I
der than those of Elaphrosaurus (Janensch, 1925). I
The crania of the ornithomimids are all very similar J
and, at this point. most ofthe distinguishing characters --
are found in the postcranial skeletons, although the i
braincases have not yet been closely compared. The 1
i
i
 SMITH AND GALTON- OSTEOLOGY OF ARCHAEORNITHOMIMUS ASIATICUS
cranium of Struthiomimus samueli (ROM 840) was
compared with a cast of Gallimimus bullatus. In Gal-
limimus, the posterior end of the mandible is deflected
down rather than being straight as in Struthiomimus.
Correspondingly, the jugal angles down in Gallimimus.
In Gallimimus, a horizontal crest is formed by the
prefrontal and frontal bones over the orbit. This struc-
ture was not observed in the other ornithomimids.
In the coracoid of the Asian genera, including Ar-
chaeornithomimus (Fig. 2A), the anterior margin is
dorsoventrally more convex than it is in the North
American genera and the posterior notch is wider. This
element is proportionally longer and more wedge
shaped in the North American genera than it is in the
Asian ornithomimids (Osborn, 1917; Sternberg, 1933;
Osmolska and Roniewicz, 1970; Osmolska et al., 1972;
Russell, 1972; Nicholls and Russell, 1985). This ele-
ment is incomplete in Elaphrosaurus (Janensch, 1929).
The biceps tubercle is less well developed in Deinochei-
rus (Osmolska and Roniewicz, 1970) and Gallimimus
(Osmolska, et al., 1972) than it is in Archaeornithomi-
mus.
The deltopectoral crest of the humerus is lower in
Elaphrosaurus than in the later ornithomimids. There
is a more pronounced deltopectoral crest in Gallimi-
mus and Deinocheirus (Osmolska et al., 1972) than in
Archaeornithomimus and the North American or-
nithomimids. In Archaeornithomimus, metacarpal I1
is the longest and metacarpal I the shortest. Metacar-
pals I1 and I11 are subequal in Struthiomimus. Meta-
carpal I is the longest in Ornithomimus (Nicholls and
Russell, 1980). Metacarpal I is the shortest one in Gal-
limimus (Osmolska et al., 1972) and Deinocheirus (Os-
molska and Roniewicz, 1970). Metacarpal I11 is longer
than I1 in Deinocheirus (Osmolska and Roniewicz,
1970) and they are subequal in length in Gallimimus
(Osmolska et al., 1972). All of the metacarpals are
longer relative to the ulna in the North American gen-
era than they are in Archaeornithornimus. Metacarpal
ratios for selected genera are given in Table 3. The
unguals are more massive in Gallimimus than in
Archaeornithomimus or any of the North American
genera and are more similar to those of Deinocheirus
and Albertosaurus.
The ventral surface of the pubic boot is anteropos-
teriorly convex in Ornithomimus (Russell, 1972), Stru-
thiomimus (Osmolska et al., 1972), and Dromiceiomi-
mus (AMNH 5201). This surface is flatter in
Gallimimus (Osmolska et al., 1972), Archaeornitho-
mimus (Fig. 3B), and Allosaurus (Gilmore, 1920). In
Archaeornithomimus, the obturator process of the is-
chium is larger than in Dromiceiomimus (AMNH
5201). The distal end of the ischium is more robust
and extends anteriorly in front of the shaft more than
the blade-like character of the other ornithomimids.
The shaft of the pubis is straighter in Gallimimus than
in other ornithomimids.
The neck of the femur is less well deyeloued in Gal-
limimus (Osmolska et al., 1972) and an isolated or-
nithomimid femur from North America than in Ar-
263
TABLE 3. Metacarpal ratios of selected theropods. Values
for metacarpal I1 are reduced to unity. Ratios for Gallimimus,
Deinonychus, Syntarsus, and Dilophosaurus calculated from
figures in Osmolska et al. (1972), Ostrom (1969), Raath (1969)
and Welles (1984), respectively.
Taxon
Archaeornithomimus (AMNH 6569)
Ornithomimus (ROM 85 1)
Gallimimus (GI DPS 100/11)
Allosaurus (ROM 509 1)
Albertosaurus (ROM 672)
Deinonychus (YPM 52 11)
Syntarsus (QG/ 1)
Dilophosaurus (UCMP 37302)
chaeornithomimus. In the femur of Elaphrosaurus, the
lesser trochanter is further from the proximal end than
it is in the other ornithomimids. The tibia of Ar-
chaeornithomimus (Fig. 30-R) has a rounded cnemial
crest that extends above the femoral articulation. In
Struthiomimus, the crest is wedge shaped and level
with the articular surface. The metatarsals ofdrchaeor-
nithomimus (Fig. 4A-G) are more robust than those
of Gallimimus and the North American genera. Unlike
the other ornithomimids, the proximal e n a f meta-
tarsal I11 of Elaphrosaurus and Harpymimus are vis-
ible from an anterior view when the metatarsals are
articulated together. The pedal unguals are more re-
curved in Gallimimus than in other ornithomimids.
DISCUSSION
Familial Diagnosis
Russell (1972) cited the following characters as being
diagnostic for the family:
[I] Skull relatively light and edentulous. [2] Maxilla
excluded fiom external narial border, secondary pal-
ate present. [3] Orbit large. [4] Temporal region re-
duced, alae from squamosal may nearly enclose su-
pratemporal fenestra. [5] With possible exception of
anterior cervicals, presacral vertebrae lack pleuro-
coels. [6] Six centra co-ossified in adults to form
sacrum. [7] Distal caudal vertebrae with elongate
anterior zygapophyses. [8] Length of humerus great-
er than half length of femur. [9] Antebrachium elon-
gate, carpalia reduced, and unsuited for extensive
mediolateral movement. [lo] All 3 metacarpals ap-
proximately equal in development. [ l l ] Digits of
manus elongate and approximately equal in length.
[12] Length of pubic boot less than half length of
femur. [I 31 Ischia distally expanded and ventrodis-
tally recurved. [14] Pelvic canal broad. [I 51 Tibia-
astragalus longer than femur. [l6] Metatarsal I11 con-
stricted proximally. r1713rst phalanx not present
in pes, [18] phalanges of fourth digit unusually ab-
breviated. [19] Pedal unguals ungulate.
 JOURNAL OF VERTEBRATE PALEONTOLOGY, VOL. 10, NO. 2, 1990
However, many, if not all, of these characters need to
be revised to diagnose the Ornithomimidae adequately
and to include Elaphrosaurus (Janensch, 1925) and
Archaeornithomimus. Harpymimus (Barsbold and
Perle, 1984), from the Aptian-Albian Shinehuduk For-
mation of Southeastern Mongolia, and Garudimimus
(Barsbold, 1981) should also be considered ornithomi-
mids. A light edentulous skull with a large orbit was
also present in Oviraptor, which possesses numerous
other characters excluding it from the ornithomimids.
In Deinonychus, the maxilla does not border the nares
(Ostrom, 1969). The ornithomimid diagnosis should
be revised to include the possible presence of teeth,
which may have been present in Elaphrosaurus (Ja-
nensch, 1925) and were definitely present in Harpymi-
mus, and five or six sacral vertebrae, with five present
in Archaeornithomimus and Gallimimus. Most of the
presacral vertebrae of Compsognathus also lack pleu-
rocoels (Ostrom, 1978). Elongate prezygapophyseswere
present on the distal caudal vertebrae of many thero-
pods, such as Allosaurus. The length of the humerus
is greater than half the length of the femur in Comp-
sognathus (Ostrom, 1978). Compsognathus also ex-
hibits a ventrodistally recurved ischium with a distal
expansion. Metacarpal I of Harpymimus is signifi-
cantly shorter than metacarpals I1 and 111. In Comp-
sognathus, the tibia is also longer than the femur (Os-
trom, 1978). Metatarsal I11 is somewhat constricted in
Elaphrosaurus and Harpymimus, but the proximal end
is visible from an anterior view when the metatarsals
are articulated, unlike the other ornithomimids. This
extreme metatarsal I11 proximal constriction, however,
also holds for Tyrannosaurus. In Archaeornithomimus,
characters 10 and 16 do hold, but 1 is unknown. Char-
acter 17 presumably refers to the absence of the first
digit rather than the first phalanx.
In the diagnosis for Harpymimus, Barsbold and Perle
(1984) stated that the bones are very similar to those
of ornithomimids, but they cited the presence of teeth
as a definitive characteristic for the new family Har-
pymimidae. This feature is not diagnostic since teeth
were probably present in Elaphrosaurus and would be
expected in a conservative ornithomimid, so Harpymi-
mus should be included in the Ornithomimidae.
I
Status of Archaeornithomimus a@nis
i 1~
I 11
11 11
A recurved pedal ungual, two distal caudal verte-
brae, a vertebral centrum, an astragalus, and two in-
I
I
complete metatarsals from the Lower Cretaceous
Arundel Formation of Maryland were made the syn-
111 1 Ill
types of Ornithomimus afinis by Gilmore (1920). This
material was illustrated by Gilmore (1920, 1924) and
I1 l 1l Lull (19 1I), who described the specimens as the cotype
II of the ornithopod dinosaur Dryosaurus grandis Lull
(19 11). However, Gilmore (1920) drew attention to
I l1
I the probable ornithomimid characters of the- m-aterial Barsbold, R . 198 1. The edentate carnivorous dinosaurs of
and, since the name grandis had already been used,
I renamed the material Ornithomzmus a8nzs. ~ u s s e l ~
(1972) moved the material to his new genus Archaeor-
I
nithomimus as A. afinis (Gilmore, 1920) on the basis
of a recurved pedal ungual. However, the pedal unguals
of Archaeornithomimus asiaticus are straight as in oth-
er ornithomimids, and the recurved Asian phalanges
are probably referable to Alectrosaurus, the only other
theropod known to be present in the Iren Dabasu fauna
(Gilmore, 1933). The recurved pedal ungual from the
Arundel Formation of Maryland (USNM 6 107; Gil-
more, 1920:fig. 76) is probably from the theropod
Dryptosaurus (?) that is also described from there (Gil-
more, 1924).
The Arundel material, except for the two partial
metatarsals, is insufficient to characterize beyond the-
ropod incertae sedis. The two metatarsals are hollow,
but the proximal ends are not present, so it is not
possible to determine whether or not the incomplete
metatarsal I11 is constricted at that end. On this basis,
these two elements can only be referred to a coeluro-
saurian grade.
Status of Elaphrosaurus bambergi
Nopsca (1928) classified Elaphrosaurus as a primi-
tive ornithomimid, and Russell (1972) and Galton
(1982) listed the following characters as ornithomimid:
1) a straight humerus with a small deltopectoral crest,
2) long prezygapophyses on the distal caudal vertebrae,
3) abbreviated phalanges on the fourth toe, and 4) the
form of the presacral and proximal cau_rl,aL-wrteb~ae.
An additional feature that Elaphrosaurus shares with
the Ornithomimidae is the distal expansion of the is-
chium.
ACKNOWLEDGMENTS
We thank Dr. Eugene Gaffney and Charlotte Holton
(AMNH) for their invaluable assistance on this project,
and Drs. Michael Somers (University of Bridgeport),
Elizabeth Nicholls (University of Calgary), Halszca
Ozmolska (Polish Academy of Sciences), David Weis-
hampel (Johns Hopkins University), and Philip Currie
(Tyrrell Museum of Paleontology, Drumheller, Alber-
ta, Canada) for their comments on this paper. Drs.
Gordon Edmund, Andrew Leitch, and Chris Mc-
Gowan (ROM) permitted study of the material and
use of equipment at the Royal Ontario Museum. Dr.
Robert Grant (University of Arizona) and Steve
Walmsley arranged and provided lodging while in To-
ronto. Donald Livingston, Jr. (Modem Languages De-
partment, Arizona State University, Phoenix, Arizona)
translated articles from Russian. The manuscript was
typed by Jennifer Monahan (University of Bridgeport).
This project was supported by the Biology Department
of the University of.Bridgeport.
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Received 21 September 1987; accepted 11 August 1989.