Cretaceous Late Coniacian Santonian
Saurischia Theropoda Tetanurae Coelurosauria Oviraptorosauria
Barun Goyot Formation, Omnogov, "Djadochtinskaya" Svita, Ovorkhangai Mongolia; Iren Dabasu Formation, China
< 1 meter
Genus - Typespecies - Skull
seems to have been a bird-like, running dinosaur. It was a lightweight,
long-legged creature with a short, deep head, toothless beak, long,
curved neck, well-developed tail, and short forelimbs. No fossil
feathers have been found, but the rough ridge on one foramen bone
resembles a modern bird's attachement for flight feathers
3 partial skeletons.
Kurzanov, S.M. (1981) An unusual theropod from the Upper Cretaceous of Mongolia Iskopayemyye pozvonochnyye Mongolii (Fossil
Vertebrates of Mongolia). Trudy Sovmestnay Sovetsko-Mongolskay
Paleontologiyeskay Ekspeditsiy (Joint Soviet-Mongolian Paleontological
Expedition) 15:39-49. Nauka Moscow, 1981
Translated by Catherine Siskron and Samuel P. Welles minor editing by Matthew Carrano, January 2000
During the exploration of the Joint Soviet-Mongolian paleontological
expedition in the summer of 1973, at the Udan-Sayr (southern Gobi)
location, was found a fairly complete skeleton of a bird-like theropod,
which belonged to the new genus and species Avimimus portentosus. With the exception of a crushed skull fragment, all the remaining bones are very well preserved.
The location of Udan-Sayr is 75 km. south of Hovd-somon
Ubur-Hangayskaymak, in the foothills of the Gurvan-Sayhan mountain
range. The osseous layers of this location are in the red colored sands
and sandstone with rare interlayers of clay and conglomerates. The
visible thickness of the deposits is approximately 15 m. The age of the
deposits can be established as Nemegetinsk. This is indicated by the
presence in it of teeth of Tarbosaurus, a carnosaur, known from the deposits of that age from various locations in the Southern Gobi.
Besides the Udan-Sayr location, Avimimus
remains were also found in the Shara-Tsav location, 7 km north from the
Bayshin-Tsav in the southeastern Gobi. Besides this, here was also
found an almost complete skeleton of the ornithomimid
Gallimimus (classification by Barsbold) and the skull of the sauropod
which are also highly characteristic for the Nemegetinsk deposits of
the southern Gobi. The fauna found in this region clearly indicates
that the deposits in this location are of the Nemegetsk suite.
The discovery of Avimimus
is of particular interest. On one hand this is the first dinosaur that
has such clearly expressed bird features, in such large numbers, on the
other it is also the first among the theropods with such unusual
structure of the pelvis. Such a combination of unique features places Avimimus in a category all its own among the dinosaurs and places it in a new family, [Avimimidae], fam. nov.
Family [Avimimidae] Kurzanov, Fam. Nov.
Osmolska, Currie & Barsbold, 2004 placed A. portentosus as a basal member of the oviraptorosaur clade
theropods, 1-1.5 m in length. The skull is small with relatively large
orbits. The frontal and parietal bones are fused. The cervical
vertebrae increase noticeably in site as they approach the trunk. On
the last two cervical and the first three dorsal vertebrae, the
hypapophyses are developed. There are no fewer than 11 cervfcal
vertebrae. The pelvis is wide. The ilium, ischium and pubis are fused
around the acetabular (orig. “vertlugian”) cavity. The posterior
branches of the ilium are directed almost horizontally. On the femora
are developed special additional condyles for the articulation with the
fibula. The latter are highly reduced and are fused with the tibia at
their distal ends. The bones of the distal row of the tarsus are fused
with the metatarsus. The second and fourth metatarsals fuse proximarly
with each other, the fifth fuses fully with the fourth. the phalanges
of the external toes are highly shortened.
Family composition .
Single genus Avimimus gen. nov.
The closest to avimimids are members of the family [Ornithomimidae - Ornithomimosauria]. The number of cervical vertebrae in ornithomimids is the same as that of other theropods, and does not rise above 10, while Avimimus
has no fewer than 11. Until now hypapophyses were not observed on
cervical and dorsal vertebrae among theropods, but they are present in
the case of Avimimus.
The same is true of
the fusion of the parietal and frontal bones. One of the main features,
which differentiates avimimids not only from ornithomimids, but also
from other theropod families, is the structure of the pelvis and the
The rather wide pelvis of Avimimus,
if one is to judge by the distance between the acetabular cavities – in
relative dimensions, it is almost twice as wide as that of
ornithomimids. Practically all theropods have an ischium that is
directed posteriorly, and only in Avimimus is it projected anteriorly and ventrally, parallel to the pubis.
Also, all theropods, with the exception of Avimimus, have pubic bones which participate in the formation of the acetabular cavity. Another first among theropods is that Avimimus has an additional third condyle on its femur. Undoubtedly, among features peculiar to Avimimus
is the full fusion of the proximal row of bones of the metatarsus, the
proximal fusion of metatarsals II, IV and V, and the highly shortened
external toes, Such features are also completely unknown among
Avimimus Kurzanov, gen. nov.
Name of genus .
From avis, Latin, bird and mimus, Latin, to mimic (?),
Type species .
Avimimus portentosus, Upper Cretaceous, Nemegetinsk suite, Mongolian National Republic.
Species composition .
Upper Cretaceous, MNR.
Avimimusportentosus Kurzanov, sp. nov.
Name of species .
From portentosus, Latin, unusual.
PIN, #3907/1, incomplete skeleton; MNR, Udan-Sayr, Upper Cretaceous, Nemgetinsk suite.
the holotype, paratype #3906/1 from the location Shara-Tsav, individual
fragments of the postcranial skeleton; MNR, Upper Cretaceous,
Almost all the vertebrae, girdle of the forelimbs, and pelvis were
found separated, while the rest of the bones were found articulated.
They were found in the perimeters of a rather small area, and
undoubtedtly belong to the same specimen, the most characteristic in
size. At the Udan-Sayr location, Shara-Tsav, were discovered at the
same time a large quantity of individual bones from other specimens of Avimimus, practically all of similar dimensions.
Skull. Only the badly crushed skull roof was preserved along with part
of the occipit and the lower part of the left wall of the cranium. Some
bones were displaced. However, the fact that they mostly preserved the
shape and surface, allowed a partial reconstruction of the skull of Avimimus.
The skull is small, its length from the top of the upper occipit to the
begining of the conjectural nasal openings is approximately 50 mnm. In
addition to this, the possible size of the orbit is no less than 25 mn,
that is the orbit, as in the case of birds, is proportionally very
large, and started also as in the case of birds almost immediately
behind the nasal opening. The upper temporal cavities are very small,
their greatest diameter was not above 10 mm. Judging by the preserved
posterior edge, the nasal openings were narrow and elongated in length.
The frontal and temporal bones are
fully fused with each other, sutures with adjoining bones are partially
obliterated. They form the convex skull cupola and the medial parts of
the upper temporal fenestrae. The single fronto-temporal bone extended
anteriorly almost to the nasal openings.
The postorbital bones form the posterior half of the upper edge of the
orbit. Medially they become quite thick. Between them and the
fronto-parietal on both sides there is a bend. Anteriorly they rest on
the posterior part of the lacrimals.
Lacrimals comprise the anterior half of the upper part of the orbit.
Narrow, short, apparently, with a descending process. Anteriorly they
rest on the nasals.
nasals were preserved only the proximal part of the left nasal with the
surrounding smooth cavity, gradually passing, apparently, into the
nasal opening. It is important to note that the supposedly long, narrow
nostrils were located not so much on the lateral as on the dorsal part
of the skull, in a position very close to that of the birds.
The upper occipital bone occupied almost the whole upper half of the
occipital curface. In the middle of its upper edge is located a small,
symmetrically located swelling. As a whole the bone is wide and flat,
and forming a considerable part of the occipit, makes it also flat and
even, as in birds. Ventrally the supraoccipital fuses firmly with the
exoccipitals, anteriorly and laterally with the prootic.
The highly fragmented prootic, which forms the left wall of the
endocranial cavity, is visible only from the medial side. A
considerable part of it is occupied by the lower half of the large
circular cavity, located directly inside the osseous labyrinth (along
its edge pass the semicircular canals) and the corresponding recessus
intracucticus of some theropods (Kurzanov, 1976). An analogous formation is noted also in the cranium of the Upper Cretaceous sea bird Hesperornis regalis (Marsh, 1880),
in which the endocranium as a whole is similar to that of theropods
from the famny Itemiridae. This structure is present also in
contemporary birds, although it is relatively smaller in size, and
accommodates special excrescence of the cerebelum – flocculi.
Apparently, an analogous function was performed by the otic cavity of Avimimus, and there is no basis to associate its existance with the lymphatic or venous systems, as was supposed earlier (Kurzanov, 1976).
Immediately under the otic cavity is located a horizontally elongated
cavity, on the bottom of which are located two openings for the otic
nerve, through the anterior of which passes the vestibular, and
posteriorly the cochlear nerve. Directly anteriorly and somewhat above
it is located the opening for the facial nerve. Posteriorly from the
otic capsule, the endocranial cavity remains equally high, and the
medial curvature of the otic bone indicates that it becomes narrower.
Ventrally, immediately behind the otic capsule is located a narrow high
fissure of the jugular opening, and directly in front of it a snlall
opening for the hypoglossal nerve.
The shape and relative dimensions of the preotic bones, and also of the
fronto-temporal allow to speculatively judge the shape of the brain.
The cupola-shaped and wide roof of the skull, and the lateral curvature
of the anterior part of the preotic bones, indicate an unusual
development of the large hemispheres of the brain.
If one is to judge by the height of the preotic bone, the endocranial
cavity is relatively large and high behind the otic capsule, as is the
cerebellum. Such a structure of this part of the cranium suggests the
possibility that the height of the endocranium may be connected with
the double curvature of the elongated brain, which is characteristic
The small pieces preserved of the
orbitosphenoid bones carry large foramina for the optic nerves. The
orbitosphenods are almost at a right angle and meet in the middle,
where they contact the unpaired ethmoid. The latter at this point
becomes thin, forming the fissura craniofacialis, which separates the
facial part of the skull from the cranium. Further forward the ethmoid
suddenly thickens. The sutures between the orbitosphenoids and the
ethmoid (?) are obliterated.
Vertebral column. Only seven cervical vertebrae have been preserved (2,
4, 6, 8-11), and also nine dorsal vertebrae, in sequence. The vertebrae
of both sectors are platycoelous, and the articular surfaces are
of the cervical vertebrae slightly increase toward the trunk. The
length of the axis is 17 mm, that of the ninth vertebrae is 24 mm. The
change in the size of the vertebrae allowed us to establish their
approximate number: a minimrirn of 11 cervical vertebrae.
The body of the vertebra is elongated. In the middle is located a
fairly large oval opening, which leads to the cavity inside the body.
On vertebrae from 6th to the ninth an additional opening is located on
the anterior edge.
The lower surface of the
bodies, starting with the axis, with a pointed crest in the middle,
which gradually flattens, and then on the tenth vertebrae again becomes
pointed and remains pointed on all the dorsal vertebrae. On the
anterior edge of the supposedly last (11 th) cervical vertebra this
crest immediately becomes the hypapophysis, fairly thick, with a
Judging by the site of this
hypapophysis, it is perfectly logical to suppose that a somewhat
smaller one was found also on the tenth vertebrae, the ventral surface
of which was destroyed. The arches are low, with very short transversal
processes, immediately under which is found a cavity with three
The articular surfaces of the majority of
the cervical vertebrae are not displaced in relation to one another;
and only the fourth (possibly, that this was true also of the 3rd and
5th which were not preserved) the anterior articular surface is raised
dorsally in relation to the posterior, creating the necessary natural
curvature of the neck.
vertebrae. In differentiation from the cervicals, are of equal
dimensions. The body of the first vertebrae in cross-section have an
elongated-oval shape, while close to the tail they become rounded. Up
to the 9th dorsal vertebra a pointed crest is present on the ventral
surface. It is especially well expressed on the dorsal vertebrae, where
on the first and third it turns into a hypapophysis on the anterior
edge, a very large one on the first.
facets of the rib head are well expressed and gradually move from the
anterior edge to the middle of the body (first vertebra) to the level
of the anterior articular processes, to the arch (starting with the
seventh). In the articulated state the dorsal vertebrae form a gently
sloping arch. The lateral sides of the body at first are almost flat,
then become more and more curved in the anteroposterior direction. The
neural canal is relatively large, its diameter is approximately two
times smaller than than the body af the vertebra. The anterior
articular processes are short, barely project beyond the surface of the
As is apparent from the description, the most characteristic features of the vertebra of Avimimus
are: considerable increase in the length of the cervical vertebrae and
the first dorsal vertebrae, the appearance of which is probably
connected with the strenghtening of the m. rectus capitis.
Pectoral girdle. The scapula and the coracoid are fused without a
visible suture. Only a wide strip, slightly roughened, which passes
vertically from the middle of the upper edge of the glenoid cavity,
indicates that place (where the suture ought to be). The scapular part,
just as is the case in all theropods, is wide ventrally and slightly
more sharply narrowed in the dorsal part. What is important is that
there are no sharply expressed boundaries between these parts, which is
most characteristic of birds and theropods from the family
The glenoid cavity is fairly
narrow, elongated, directed straight down, thus indicating the position
of forelimbs vertically below, directly under the body. The
supraglenoidal thickening is not very well expressed. The most
characteristic peculiarity of the coracoidal part is, apparently, its
elongated shape. Thanks to this it is possible to approximately measure
the angle between the scapula and the coracoid – a blunt angle of
approximatety 160° which is quite characteristic for running birds,
which is also true of the fact that the coracoid is elongated. In front
and below the coracoidal opening is located a large pointed tubercle,
the existence of which is probably connected with the attachment of m.
biceps brachii, the most powerful flexor of the antebrachium. Directly
below this tubercle is located a large concave surface – most likely
the area of attachment of m. coracohumeralis.
Humerus. The humerus of Avimimus
is noticeably differentiated in structure from that of small theropods.
The deltopectoral crest is well developed and its length is greater
than 1/3 the length of the humerus. On its dorsolateral surface, along
the entire length of the thickening, are noted the beginnings of m.
deltoideus, which performed the function of pronation and supination of
the shoulder. On the distal end of the deltopectoral crest remained a
small swelling from the m. pectoralis.
proximal end of the humerus dorsally was preserved a rough, slanted,
wide strip of m. coracohumeralis, which passed from the upper edge of
the deltopectoral crest in the direction of the tuberculum mediale. On
the dorsolateral surface, exactly in the middle, is located a slanted
seam – an approximate line for the attachment of m. triceps brachii,
and quite possibly also of m. latissimus dorsi.
It is possible that the most important feature of the Avimimus
humerus is the characteristic three-segmented distal epiphysis, which
is divided into three unequal parts. The largest of them is the one in
the center, and the one that projects the most is the medial. Most
likely the middle condyle, being the largest, corresponds to the ulnar,
while the lateral corresponds to the radial, which is indicated by its
position in relation to the ulnar condyle and also its elongated shape,
which allows a minor displacement of the proximal end of the radius
during pronation and supination.
The deve!opment of
the third, medial part of the distal epiphysis, which cannot be named a
condyle and does not exist in other theropods, possibly is connected
with the attachment of muscles, which accomplish pronation of the
antebrachium – m. pronator and and flexor – flexor carpi and metacarpi.
A quite similar construction of this sector of the humerus can be found
Immediately above the
central condyle, the distal joint the humerus is noticeably indented,
which apparently considerably increased the flexing possibilities of
the radial joint. Imnediately above was found the beginning of m.
brachialis anticus, the most powerful flexor of the antebrachium. The
external edge directly above the radial condyle is greatly thickened
and rough, indicating the attachment of m. extensor digitorum comnunis.
Immediately below, directly to the lateral condyle, is attached a
supinator, simultaneously also the flexor of the antebrachium.
As can be gathered from the above information, the clearest muscle
imprints are connected mostly with the flexors of the antebrachium,
flexors and extensors of the hand. The approximate, due to the
incomplete preservation of the forelimbs, analysis of the shoulder
muscles indicates a definite tendency to the amplification and possibly
predominance of the flexing-extending motion of the limb. The ulna, of
which we have only the compressed proximal epiphysis with a relatively
low olecranon process. On the external side, closer to the anterior
edge, near the joint is located a triangular rough surface for the m.
brachialis anticus. From the internal side, adjoining the radius, the
ulna is slightly concave.
Pelvic girdle. The hind limbs of Avimimus
are long and slender, with a tibia considerably enlarged relation to
other sectors. The length of the femur has the ratio to the tibia and
to the length of the metatarsus of 1:1.37:0.81. The distal parts of the
limb sectors are noticeably thinner than the proximal, just as the
whole limb becomes thinner distally. All articular surfaces are very
The femur, is
insignificantly curved, with very well defined joints. Its length is
188 mm. The head is ball shaped, with a rounded cross-section thinner
than the diameter of the head and neck. It is located at a right angle
to the axis of the femur, so that just like the shape of the head, it
is highly characteristic for the majority of birds. A little above the
middle of the medial surface of the head is located a small indentation
for the attachment of ligamentum teres.
large greater trochanter surpasses the head 1.5 times in width. It is
separated by a deep, narrow fissure from the lesser trochanter, which
is a high, subquadrate in cross-section, projection, which does not
project up beyond the surface of the greater trochanter. On the lateral
side of the latter is located a triangular outgrowth, apparently
connected with m. ilio-femoralis. The distal condyles of the femur are
large, the external is more developed, thus establishing the position
sector of the limb – somewhat medially, under an obtuse angle to the
The intercondyle furrow is wide, with a
flat bottom, forming a large area for the cruciate ligaments of the
knee joint, analogous to that of birds. A remarkable feature of Avimimus,
which is found for the first time.among dinosaurs in general, is the
development of a special additional supracondyle on the external
condyle of the distal epiphysis of the femur, which forms the joint for
the reduced fibula. The proximal head of the latter moved in the furrow
between the additional supracondyle and the lateral condyle.
Apparently, this is connected with the reduction of necessity in the
rotary motions and the strenghtening of the joint at the expense of the
full transfer of support to the tibia.
From the internal condyle, along the anterior side of the femur, passes
a low, but clearly defined, crest, apparently one that marked the line
of attachment of m. femorotibialis. On the posterior side, immediately
above the distal joint is located a relatively deep fossa, which is
also found in birds – fossa poplitea. A moderately well developed
fourth trochanter is displaced proximally.
The tibia, 257 mm long, is in every sense of the word a tibiotarsus due
to the full fusion with proximal elements of the tarsus. Such a
structure was previously observed only among birds. There is no condyle
between them, and they fuse smoothly. The cnemial crest is large,
laterally triangular in shape, suggesting strong development of the
knee flexor (partly m. femorotibialis), which is confinned by the
clearly expressed traces of flexors on the femur.
below the cnemial crest on the external side is developed an elongated
right-angled processus fibularis. The trunk of the tibia is absolutely
straight, semi-circular in cross-section. The distal end with two
almost equal in size condyles, is slightly turned on its long axis
inward. There are hardly any signs left of the fusion between the crus
and tarsus, although on the external side of the tibia is clearly
visible the distal end of the fused fibula.
articular surface of the intertarsal joint is enlarged, allowing the
turn of the metatarsus to at least 120°. The distal condyles project
forward to almost half the thickeness of the bone, and above them is
formed an articular area, which is also reminiscent of that of birds.
Their joint width is considerably greater than the width of the trunk,
which is reminiscent of the same region in birds.
The fibula, as was mentioned, is highly reduced. Approximately along
one-third of its length is fused to the tibia. The proximal end is
flat, elongated – triangular in shape with a small process for the
attachment of the fibula.
metatarsus is almost analogous to that of birds, for the fusion of
distal elements of the tarsus and metatarsus occurs, and as well as the
fusion of the elements of the latter with each other, but only
proximally. This, it is true, does not affect the rudiment of the 5th
tarsal, which fuses fully, and the 3rd which is proximally reduced and
ends approximately in the middle of the tarsus.
posteriar proximal surface of the tarsus due to the fusion of the
metatarsals II and IV, forms a monolithic area with a rough shagreene
from the attached m. gastrocnemius and short toe flexors. Although a
process similar to that of ground birds is not formed here,
nevertheless the articular surface for the attachment of muscles for
the flexing of toes, the crurotarsal and the knee joints are noticeably
enlarged in comparison with ornithomimids. The length of the tarsus is
The toes are shortened at
the expense of the length of the distal phalanges (this is especially
true of the outside toes). Compared to the ornithomimids the basic
phalanges are also shortened, which is most visible in the second toe.
The opposite is true of the middle toe, where the basic phalanx is
considerably elongated, and the others remain of about the same length
as those of the ornithomimids.
This is why the
third toe is considerably longer than the external ones, especially if
one is not to take into account the claw phalanges. In this case, with
the normally articulated metapodials, the second toe reaches only the
middle of the first phalanx of the third, and the fourth did not reach
to the articular fossa on it. The articular surfaces of the phalanges
of the external toes are peculiarly arranged.
example, on the fourth toe of the right foot the external part of the
articular surface of each phalanx is slightly more developed than the
internal, allowing the toe as a whole to have some curvature to the
inside. An analogous situation can be seen in the case of the second
toe of the left hind limb. It develops that the foot as a whole, due to
the curvature of the external toe, narrows toward the middle one. With
the exception that in the shape and development of articular surfaces
the phalanges of Avimimus are quite similar to those of ornithomimids.
this does not apply to the claw phalanges of the outside toes. The
latter are differentiated by a ventral suiface that is pointed
laterally and from such a peculiarity are reminiscent of claws. Besides
that, they are relatively compressed laterally and above the claw
phalanx of the third toe. The latter has a rather common shape – it is
smaller than the external, symmetrical, with a flat lower edge, hardly
curved at all. Also the claw phalanges of the external toes are only
Pelvis. Only the
posterior parts of the ilia, starting at the edge of the acetabular
cavity, remain. The edge of the acetabular cavity is thick at the
expense of the development in this place of a thick antitrochanter
which is triangular in shape if viewed laterally, with a slightly
convex surface. On deep bending of the femur the antitrochanter
articutated with the greater trochanter and the neck, giving support
for the femur, for in this case the extensors developed a considerable
lateral motion, which aimed to push out the head from the acetabulum,
which was resisted by the antitrochanter.
lower posterior edge of the femur is practically straight, only in the
middle can be noted a small tubercule, from which starts a large
triangular roughness of the flexor of the tibia, m. flexor tibialis.
Slightly toward the anterior, on the inner side are preserved two fully
fused processes, which connected with the sacral vertebrae. In the most
posterior part of the lower edge is visible the roughness of the
supinator of the tibia, m. ilio-fibularis. All of the lower part behind
the acetabulum is considerably thickened in relation with the majority
of theropods, suggesting an intensive development of the extensors of
The ischium is rather
common for theropods, with a very well expressed surface of the
acetabulum. The posterior process, for the articulation with the ilium,
and the anterior are developed practically to the same extent and
jointly form slightly less than half of the acetabular cavity. From the
lateral side, is proximally devetoped a wide, short and rough crest,
which marks the area of attachment of the m. puboischiofemoralis
internus. The distal parts of the ischium were not preserved, but if
one is to judge from the character of the break, apparently became flat
and turned into flat lamellae, which contaced the posterior distal
surface of the pubis.
has a short, for theropods, but wide "boot". Distally, the pubic bones
diverge very widely, relatively wider than other theropods with the
exception of Segnosauridae. For example, wider than in the ornithomimid
Gallimimus bullatus, specimen PIN, #551/54, which surpasses Avimimus in size almost double.
from the middle the right and left pubic bones fuse. Directly from this
spot they become elongated, triangular in cross-section, whereas the
posterior edges are turned outward and the pubic become "open" to the
posterior. In this manner is formed a peculiar receptacle for the
distal ends of the ischia. The anterior edges, to the contrary, in
differentiation from other theropods, are closely approximated and
between them remains only a smarl depression. On the break it is quite
visible that the proximal pubic bones become much thinner, however
their ends were not preserved.
spite of the fact that the pelvic bones were not found in an
articulated state, and were incomplete, their position in relation to
one another can be properly reestablished. In as much as the ischial
process of the ilium and the iliac process of the ischium were
preserved, then they sufficiently clearly establish the position of
these bones, as well as the maximal posterior position of the ilium. If
one is to take into consideration of the posterior part of the ilium
approximately parallel to the spine, then the direction of the ischium
to it is at almost a 90° angle, and if one is to take into
consideration that there is no precise contact between the pelvic
bones, then the angle may have been greater. Such a situation is
completely uncharacteristic for theropods, where the angle between the
spine and the ischium ranges from 30° to 50°.
It turns out that the ischia of Avimimus
are turned anteriorly by approximately 50°, that is in a positlon that
is more characteristic for pubes. The latter were located anteriorly,
in their usual place, more or less parallel to the ischia. In as much
as in specimen #3907/1 from Udan-Sayr were not preserved proximal parts
of of the pubes, then it was only possible to establish aproximately
the character of their articulation with the pelvis after examining
additional material from the location of Shar-Tsav.
Among the bone fragments which belonged to different Avimimus
skeletons, was discovered the proximal part of an ischium, which was
fully fused with another pelvic bone. Jointly they formed the upper
anterior of the acetabulum. After the comparison of this fragment with
those of specimen #3907/1, it became clear that the pelvic bone in
question was a small piece of the ilium. In this manner the acetabular
cavity was formed by the ischium and ilium, while the pubis had no
participation in its formation.
In the same manner
that the anterior position of the ischium, the exclusion of the pubis
from the acetabular cavity has no analogues among the dinosaurs in
general. Due to this fact, and to the absence of proximal ends of the
pubes, the manner of its articulation with the pelvis is not very
clear. For the time being, the most likely supposition is the
following: approximately at the point of contact of the ischium and the
ilium on the external side is located a small protuberance, to which
possibly was attached, with the help of ligaments, the pubis.
Indirectly this type of attachment is supported by the rather sudden
decrease in the thickness of the pubes. At the present time the above
described pelvic structure appears likely and doubts can be had only in
relation to the articulation of the pubes.
Geologic Age . Upper Cretaceous, Nemegetinsk suite, Maastrichtian.
Location . Mongolian National Republic, southern and southeastern Gobi.
As is apparent from description, the pelvis of Avimimus
has a rather unusual construction, which until now has not been
observed in theropods, or dinosaurs in general. This first of all is
true of the up and down direction of the ischia, which together with
the ilia formed the acetabulum. To a lesser degree this also applies to
the width of the pelvis.
along the distance between the centers of the acetabular cavities, it
seemed equal to 10 cm. For an animal of such small proportions as Avimimus,
the pelvis is relhtively much wider than in the majority of theropods,
with the exception of the representatives of a new family,
1979). Such a shape of the pelvis unquestionably adds stability to the
body during bipedal locomotion, and is characteristic, continuing the
comparison with birds, of running and climbing animals.
In as much as there are no climbing features found in Avimimus, the majority of the other peculiarities of the sketeton indicate it to be a running form. It is known (Gambaryan, 1972) that the greatest development in running is achieved in the extensors of the pelvo-femoral, knee and tibia-foot joints.
It is precisely this type of development that we find in Avimimus:
an unusually thick, massive wing on the ilium, a large cnemial crest, a
large area for the posterior part of the metapodials. The strenghtening
of the extensor musculature of the femur occurred at the expense of the
increase of the area of its attachment on the posterior branch of the
ilium, for which the latter occupied at most a horizontal position.
The place for the attachment of the extensors also has great
significance for increasing the speed of locomotion. The closer it is
to the proximal sector of the linb, and the shorter are these proximal
sectors, the greater will be the gain in speed, and the greater the
loss in strength. This applies to both the relative shortening of the
femur of Avimimus, as well as the proximal displacement on it of the fourth trochanter.
However due to the turn of the ischia forward occurred some weakening
of the extending capability of m. puboischiofemoralis. With such an
arrangement the posterior muscles change the direction of their motion
and the extensor-adductor becomes mainly an adductor.
as was already mentioned, a considerable strengthening in the femur
extensors compensated a similar weakening. Otherwise the turn of the
ischia forward should not introduce basic changes in the mechanics of
the pelvo-femoral muscles. One of the possible explanations of the
unusual arrangement of the Avimimus pelvis makes it similar to the open pelvis of birds, which is adapted to the laying of relatively large eggs.
Approximately the same types of tendencies for adaptation to running
are also observed in the structure of the foot, the metatarsal bones of
which have the thinnest, most slender features, which is characteristic
for the distal sector of the limbs of running birds.
adaptation to running is also reflected in the relative elongation of
the first phalanges of the third toes and the shortening of the toes
themselves, which is particularly noticeable on the second.
Undoubtedly, this cannot yet be called a reduction, but apparently it
is already beginning to take place, in as much as the second metatarsal
is smaller and thinner than the fourth.
already noted that the reduction of the toes went quite far. Thus, if
in the forward position of the foot the support fell on all the
phalanges, with the exception of the first exterior toes, then in the
back position – only an the 2 nd-4 th-phalanges of the third, while the
external, most likely, did not touch the ground at all. If one is to
take into consideration some curvature of the external toes toward the
middle one, then the support in the forward position of the foot was
very narrow. A
ll this to some degree is reminiscent
of the foot arrangement of the ostrich. It is interesting to note that
in differentiation from running birds the claw phalanges of the hind
limbs of Avimimus are practically straight, and the flexor
tubercles are not developed on them, but there is no viable explanation
for this at the present time.
However in spite of the similarities that were stressed between Avimimus and running birds, neither Avimimus, and even less so ornithomimids, are not running animals in the same sense as the contemporary Gradientes. Even in Avimimus,
the closest to the latter, the limb proportions are far from being the
same, which is also true of the development of the joints and
musculature of the knee and crurotarsus, which are not as well
For example, the ratio of length of the femur to the length of the tibia and to the metatarsus in Avimimus
is 1:1.37:0.81, while in rheas it is 1:1.84:1.70. In ornithomimids,
which we are used to considering running animals, as in the case of
ostriches, all the features related to this are expressed more weakly
than in Avimimus.
One one hand, is clear
the conditional aspect of the definition of “running” animals, on the
other would it not be productive to look for analogies of a different
type to compare the way of life of the two groups. Especially since in
all the previous views of manner of life of the ornithomimids, only an
insignificant role was assigned to the fore limbs, which both in
ornithomimids and in Avimimus are well developed and must have served some function.
It is possible to say that, up to the present, Avimimus
is the most bird-like dinosaur of all the theropods and of all the
known dinosaurs as a whole. Such a degree of of development of
bird-like characteristics in a theropod allows one to speak not only
about the accepted kinship of theropods and birds, but, what is more
important, about the possible formation of bird anatomy among the
running forms of dinosaurs.
Such thoughts were
expressed a long time ago (Furbringer, 1888; Broom, 1913) and, it
seems, find confirmation in the skeletal structure of Avimimus. However, it is important to note that a considerable similarity is observed between Avimimus
and birds in general, and with running birds only in some aspects. This
is why it would seem that such similatities in structure would indicate
kinship ties, rather than analogues in manner of life.
Apparently, supposition concerning the origin of birds from running, bipedal theropods is correctly made. Avimimus
is not an ancestor of birds, not even probably the running birds, in as
much as in the structure of the pelvis are noted features which are not
characteristic for them (the exclusion of pubes from the acetabulum,
the turning of the ischia forward). Nevertheless the very fact of the
formation of numerous typical bird features among dinosaurs suggests
another proof of the possible origin of bitds both from the running
bipedal theropods as well as the climbing forms, which in recent years
is the dominant point of view.
features we should mention the cupola-shaped skull, formed by fused
frontal and parietal bones, the close positioning of orbits and nasal
openings, the presence of the third additional condyle on the femur for
articulation with with the fibula, a wide pelvis, fusion of the tarsus
and the metatarsus, the presence of the hypapophyses on the last
cervical vertebrae and the first dorsal, reduction and fusion of the
fibula with the tibia, and also a series of other, less significant
features. The formation of such features is hard to explain as parallel
development, but rather seems to indicate kinship wfth birds. In
addition, the more abundant are these types of features, the closer in
the bunch of parallel lines of development will stand the forms being
compared. It is possible to find more bird features in Avimimus than in the well-known
and if it were not for the time factor, then there would be less basis
for including the latter in the ancestral group for birds.
At the present time it is still premature to talk of some definite kinship ties of Avimimus
with other theropods, since the comparison even with the closest of
those ornithomimids is expressed mostly in the structure of hind limbs,
and carries, most likely, a clearly converging character. Much light
could be thrown on this question by a find of a more complete skull and
The non-reptilian structure, and possibly manner of life, of Avimimus
is suggested by both the high level of exchange of matter,
characteristic only for animals with a constant body temperature,
although in the present case are absent facts indicating a
temperature-conserving cover, which are crucial in the resolurion of
Source: Polyglot Paleontologist