Biological Journal of the Linnean Society. 8: 91-182.

With 36 figures

J une 1976

Archaeopteryx and the origin of birds

JOHN H. OSl’KOM

Accepted for publication October I975

The question ot the origin of birds can be equated with the origin of Archueopte,:vx. thc oldest
known bird. Analysis of the five presently known skeletal specimens of Archaeopteryx. and
comparison with the skeletal anaromv of the several reptilian groups that have heen proposed as
possible ancestors of birds (Ornithopoda. Thcropoda. Pscudosuchia and Sphenosuchidac).
confirm the conclusions (long rejected by most suhscqucnt workers) of tleilmann (1926). I.owe
(1935, 1944) and Holmgren (1955). namely. that the skeletal anatomy of Archaeopteryx is
extraordinarily similar to that of contemporancous and succccJing coelurosaurian dinosaurs.
Rejection of these similarities as adaptive structures only (parallel o r convergent similarities),
and therefore of no phylogenetic importance, is hcre considlered invalid. I4eilmann was thc first
to identify the only evidence that has been cited so far for dismissing coelurosaurian-avian
ancestral-descendant relationships. the supposed absence of clavicles in all thcropods, and on
that basis suggested a common Archaeopteryx-dinosaur ancestry among pseudosuchian
reptiles. That evidence is negative and thus inconclusive. and is now known to he false.
With the exception of fused claviclcs and unique ischial morphology, virtually every skeletal
feature of Archaeopteryx is known in several contemporaneous or near-contemporary
coelurosaurian dinosaurs and many of these conditions are unrelateQ specialized features ( t h e
detailed morphology of the manus. metacarpus, carpus. humerus, scapulocoracoid. pes,
metatarsus, tarsus, femur, pubis, ilium, skull and mandibles). The presence of so many derived
characters in common clearly establishes that the closest ancestral affinities ot Archaeopteryx
are with coelurosaurian theropods. There is n o contrary evidence and any other explanation is
illogical.
Ornithopod-A rchaeopteryx ancestral-descendant affinities may be dismissed becausc of the
false “avian” organization of the pelvis in the Berlin specimen of Archaeopteryx and the merely
superficially bird-like construction of the ornithischian pelvis. The suite of specialized
characters unique to ornithischians (e.g., predentary, tooth morphology), that occur even in
Triassic representatives, is further evidence for dismissing close affinity between ornithopods
and Archaeopteryx. The supposed close relationship between birds and pseudosuchians is
judged to be remote at best, due to the completely primitive nature of the few anatomical
features which pseudosuchians have in common with Archaeopteryx. Sphenosuchus. a primitive
and early archosaur, is also a potential avian ancestor, b u t existing evidence consists of primitive
archosaurian features plus a few similarities with certain modern birds. These similarities. which
are present in two groups that are separated from each other by more than 200 million years,
and which cannot be demonstrated in Archaeopteryx, are considered irrelevant to the origins of
Archueopteryx and subsequent birds.
All available evidence indicates unequivocally that Archaeopteryx evolved from a small
coelurosaurian dinosaur and that modern birds are surviving dinosaurian descendants. Stated
simply, avian phylogeny was: Pseudosuchia Coelurosauria - - + Archaeopteryx
- - + ---+ higher
birds.
7 91
92 J . I I. OS'IROM

CONTI'N'IS
Introduction . . . . . . . . . . . . . . . . . . . . . . 93
llistorical rcvicw . . . . . . . . . . . . . . . . . . . . 97
Ilvitlencc o n the origin o f Archueopreryx . . . . . . . . . . . . . 100
'fhe ornithopod evidence . . . . . . . . . . . . . . . . 101
The theropod evidence . . . . . . . . . . . . . . . . . 109
Manus and forelimb . . . . . . . . . . . . . . . . 109
Manus . . . . . . . . . . . . . . . . . 111
Metacarpus . . . . . . . . . . . . . . . . 111
Carpus . . . . . . . . . . . . . . . . . 113
Radius and ulna . . . . . . . . . . . . . . . 113
tlumerus . . . . . . . . . . . . . . . . . 114
1;orelimh summation . . . . . . . . . . . . . 114
Pectoral girtlle . . . . . . . . . . . . . . . . . 115
Scapula . . . . . . . . . . . . . . . . . 116
Coracoid . . . . . . . . . . . . . . . . . 116
Iiindlimb and pes . . . . . . . . . . . . . . . . 119
Pes . . . . . . . . . . . . . . . . . . 119
Metatarsus . . . . . . . . . . . . . . . . 119
Tarsus . . . . . . . . . . . . . . . . . 121
Tibia and fibula . . . . . . . . . . . . . . . 123
Femur . . . . . . . . . . . . . . . . . 123
Ilindlimb summation . . . . . . . . . . . . . 124
Pelvic girdle . . . . . . . . . . . . . . . . . . 124
Pubis . . . . . . . . . . . . . . . . . . 126
Ilium . . . . . . . . . . . . . . . . . . 126
lschium . . . . . . . . . . . . . . . . . 129
Skull and jaws . . . . . . . . . . . . . . . . . 131
Vertebral column . . . . . . . . . . . . . . . . 135
Other skeletal elements . . . . . . . . . . . . . . . 137
Sternum . . . . . . . . . . . . . . . . . 137
Clavicle-furcula question . . . . . . . . . . . . 138
Gastralia . . . . . . . . . . . . . . . . . 139
Summary o f theropotl evidence . . . . . . . . . . . . 140
The pseudosuchian evidence . . . . . . . . . . . . . . . 140
Manus and forelimb . . . . . . . . . . . . . . . . 143
Manus . . . . . . . . . . . . . . . . . 143
Metacarpus . . . . . . . . . . . . . . . . 143
Carpus . . . . . . . . . . . . . . . . . 144
Radius and ulna . . . . . . . . . . . . . . . 145
tlumerus . . . . . . . . . . . . . . . . . 145
Forelimb summation . . . . . . . . . . . . . 146
Pectoral girdle . . . . . . . . . . . . . . . . . 147
Scapula . . . . . . . . . . . . . . . . . 147
Coracoid . . . . . . . . . . . . . . . . . 148
Clavicle and interclavicle . . . . . . . . . . . . 148
Sternum . . . . . . . . . . . . . . . . . 148
Hindlimb and pes . . . . . . . . . . . . . . . . 148
Pes . . . . . . . . . . . . . . . . . . 148
Metatarsus . . . . . . . . . . . . . . . . 149
Tarsus . . . . . . . . . . . . . . . . . 150
Tibia and fibula . . . . . . . . . . . . . . . 152
Femur . . . . . . . . . . . . . . . . . 153
Hindlimb summation . . . . . . . . . . . . . 153
Pelvic girdle . . . . . . . . . . . . . . . . . . 154
Pubis . . . . . . . . . . . . . . . . . . 155
ilium . . . . . . . . . . . . . . . . . . 155
lschium . . . . . . . . . . . . . . . . . 156
Skull and jaws . . . . . . . . . . . . . . . . . 156
Vertebral column . . . . . . . . . . . . . . . . 158
Other skeletal elements . . . . . . . . . . . . . . . 159
Gastralia . . . . . . . . . . . . . . . . . 159
Scutes . . . . . . . . . . . . . . . . . 159
Summary of pseudosuehian evidence . . . . . . . . . . . 159
The Sphenosuchus evidence . . . . . . . . . . . . . . . 159
 ARCHAEOPTERYX A N D THE ORIGIN OF HlKDS 93

Kvidencc supposedly contrary t o a theropocl ancestry t o r Arcliueopteryx . . . . . 163

After Ileilmann . . . . . . . . . . . . . . . . . . . . . 168
Summary . . . . . . . . . . . . . . . . . . . . . . . 173
Acknowledgements . . . . . . . . . . . . . . . . . . . . 174
Hcferrnccs . . . . . . . . . . . . . . . . . . . . . . 174
Addendum and Reference . . . . . . . . . . . . . . . . . . 179
Appendix: Systematic list of taxa . . . . . . . . . . . . . . . . 179

INTRODUCTION

For nearly half a century, a general consensus has placed the origin of birds
among pseudosuchian thecodontians, a group of primitive archosaurian reptiles
of Triassic age that also are believed to have given rise to the two orders of
dinosaurs, flying reptiles (pterosaurs), and crocodiles as well. The
pseudosuchian ancestral theory, first explicitly suggested by Broom (1 9 1 3 ) ,
achieved general acceptance with the publication of Heilmann’s classic
monograph on Tlic origin oj’ birds in 1926. Since that time, few alternative
theories on bird origins have been proposed and today the pseudosuchian origin
enjoys almost universal acceptance.
A pseudosuchian ancestry of birds, and of higher archosaurs, quite probably
is ultimately correct, considering that pseudosuchians are among the oldest and
most primitive archosaurs known. But it is possible now t o identify a more
immediate ancestry of birds more precisely-one that is post-Triassic and
post-pseudosuchian. In the half century since Heilmann assessed the various
reptilian groups that might have given rise to birds, critical new evidence has
come to light, the most important of which is the discovery or recognition of
three more specimens of Arcliucwptcrj3.u. Ornithologists have long recognized
that various anatomical features of modern birds suggest that they arose from
reptilian stock, but the most persuasive evidence of all rests in the five
presently known specimens of that archaic bird.
Possibly no other zoological specimens, fossil or Recent, are considered so
important as are those of Arcliacopteryx litliographica (see Figs 1, 2 and 3).
Certainly few other specimens have generated such widespread interest or
provoked as much speculation and controversy. The reasons are several: these
specimens are the oldest (Tithonian = Late Jurassic) known fossil bird remains;
they are extremely rare, only five specimens (excluding the solitary feather) are
known at present; several of these preserve remarkably detailed impressions of
feathers and an extraordinary mixture of reptilian and avian characters; and
most important of all, because of the last fact, out of all presently known fossil
and living organisms, these specimens are widely recognized as constituting the
best example of an organism perfectly intermediate between two higher
taxonomic categories-representing an ideal transitional stage between ancestral
and descendant stocks. Archaeopteryx may well be the most impressive fossil
evidence of the fact of organic evolution.
The objective of this paper is to review and evaluate all available fossil
evidence pertaining to the immediate, rather than the remote Triassic, ancestry
of Archaeopteryx and to offer an up-dated theory of the origin of birds. The
data, interpretations and conclusions that follow are founded on the single
critical assumption that Archaeopterjsx holds the key to bird origins, wlietlirr
or not it occupied a positiorz directly ancestral to later birds. After extensive
study of all five skeletal specimens, it is my firm conviction that
Figure 1. The first two skeletal specimens of Archaeopteryx lithographica to be recognized. A. The type specimen, now in the British Museum
(Natural History). London; B. the Berlin specimen, once designated Archueomis siemensi. now in the Humboldt Museum fur Naturkunde. East
Berlin. Scale = 10 cm.
ARCffAEOPTERY X A N D THI< OKIGIN OF RlKDS 95
96 J. 11. OSTKOM
 ARCffAEOPTER Y X A N D ‘ l l i l < ORIGIN 0 1 ; HIKDS 97

pseudosuchians were remote, not only temporally but phylogenetically as well,

from the origin of Arc,liuc’ol)te,:1,s. ‘I‘he evidence in these five specimens points
unequivocally to an immediate ancestor among the small coelurosaurian
theropod dinosaurs (Ostrom, 1973, 1975a,b).

HIS’TOKICAI. K K V lliW

Early in the last century, some scholars believed that birds had existed as far
back as ‘Triassic times. That belief stemmed from the abundant occurrence of
bird-like footprints in the Late ’I’riassic strata of the Connecticut Valley in
North America. These are now believed to be dinosaur footprints, but they
were not recognized as such until after the discovery of the first specimen of
Arc.liuropterj-.uin 1861. That discovery apparently led to a revised conclusion
that birds had not originated until sometime after the close of the Triassic.
Despite the curious plume-like structures in the Early Triassic, presumed
thecodontian L o r i g i ~ q z ~ u ~(Sharov,
~iu 1970), n o contrary evidence has turned up
as yet. That does not, however, rule o u t the possibility that ’I’riassic
“feathered” vertebrates existed; feathers obviously existed before
A rcli uco1)t erjas.
A brief reference by von Schlotheim (1820) t o feathered fossils from the
limestone beds near the towns of Pappenheim and Solnhofen in Bavaria is the
first published record of the possible existence of birds during Jurassic times,
although, in 1820, little of the geologic column had been deciphered and even
less was known about geologic time. Unfortunately, the whereabouts of von
Schlotheim’s feathered fossils are unknown today. The first still-verifiable
evidence of Jurassic birds is t h e imprint of a solitary feather in a small slab of
these same Solnhofen limestones (Fig. 2A). This find was reported by von
Meyer (1861a) in a letter to Professor H. Bronn, published in Bronn’s Nerres
Julirbircli fiir Mirierulogie (p. 561). Less than two months later, von Meyer
(1861b) reported the discovery in the same limestone strata of a partial
skeleton associated with distinct impressions of feathers. This find, the now
well-known London specimen (Fig. l A ) , is currently in the British Museum
(Natural History) in London. At first, some scholars questioned the authen-
ticity of both specimens, b u t von Meyer (1 862) established them as genuine.
Early debate centered on the question of the proper systematic assignment
of the skeletal remains. Were they the remains of a true Jurassic bird or merely
those of a feathered reptile? Wagner (1861), who accepted the specimens as
authentic fossils even though he never saw them, finding it impossible to
conceive of a “reptilian bird” declared the remains t o be those of a “feathered
reptile”, which he proceeded to name Gripliosaitrirs problematicus. Most
scholars, however, quickly accepted the opinions of Owen (1862, 1863) and
Huxley (1868a) that Arcliaeoptc~ryx was indeed a true bird, albeit very
primitive, of great antiquity.
While the avian versus reptilian controversy derived chiefly from the mixture
of avian and reptilian characters preserved in the London specimen, another
major contributing factor was the particular time of that discovery-1861-
98 J. H. O S f H O M

Barely two years after publication of Darwin’s Tlie origiii of specics (1859). In
that light, some of Wagner’s comments are of special interest:
“In conclusion, I must add a few words to ward off Darwinian
misinterpretations of our new Saurian. At first glance at the
Cripliosurtnis we might certainly form the notion that we had before
us an intermediate creature, engaged in the transition from Saurian
to the bird. Darwin and his adherents will probably employ the new
discovery as an exceedingly welcome occurrence for the justification
of their strange views upon the transformations of animals. But in
this they will be wrong.” (Translated from Wagner, 1861: 146)
Despite the protests of Wagner and of other anti-evolutionists (even recently ;
see Armstrong, 1966, and Armstrong & Kroll, 1967 for two recent exorcisms),
Archaeopfer?ix has long been recognized (Huxley, 1868a) as the most
persuasive-if not compelling-evidence for a reptilian ancestry of birds.
By the time the second skeleton of Archucwpfwj~x,the now famous Berlin
specimen (Fig. lB), was discovered in 1877, the debate had shifted (as Wagner
expected it would) from the question of the proper systematic position, to that
of the origin of birds and the particular reptilian affinities of Arcliueopferys.
Over the years, Archueoptrryx has been linked with a variety of reptiles
including lizards, pterosaurs, ornithopod dinosaurs, theropod dinosaurs and
pseudosuchian thecodonts*. Most recently, Walker (1972) has suggested an
affinity with primitive, Triassic crocodilomorphs. At first, dinosaurian affinities
were favoured, owing largely to the works of Cope (1867), Huxley (1867,
1868b, 1870), Marsh (1877, 1881b), Gegenbaur (1878), Williston (1879), Vogt
(1879, 1880), Baur (1883, 1884a,b, 1885a,b) and Abel (1912). Opposition to
the dinosaurian theory was expressed by Seeley (1881), Dollo (1882, 1883),
Dames (1884, 1885) and Parker (1887). Owen never published his views on this
question, but apparently he, too, opposed dinosaurian affinities.
Furbringer (1888) was the first to suggest what might be called a
compromise theory which postulated an unspecified common ancestor for
birds and dinosaurs. The common ancestor hypothesis was advocated in later
years by Osborn (1900), Broom (1906, 1913), Heilman (1926)’ Tucker
(1938a,b) and, in modified form, by Galton (1970). I t is the preferred theory
today, although a few contrary schemes have been presented by Boas (1930),
Lowe (1935, 1944) and Holmgren (1955). Heilmann’s superb study seems t o
have stilled the debate, for nearly all recent authors have accepted bird origins
among Triassic pseudosuchian thecodontians (de Beer, 1954a,b, 1964; Bock,
1969a; Swinton, 1958, 1960, 1964; Piveteau, 1950, 1955; Welty, 1962;
Romer, 1966, 1968; George & Berger, 1966; Van Tyne & Berger, 1959;
Pettingill, 1970).
Today’s high cost of publication prohibits a detailed review of the rise and
fall of the various theories on the relationships of Arclzaeopteryx and the origin
of birds (readers are referred to the bibliography at the end of this paper), but a
brief summary is in order. As noted above, prior to the pseudosuchian theory,
dinosaurian affinities were accepted by many. But the fragmentary fossil record

* See Appendix for a summary classification of the taxa referred to herein.

 ARCIfAEOPTER Y X A N D T l I I i ORIGIN O F RIRIIS 99

and the less complete roster of then known dinosaurs* prompted Mudgc
(1879) to observe that:
“The dinosaurs vary so much from each other that it is difficult to
give a single trait that runs through the whole. But no single genus or
set of genera, have many features in common with birds, or a single
persistent, typical element or structure which is found in both.”
(Mudge, 1879: 226)
That was followed by Furbringer’s (1888) conclusion that direct descent of
birds from any known type of dinosaur was not possible and all resemblances
between dinosaurs and birds were “parallels” and “convergent analogies”.
Broom (1906) argued that birds had arisen “from a group immediately
ancestral to the Theropodous Dinosaurs” and in 191 3 he specified:
“The Pseudosuchia, now that it is better known, proves to be just
such a group as is required. In those points where we find the
Dinosaur too specialized we see the Pseudosuchian still primitive
enough.” (Broom, 191 3 : 63 1 )
Thus, the stage was set for Heilmann (1926). After comparing the skeletal
anatomy of Arc/zaeoprery.u with that of various ancestral candidates, namely
pterosaurs, ornithopods, coelurosaurian theropods and pseudosuchians, he
found the closest resemblance to be with coelurosaurian dinosaurs. Yet, he
rejected a coelurosaurian ancestry of birds solely because clavicles, the
precursors of the avian furcula, were unknown in theropods. Following the
suggestion by Broom, Heilmann, too, concluded that birds probably arose from
pseudosu ch ian s.
Not since Lowe (1935, 1944) has anyone denied the avian identification of
Archaeopteryx, or its importance for avian origins. That is not to say, however,
that everyone accepts it as the ancestral bird. Lowe (1935), Tucker (1938b),
Swinton (1960) and George & Berger (1966), to cite just a few, considered
Ardtaeopteryx t o be an aberrant form, well removed from the main line of bird
evolution. The extreme view was that of Lowe (1935) who believed
Arc/iaeopter~~.r to be a feathered dinosaur (!) while Swinton (1960) argued
“there is no justification for making Archuc~opterq,x the progenitor of all
subsequent birds”, pointing out that it would be extremely improbable if the
most ancient bird known to us also happened to represent any stage of the
main avian lineage. Simpson (1946) and de Beer (1964) on the other hand,
concluded that Archaeopteryx probably was on the direct line of evolution
from reptilian ancestors t o modern birds. Although I accept Swinton’s logic
(but not his conclusion), it must be pointed o u t that a “main line” position for
Archaeopteryx is not impossible. Not one feature of the skeletons or of the
plumage impressions of any of the known specimens precludes such a central
ancestral position. No other contemporaneous or more ancient candidates are
known (except possibly the indeterminate specimen of Laopteryx; see Marsh,

* The roster of dinosaurian genera has more than trebled since 1879, according to White’s ( 1 9 7 3 )
Cutalogue of the Generu of Dinosaurs. It should also be pointed out that the meaning of the term
“dinosaur” has changed since that time. Prior to the turn of the century-, although applied to many of the
same taxonomic groups as in today’s usage, the term seems to have been visualized as encompassing a
much narrower and more closely related spectrum of taxa than is generally accepted today.
I 0 0 J. 11. OS'I'KOM

1881 and Simpson, 1926). lmprobable though it may be, the possibility that
Archueoptcv-ji.x actually was ancestral to all later birds still exists, and the
critical question remains: What was the source of Arcliuewpfc~rj~s?

lVII>I:NCI< O N '1'1111 ORIGIN O F AHCHAEOPTER Y X

At the present time, three distinct reptilian groups may be considered as

poss ib 1y an ce st r al to A rdt ucwp f crj '.r: The Or n it h o p od a (Or d e r Or n i t hi sch i a * ,
Theropoda (Order Saurischiat) and Pseudosuchia (Order Thecodontia? ). A
fourth candidate, sphenosuchid crocodilomorphs, has been suggested by Walker
(1972) as close to the origin of both birds and crocodiles. There is n o evidence
indicative of close phyletic relationships between Arc/iucoptrrjv.x and either
pterosaurs or lim-ds, hence neither of these groups will be considered further
here. The only relevant evidence available to us for deciding which (if any) of
the above suborders is the true and immediate ancestral group consists of the
skeletal anatomy preserved in all of the known representatives of these four
groups and that of the five presently known specimens of Arc/iac.o/if~'r!,.r.The
following material is organized o n that basis. The anatomy of modern birds, so
highly specialized and so remote in time from the Class origins, is of n o value in
seeking the origins of Arcltueoprerj's, and is also excluded from further
consideration here.
Precisely what constitutes valid evidence of close phyletic relationship
between t w o or more taxa has been, and still is, the subject of intense debate
(see Bock, 1969b,c,d; Brundin, 1968; Colless, 1967, 1969a,b; Cracraft, 1967;
Ghiselin, 1969; Hennig, 1966; Hull, 1967; M a s h , 1952; Nelson, 1970; and
many others). Nevertheless, structural similarity, whether it be at the genetic,
molecular o r anatomical level, is widely accepted as t h e most reliable index of
phylogenetic affinity. The difficulty is not in recognizing the degree of
resemblance, b u t in distinguishing between those resemblances that are
homologous and those that are not. Unanimity is rarely achieved because of the
difficulty o r impossibility of proving to everyone's satisfaction either the
homology o r non-homology of similar features. Such "proof" requires evidence
that is rarely, if ever, available-that is, full and complete knowledge of the
entire phylogenetic series. In the absence of that kind of documentary
evidence, the only reasonable working hypothesis remaining is that such
resemblances are homologous in the absence of contrary evidence, and the more
extensive and detailed t h e structural similarities, the closer the phylogenetic
relationships. In Hennigian terms, the greater the frequency of derived
characters in common, as opposed to primitive characters, the closer the
relationship.

* Order Ornithischia
Suborder Ornithopoda: Families; Fabrosauridae, Heterodontosauridae, Hypsilophodontidae.
Iguanodontidae, tlatlrosauriclae.
t Order Saurischia
Suborder 'rhcropoda (Infraorder Coelurosauria): Families; Proconipsognathidae, Coeluridae,
Segisauridae. 1)roniaeoFauri~lae; Ornithomimidae, (Infraordcr Carnosauria): Megalosauridae.
Tyrannosauridae.
$ Order Thecodontia
Sulmrder Pscudosuchia: Families; Euparkcriidae, Ornithosuchidae, Prestosuchiilae, Sclero-
rnochlitlac.
 AKCIIAt‘OPTER Y.Y A N D T t I K ORIGIN 0 1 : HlliDS 101

If ornithopods, theropods and Arcliucwpterj3s include a pseudosuchian stage

somewhere in each of their ancestries (which few authorities would challenge),
thc matter before us may be reduced to t w o possible alternatives. First,
Arc1iueoptcyv.y evolved directly from a pseudosuchian ancestor independently
of the contemporaneous ornithopod and theropod lineages. Or, second,
Arc.liacioptcr?,.r evolved from a pseudosuchian ancestry h!. I V L L ~ ’ o f an
intermediate theropod o r ornithopod stock. The second alternative concludes
that the similar features of AI.(,liacoptcrj,s and theropods (or ornithopods) are
homologues. The first alternativc requires that such similarities be
non-homologous and independently derived in parallel.
Parallel evolution may be defined as the similar response (adaptive change)
of a common heritage in two o r more related lineages to similar environmental
conditions (selective pressures). Visualize, if you will, two “sibling” lineages
diverging from a common ancestor. They possess certain shared primitive
characters of their common ancestor, plus the latent, but as yet unexploited,
potential to develop similar specialized adaptations (derived characters) as a
result of experiencing the same or very similar environmental conditions and
ecologic opportunities-in more or less the same sequence. The essential
criterion of parallelism is that derived characters in common among related
(sibling) descendant groups are riot present in the common ancestor. In other
words, the postulated relationship between “ancestor” and “descendant”
lineages (species) is based entirely upon the occurrence of /irir~iitiwc1iaructer.s
in common, whereas the common occurrence of clcrivctl c.liaructcrs is taken to
mean “sibling” relationship o t i l i , . ‘lo express this yet another way: parallelism
is a purely theoretical explanation to account for the absciicv in any known
antecedent of certain derived characters that arc present in the supposed
parallel groups. While I accept t h e concept of parallel evolution, in my opinion,
the easily explained gaps in the known fossil record d o not validate the negative
evidence upon which the concept of parallelism seems to rest.
I t is conceivable that solitary specialized (derived) features, or even several
component features of a single structural complex, may arise more than once in
parallel. But the probability of multiple near-identical structures of several
independent structural complexes evolving in parallel seems very remote
indeed. I t is quite illogical to me to dismiss a (phylo)-genetic explanation of
multiple derived characters in common in favour of coincidental environmen-
tally imposed likeness. The critical question before us is: Which of the three
possible ancestral groups possesses the highest incidence of
“Arcliacopter~.u-like”derived characters?

The orr I itli opod e viden ce

The only advocate of an ornithopod ancestry of Arcliaeopteryx was Baur
(1883, 1884a), although Galton (1970) suggested a common ancestor for
Archaeopteryx and ornithischians. Baur based his conclusions chiefly o n the
evidence of the tarsus and pelvis in various dinosaurs which he contended
approached the condition found in modern birds. However, most of the taxa
cited by Bau r ( Am pli isaimis, Zar I clodon, Cornpsogna t h iis, Cera tosaiirirs ) are
now known to be saurischian rather than ornithischian! There was a tendency
in some ornithopods (Tliescelosairrus. Laosaunrs, Camptosaurus, Hypsilo-
102 1. tl. OS'I'HOM

C 5"

Figure 4. (:omparison ot the left toot and metatarsus in Arcliaeoprerys ( C ) with that o f t w o
ornithopod ornithischians, Luosuurus consors, Y .P.M. 1882 (B) and Tenonrosaurus rilleti*, P.U.
16338 ( R ) . For added comparison, the feet o f two coelurosaurian theropods are included;
Coeloplrysis longicolis. A.M. N. t 1. 7224 ( A ) and Compsogna thus lorrgipes. Bayerische
Staatssammlung, Munich (D). All specimens are reproduced to unit length of thc metatarsus for
easier comparison. Notice the more masive construction of the ornithopod feet, regardless of
size. as compared with those of Archaeopteryx and the theropods. Notice also that the hallux
(digit I ) is not reversed in the ornithopod foot as it is in Archaeopleryx and theropods. Vertical
scales = 3 em. See also Fig. 15. A.M.N.H., American Museum; P.U., Princeton University;
Y.P.M., Yale Peabody Museum.
 ARCHAEOPTERYX A N D T t l l i OKIGIN O F HlKDS 103

pliodon, Tc~)iont;sairrirs") for reduction of the first toe, but the hallux was
never reversed to a position behind digit I1 as occurs in Arc~/iuc~optrr~~.u,most
modern birds and in nearly all theropods as well. As Fig. 4 shows, with the
exception of the absence of digit V, there is little similarity between the pes of
Arcliacwptcryx and those of typical ornithopods. The latter tend to be broader
and more massive and while the tarsus does feature a mesotarsal ankle joint, the
astragalus lacks an anterior ascending process.
The superficial resemblance of the ornithischian pelvis to that of modern
birds (Fig. 5 ) , and also perhaps to that of Arcliuroptrrj-.u (but see later
comments on this) has been the most important factor behind suggestions of
close evolutionary relationships between birds and ornithischian dinosaurs.
That similarity led Galton (1970), and others earlier, to equate t h e
ornithischian postpubic rod (posterior ramus) with the posteriorly directed
pubis of birds. The ornithischian prepubic process (absent in all Triassic and
Early Jurassic ornithischians; see Fig. 5 ) Galton equated with the pectineal
process of modern birds, which is developed o n the ilium, but which is absent

iI

Figure 5 . Pelves. in left lateral view. o f several ornithischians compared with t h e pelvic
organization of a modern h r d ( A ) a n d that ah presen>ed in the Berlin A r ( , / i u e o p r e r w (I-').T h e
post-puhic rod (PO) of ornithischians has I x e n equated with t h e avian pubis ( p u ) . T h e pre-puliic
process ( p r ) is considered a new structure. A. Colionbu; H. Scelidosuurrrs; C , C'uwiprosuirnts; D .
Tliescelosuirnts; E, Sregosuitnrs; I:. Arcliucopteryx (Berlin \peciiiien). For I he restored puhic
orientation postulated for Arcliuropturyx. see Fig. XC. Sketches a r e not t o wale. i l l Ilium: isc,
ischium: po, po\t-pul)ic rainus: pr, prepubic ramus; pu, pubis.

or only very weakly developed in Archaeopteryx. Galton related the backward

shift of the ornithischian and avian pubis to the development of bipedal
locomotion and suggested that this backward shift may have occurred only
once. For this reason, he postulated a common ancestor for birds and
* Tenontosuunrs is an iguanodontid ornithopod recently described (Ostrom, 1970a) f r o m t h e Early
Cretaceous of western North America.
104 J . 14. OSTIIOM

ornithischians, clearly acknowledging that birds could not be descendant from

any presently known ornithischian. Galton also recognized that the very
different pelvic arrangement in bipedal theropods posed some difficulties for
his theory of bipedal locomotion in the origin of the avian and ornithischian
pelvic arrangement, but he offered n o further explanation. For additional
discussion of the ornithischian (and archosaurian) pelvis, see the excellent
paper by Charig (1972).
I t now appears that the evidence of the pelvis is not so important after all.
Despite the similarity in the orientation of the pubis in modern birds and
ornithischian dinosaurs, there is substantial evidence, unrecognized until
recently, that the pubis of Arcliuroptcr.vs almost certainly was not directed as
sharply backward as the Berlin specimen seems to indicate. This, coupled with
the absence of any other bird-like, or, more appropriately, Arc,/zuc.opferl,.\-like,
features in any known member of the Ornithischia (a fact which Heilmann
stressed) greatly diminishes the probability of close phyletic relationship
between ornithischians and birds. Galton attempted to explain the absence of
any other avian features in ornithischians as the result of subsequent
specializations and a shift to herbivory.
The Berlin specimen of Archacwpteryx seems to show the pelvic bones in
natural articulation, with the pubis extending down and backward nearly
parallel to the ischium, very similar to the condition in modern birds. The first
indication that this "bird-like" arrangement might not be correct was suggested
by the Teyler or Haarlem specimen, recognized as Arclrac~opteri*sin 1970
(Ostrom, 1970b, 1972). As Fig. 6 shows, parts of the shaft and the distal
extremities of the pubes in that specimen are preserved between the shafts of
the femora in what appears to be natural position. However, there is n o
indication whatever of the ischium adjacent t o these pubes as there should be if
both elements were originally positioned as they are prcwriwl in the Berlin
specimen. Since all other bones of the Teyler specimen are preserved in
articulation, it seems unlikely that the ischia only were disarticulated. Also
important in this specimen is the orientation of the pubis. The pubic shaft and
extremities form an axis that is nearly perpendicular to that of the posterior
dorsal vertebrae (Fig. 6), in contrast t o the 130" to 140" angulation preserved
in the Berlin specimen (Fig. 7A). It is quite possible, of course, that either the
pubes or the ischia of the Teyler specimen were displaced from their natural
positions, but there is n o evidence of disarticulation. More importantly though,
other independent evidence exists which supports the non-avian pubic
orientation in A rclr aeop tcrys.
This evidence is to be found in the Berlin specimen, the Maxberg specimen,
and in the recently described fifth specimen of Arclzaeopteryx (Mayr, 1973;
Wellnhofer, 1974). Close examination of the Berlin specimen reveals a number
of details that establish beyond any possible doubt that the t w o sides of the
pelvis are displaced in relation to each other and that the right pubis is r r o f
preserved in a natural position. First, there is a distinct fracture across the
proximal part of the right pubis (Fig. 7A). Secondly, as noted by Heilmann,
there is a triangular area adjacent to that fracture that is not identifiable as
bone, consisting of fine calcite crystals. Thirdly, the shaft and distal expansion
of the right pubis are preserved a t a higher level in the slab than the right
ischium, and seem to be twisted with respect to the parasagittal plane defined
 ARC'IIAEOPTERYX A N D 'llll- ORIGIN O F BIRDS 105

Figure 6 . T h e main slab of the Teylcr speclmen of Arcluwoptervx to show thc angular
relationship (approx. 9 0 " ) o f the putis t o t h e posterior dorsal vertebrae. Notice that no sign of
the ischium is preserved in the expected position. Compare with Figs 7 and X.
Figure 7
 ARCHAEOPTERYX AND THK ORIGIN O F BIRDS 107

by the position of the right ilium. As shown by the London specimen, where
both pubes are still joined, although disarticulated from the rest of the pelvis,
there is a long pubic symphysis that would seem to require that the pubis in its
natural position be situated medial t o the ischium. Consequently, unless
displaced, it should have been preserved at a lower level in the slab than is the
case in the Berlin specimen. Finally, X-rays show that the left half of the pelvis
in the Berlin specimen has been displaced upwards and backwards relative t o
the right, as is indicated by the position of the left femur and the dorsal edge of
the left ilium (see Fig. 7A). Preparation of the underside of the Berlin specimen
(Fig. 7B) has revealed that the distal end of the left pubis is still fused with that
of the right pubis, but the two pubic shafts diverge upward to opposite sides of
the right acetabulum. The X-rays (Fig. 7C) also show these discordant traces of
the two shafts. Because of the firm union at the pubic symphysis, the upward
and backward displacement of the left half of the pelvis appears to have pulled
the right pubis (but not the ischium or ilium) up and backward also, fracturing
it just below the ilium.
In the new Eichstatt specimen, the pubes are both present (Fig. 8A), but
appear to be slightly rotated about a vertical axis. They are oriented at about
100" to 110" to the trace of the posterior dorsal vertebrae. This orientation is
very close to that seen in the Maxberg specimen (the third specimen of
Arcliueopteryx), where, as shown in Heller's X-ray (1959, pl. 14-l), but not
previously noted, the angle between the long axis of the ilium and the pubic
shafts is also about 100" (see Fig. 8B). Thus, the last three specimens t o be
found all seem to confirm a displaced position of the pubis in the Berlin
specimen.
It is not surprising that this condition was not recognized before, because
obviously there is nothing surprising about a bird-like orientation of the pubis
in an ancestral bird. In fact, it is improbable that post-mortem displacement
coincidentally would have aligned this element in a bird-like orientation. The
evidence is clear, however, that the pubis in the Berlin specimen has indeed
been displaced, even though the natural position of the pubis cannot be
reconstructed precisely. The fact that it has been displaced, and the absence of
any positive evidence in any of the other specimens of Archaeopteryx that the

t:igure 7. The pelvic region o f the Berlin Archaeopteryx specimen showing the different
pobitions of the left and right pelves, evidence that the "hird-like" position of the right pubis in
this specimen is not necessarily that o f the original orientation in life. A. The upper surface of
the main slab; arrow 1: the distal expansion of the pubis; arrow 2 . the posterior extremity of
the ilium; arrow 3, the dorsal margin of the left ilium clearly displaced upward relative t o the
right ilium; arrow 4, a fracture between the right pubis and ilium; arrow B, the shaft of the right
pubis. R. Underside of the same specimen (printed in reverse for easier comparison with A ) ;
arrow 1 points to the distal expansion of the / e f t pubis still fused t o the right pubis (arrow 1 of
A ) ; pointer A indicates the / e f t pubic shaft, and pointer C points t o the head of the right femur
still articulated in the acetahulum (as can he seen in A). The reversed printing of B clearly
shows that the shafts of the left pubis (arrow A) and the right pubis (arrow B) diverge upward
to opposite sides of the acetabulum. C. X-ray image of the same region (to approximately the
same scale as A and B ) showing the dissimilar positions of the two pubes. Arrows 1 and 2
respectively indicate the distal expansion of the puhes and the posterior extremity of the right
ilium. Arrows A and B point t o the shafts of the left and right pubes, respectively. Compare
these photos with Figs 6 and 8. See text for further explanation. Scale divisions in A = 1.0 mm;
in B = 0.5 mm. C is at the same magnification as A. X-ray provided through the courtesy of Dr
H. Jaeger, Hurnboldt Museum fur Naturkunde, East Berlin.
8
108 J . H. OSTKOM

m
 ARCIfAAEOPTERYX A N D THE ORIGIN OF BIRDS 109

pubis had a bird-like (or ornithischian-like) orientation, nullifies the most

important evidence of previous theories that related Arcliaeoptc~rj~xwith
ornithopod dinosaurs.

TI1e t h c w pod e viden cc

It has been stated (Colbert, 1969) that if it were not for the impressions
of feathers, it is unlikely that the London and Berlin specimens of
Arcliaeoptc~ryx would have been identified as “bird” remains, but instead
would have been labeled reptilian. (Recall the early debate about feathered
reptile versus reptilian bird.) 1 would go even further than that. Were it not for
those remarkable feather imprints, today both specimens would be identified
unquestionably as coelurosaurian theropods. Notice that with the exception of
the misleading orientation of the pubis in the Berlin specimen, there is only o n e
skeletal feature that is iiof currently known in any theropod specimen. This
single feature is the firsion of the clavicles into a furcula. More will be said
about this later.
Some of the coelurosaurian characters of Arcliacwpteryx, particularly of the
hand, have been discussed by several previous workers (see Lowe, 1935;
Tucker, 1938b; Holmgren, 1955), b u t not since Heilmann (1926) has there
been a comprehensive review of all the evidence, The fact is that there is much
more evidence for the theropod affinities of Archaeopteryx than has generally
been recognized. This evidence has been augmented by the three recently
recognized specimens of Arclzaeopteryx, but in past years it has been largely
overlooked because of frequently invoked suppositions of convergence and
parallelism. Another critical factor has been the discovery of a variety of new
theropods since Heilmann’s time. All these discoveries make it necessary to
re-examine the question of Arctiaeopteryx and bird origins. The following data
and interpretations are based o n my own extensive studies of the five
specimens of Archaeopteryx and of nearly all the theropod taxa cited herein.
Manus urid forelimb
Although sometimes described as “bird-like”, the hand and forelimb of
Arclzueopteryx actually are n o t like those of modern birds a t all, b u t they are
remarkably similar in a number of details to those of certain small theropods,
namely O m itholestes, Deinoriy ch us, Velocirap tor, CIi irosteriotes and probably
St enony clz osau rus and Sau ro rn it li oides. Some o f these s i m i lar i t ies h ave
repeatedly been explained as adaptive only and of n o phylogenetic

Figure 8. A. The pelvic region of the I:ichstatt specimen of Archaeopteryx. showing the
preserved pubis orientation nearly perpendicular to the long axis of the ilium. very close to the
orientation preserved in the Teyler specimen (see Fig. 6); arrow 1, distal expansion of the
pubes; arrow 2, posterior extremity of the left ilium; arrows A and B, left and right pubes lying
on top of the left femur. B. X-ray image of the pelvic region of the Maxberg Archaeopreryx
(from pl. 14-1, Heller, 1 9 5 9 ) ; 1 indicates the distal expansion of the pubes; arrow 2 points to
the posterior process of the ilium; arrow A points to the parallel shafts of the pubes, oriented
nearly perpendicular t o the long axis of the ilium, as in the Eichstatt specimen (A) and the
Teyler specimen (Fig. 6). C. My best estimate of the reconstructed natural position of the pubis
in Archaeopteryx, based o n analysis of all five skeletal specimens. D. Past traditional
interpretation of the pelvis in Archaeopteryx. based on the right side of the Berlin specimen.
Scale divisions in A = 1 .O mm.
 J. El. OSTROM

Figure 9. Comparison of the (right) hand and metacarpus of the Berlin Archaeopteryx specimen
( A ) with those (lefts) of the theropods Deinonychus antirrhopus ( B & C). Y.P.M.5206, and
Ornitholestes lrermanni (D), after Osborn (1917). Notice the relative lengths of the three fingers
and metacarpals in all three. Similar construction of the hand and metacarpus is also found in
Velociruptor motigoliensis, as is illustrated in Fig. 10C. Compare these data with Fig. 24. Scale
divisions in A = 0.5 m m ; scale lines in B, C. D = 5 cm.
 ARCHAEOPTERYX AND THE ORIGIN OF BIRDS 111

signficance-in other words, due to convergent or parallel evolution (Tucker,

1938b; Simpson, 1946; d e Beer, 1954b). Elsewhere (Ostrom, 1974a), I have
challenged that explanation and more will be said about this later.
Muni4.s. The manus of Archaeopteryx (Fig. 9A) consists of three fingers,
digits IV and V having been lost. Digit I is the shortest, 11 the longest and 111 is
intermediate in length. Digits I and I1 are the more robust and digit I l l is
slender and more delicate. The phalangeal formula is primitive (2-3-4), but the
phalangeal proportions are not. The penultimate phalanx is the longest in each
finger, rather than the proximal phalanx (the primitive condition), and that of
the first finger is especially long as compared with the terminal phalanx.
This same configuration occurs in several small to moderate sized theropods
(Figs 9C, D and lOC), such as Ornitholestes hermanni (Osborn, 1903, 1917))
Deinonychris antirrlzopw (Ostrom, 1969), Velociraptor mongoliensis (see pl. 2,
fig. 2 of Kielan-Jaworowska & Barsbold, 1972) and Chirostenotes pergradis
(Gilmore, 1924a). The chief difference between these theropod hands and
those of Archaeopter.vx is one of size, all of the theropods being larger. Also,
the fingers are relatively shorter in the theropods.
The phalangeal formulae clearly indicate that the fingers retained in both
Arclzaeopteryx and the theropods mentioned above, as well as in many other
theropods, are the first three. Any other interpretation would require that
digits I and V were lost completely in all these taxa and the remaining fingers
reduced by one and only one phalanx each, or that digits I and I1 were lost
without trace and the third and fourth fingers onlv were reduced by two
phalanges each. Retention of a splinter-like metacarpal remnant at the fourth
position in Ornitholestes and a reduced finger at that position in Coelophysis
rule out the last explanation, for those taxa at least. Since there is no clear
evidence of reduction of the first finger in any archosaur, the conservative
interpretation is that these fingers represent I, I1 and 111. The same is true of
Archaeopteryx, and probably therefore, of modern birds as well (contra,
Montagna, 1945; Holmgren, 1955, who interpreted the modern bird digits as 11,
111 and IV, on the basis of embryological evidence).
Metacarpus. The metacarpus of Archaeopteryx consists of three metacarpals
(Fig. 9A) with no sign of any other elements being preserved in any of the
presently known specimens. Metacarpal I is very short and robust, while the
second and third are very long (more than three times the length of the first)
and subequal in length. The first two metacarpals are tightly appressed
proximally (Fig. lOA), but are slightly divergent distally. Metacarpal I11 is more
slender than the others and not so closely appressed against metacarpal 11. The
first and second metacarpals may have been fused since they are preserved in
contact in one or both hands in the Berlin, Eichstatt and Maxberg specimens,
and perhaps in the Teyler specimen too. In the London specimen, however, the
left metacarpals I1 and 111 are preserved together, but metacarpal I is missing,
suggesting that it was not co-ossified with metacarpal I1 in this, one of the
largest of the five specimens. Heilmann (1926) thought that the second and
third metacarpals might have been fused proximally, but the Maxberg specimen
(the next largest) clearly shows that they were separate. The left metacarpal 111
is lacking in that specimen and that of the right side is separated by more than
2 mm from metacarpal 11. The larger dimensions of the Maxberg and London
specimens indicate that they were not immature, as compared with the smaller
112 J. H. OSTROM

Figure 10
 ARCHAEOPTERYX AND THE ORIGIN O F BIRDS 113

Berlin specimen which may be, so we must conclude that the three metacarpals
were not normally co-ossified in the adult Archacwpteryx.
Several theropods display this same basic construction of the metacarpus,
most notably, Ornitholestes, Deinonj~cliusand Velociraptor. Chirostenotes may
also have had a similar arrangement, although metacarpal 111 is not known. In
each instance, the proportions are approximately the same as in Archeopteryx
and metacarpal 111 is more slender than the others. Most significant, however, is
the form and closely appressed placement of the first metacarpal in all. In
particular, notice the distinct proximal external facet for contact with
metacarpal I1 and the internal basal expansion on metacarpal I of Deinorzgchus
(Fig. 10B) and apparently also in Vdociraptor (Fig. 1OC). These compare very
closely with similar features in the first metacarpal of Archaeopteryx.
Carpus. The carpus of Arc/zaeopter.vx appears to be composed of just three
elements, one large semi-lunate distal carpal and two small ossicles. Petronievics
(1923) first distinguished two carpals which he identified as the radiale and
ulnare. Later (1925) he suggested that the “radiale” was composed of two
fused distal carpals. Heilmann (1926) thought he could distinguish four
separate carpals in the left wrist of the Berlin specimen, but this is far from
evident in that specimen, nor does the new Eichstatt specimen substantiate this
suggestion. The large semi-lunate distal carpal (the radiale of Dames, 1884, and
Petronievics, 1923) articulates very precisely with metacarpals I and 11, but has
no contact with metacarpal 111. This condition is very clearly shown in the right
carpus of the Eichstatt specimen (Fig. 10A) and the left wrist of the Berlin
specimen. A smaller carpal is preserved at the proximal end of the third
metacarpal in the Eichstatt specimen, but whether it represents a proximal or a
distal carpal cannot be established. Heilmann (1926) interpreted this as a
centrale in the Berlin specimen, but I suspect that it is the ulnare. Another
small element is preserved between the semi-lunate carpal and the radius in
both wrists of the Berlin and Eichstatt specimens. It, presumably, is the radiale.
Among theropods, a very similar carpal construction is known in
Deinonyclzrts (Fig. 10B) and apparently in Velociraptor (Fig. lOC), and a
comparable semi-lunate carpal has been found in Stenonychosaums (Russell,
1969) and the Yale specimen of Coelurus (= Ornitholestes?). The semi-lunate
carpal of Deinonyclius has exactly the same relationships and form as in
Archaeopteryx. Also, there is a smaller carpal between metacarpal 111 and the
ulna, just as in the Eichstatt specimen.
Radius and ulna. These bones provide little detailed evidence for or against
an Archaeopteryx-theropod relationship. Both elements are long and very
slender, and distinctly bird-like, but in general proportions they are more
similar to those of “long-armed” theropods such as Deinonychus, Ornitholestes
and Struthiomimus than anything else (see Fig. 12). They are shorter than

Figure 10. Carpus and metacarpus of Archaeopteryx (A), the Eichstatt specimen. compared
with those of Deinonychus antirrhopus (B), Y .P.M. 5205. and Velociraptor mongoliensis (C).
Arrows point to the distinctive half-moon-shaped distal carpal in each, a feature that is unique
to certain theropods and Archaeopteryx. That condition, coupled with the equally distinctive
short form of the first metacarpal. also illustrated here, is considered o f critical phyletic
importance. A and B represent right wrists, C shows the left wrist and hand. Scale divisions in A
= 0 . 5 mm; scale line in B = 3 cm. Photo C provided through the courtesy o f Dr Z .
Kielan-Jaworowska and reproduced by permission of Dr R . Barsbold.
114 J . H. OSTKOM

either the manus (digit 11) or the humerus, a condition that occurs in the above
theropods and some others, but not in most modern birds or pseudosuchians.
Humerus. Although rather bird-like in its general morphology, the humerus
of Archaeopteryx also closely resembles those of several of the small theropods
(Fig. 1l), such as Coelitrus, Ornitliolestes, DrinonycCius and perhaps
V&ciraptor. The shaft in all is long, slender and slightly curved. There is a long,
high and well-defined deltopectoral crest, but little or no development of
internal or external tuberosities or of a bicipital crest, as occur in modern birds.
These same features are also lacking in the theropod humerus.

Figure 1 1 . Comparison of humeral morphology of Archaeopteryx. Berlin specimen (11); and

theropods Deinonychus antirrhopus, A.M.N.H. 301 5 ( A ) ; Ornirholesres hermanni. A.M.N.tI.
619 ( B ) ; and Microwenator celer.. A.M.N.H. 3041 (C). All humeri are right elements viewed in
dorsal aspect and drawn to the same length for better comparison. Kelative sizes are indicated
by the vertical scale lines which equal 3 cm. Compare with Fig. 25.

Forelimb summation. Individually, each of the forelimb components in

Archaeopteryx shows some degree of morphological resemblance to the
corresponding elements in certain theropods. This resemblance, as we have seen
above, is most pronounced in the distal elements, which presumably are the
more specialized components of the forelimb. Considered collectively, the
resemblance still holds and is strengthened by dimensional aspects and
intermembral proportions (Fig. 12). Tucker (1938b) and others have argued
that an invariable trend among bipeds is a reduction of forelimb length, referring
in particular to theropod dinosaurs-which, according to Tucker, obviously
could not then have had any evolutionary connection with Archaeopteryx and
bird origins. I t is indeed well known that certain theropods, such as
Tyrannosaurus, Tarbosaurus, Albertosaurus (=Gorgosaurus), and Daspleto-
saums, possessed greatly shortened forelimbs, but it should be evident now
Microvennror celer is a small coelurid theropod recently described (Ostrom, 1970a) from the Early
Cretaceous of Montana.
 ARCHAEOPTERYX A N D T H E ORIGIN O F BIRDS 11s

f:ipure 1 2 . Outline sketches o f the right forelimti skclcton of Archaeopteryx ( A ) compared with
thosr of theropocls Ornit/to/estr.s (H) and fIeittoriyc./tus ( C ) . Ilumeri are drawn t o the same
length t o minimize size-related differences. sizcs Iieinp inllicatcd Iiy the vertical scalc line.; which
equal 5 cm. Notice the cxtrenie relativc lengths o f the hands comparcd with those illustrated i n
Vip. 26.

from the abundant remains of Orriitholestes, Deinonychtrs. Velociruptor,

Ortiitliomimus, Strirthiomimus, Droniic~.iomimus,Gdlirnitiziis. Dciiioclic~irirs,
and others, that elongated forelimbs were characteristic of a number of
theropods-all of which were bipedal. Forelimb length in Deitzoriychus and
Ortiitholestcs is estimated at about 75% of presacral vertebral length. That
compares with approximately 120% in the Berlin Arcliueopteryx and nearly
140% in the Eichstatt specimen, as contrasted with only 40% to 50% in
pseudosuchians.
Pe c to ra 1 girdle
The pectoral girdle, like the manus and forelimb, is remarkably similar in
Archaeopteryx and various small theropods, a fact that has not been generally
recognized before. In fact, the only non-theropod feature of the pectoral girdle
of Archaeopteryx is the furcula, which is well preserved in the London
specimen (Fig. 23) and is partially preserved in the Maxberg and Berlin
specimens. Independent of the plumage impressions, the presence of a furcula
may be considered important substantive evidence of the avian affinities of
Archaeopteryx, but it also has been alluded to (Heilmann, 1926) as the critical
evidence against theropod relationships. Heilmann dismissed theropods as
possible ancestral stock of Archaeopteryx solely on the grounds that they
116 J . 11. OSTROM

lacked clavicles-the presumed precursors of the avian furcula. The supposed

absence of clavicles in theropods is negative evidence only, and thus
inconclusive. But more important is the discovery that clavicles were present in
at least some theropods, as will be discussed later.
Scapula. The scapula of A r c h a c o ~ t e r ~isx long and very narrow or strap-like
(Fig. 13A). The posterior end flares only slightly or not at all, as is shown by

Figure 13. Pectoral girdle o f A rcliueopreryx (A and B), as reconstructed (hy the author) from
the London and Berlin specimens, compared with those of theropods Deinonychus ( C ) . and
A llosuunts (D). A is viewed perpendicular to the plane of the scapular blade (dorsal ?). B, C and
D are viewed in lateral aspect. Compare this illustration with Fig. 27. Scapulae are drawn to the
same length, relative sizes are indicated by the horizontal scale lines that equal 5 cm. Notice the
distinctive strap-like form of the scapulae.

the London, Berlin and Maxberg specimens, contrary to the flared condition of
most reptilian (other than theropod) scapulae, nor does it taper posteriorly as
in modern birds. Its form is remarkably similar to the scapulae of
Struthiomimus, Ornitliolestes, Deinonychus, Velociraptor and most other
theropods. As far as I know, this narrow, parallel-sided, strap-like scapular form
occurs only in Archaeopteryx and theropod dinosaurs, and in slightly modified
form in modern birds.
Coracoid. The coracoid is preserved apparently co-ossified or very firmly
 ARCHAEOPTERYX A N D THF, ORIGIN OF BIRDS 117

articulated with the scapula in the first three specimens of Arc/iacoptcvy*.u.This

firm union is best indicated by the right scapulo-coracoid (unnoticed by Heller,
1959) in the Maxberg specimen, where it is preserved still united, although
displaced some 8 or 9 cm from the pectoral region and the rest of the skeleton.

Figure 14. A, Left coracoid of Archaeopteryx lithographica (London specimen, underside of

main slab; B, same-scale reconstruction o f the same coracoid seen in anterior view. For
comparison, theropod coracoids, all viewed in antero-lateral aspect; C. Deinonychus
antirrhopits, Y.P.M. 5 2 3 6 (drawn reversed for Comparison); D. Allosaurus sp., Y .P.M.
uncatalogued. Arrow 1 indicates the glenoid and arrow 2 points to the “biceps tubercle”. Scale
divisions in A = 0 . 5 mrn: scale line i f i C and D = 5 cm.
118 J. H. OSTHOM

Coracoid morphology is best seen in the left coracoid of the London specimen,
well exposed on the underside of the main slab (Fig. 14A, B). As preserved
there, the coracoid is relatively large, subrectangular and plate-like and is
shaped very much like that of theropods. I t is not elongated into the stout,
strut-like form of modern birds. Just anterior to and below the glenoid
portion and below and lateral to the supracoracoid foramen, there is a
prominent tubercle or process. This feature, referred to as the biceps tubercle
(Walker, 1972), is also well developed in many theropods, including
D e i / i o n ~ ~ / i i iand
s , has been interpreted as the probable site of origin of M.
biceps brachii (Ostrom, 1974b, 1976). Figure 14 shows the strong
resemblance between the coracoids of Archaeoptcryx and some theropods.

Hindlitnb and p e s
Like the hand and forelimb, the foot and hind leg of Arc/raeopter.vx have
been compared with those of theropods (see especially Heilmann, 1926). There
is also a marked similarity to the hind leg and foot of modern birds, which,
together with the plumage and the (false) avian pattern of the pelvis, has been
taken as strong evidence for the avian identification of these specimens.
f m Parts of the pes are preserved in all five skeletal specimens of
Archueopferyx, but the most complete and best examples are in the new
Eichstitt specimen and the Berlin specimen. The pes is a four-toed structure in
which the fifth toe has been lost and the first toe has been reversed to a
posterior position, opposing the other toes (Fig. 15A). The principal supporting
digits are 11, I11 and IV, with the third the longest and the other two somewhat
shorter and subequal in length. Digit I, the hallux, is much shorter than the
others and is somewhat elevated on the metatarsus. The phalangeal formula is
primitive (2-3-4-5) both in Arcliaeopteryx and in all adequately known
theropods. The relative lengths of the phalanges are also primitive, with the
proximal phalanx being the longest in all toes, in contrast to the phalanges of
the hands. Similar phalangeal proportions are found in theropods and in most
birds (exceptions are birds of prey, including owls, in which the penultimate
phalanges are the longest-as in the hand of Archueopreryx). Figure 1 5 shows
these similarities between Archaeopteryx and certain theropods.
Metaturszts. The metatarsus of Arcliaeopteryx consists mainly of three
complete elements, metatarsals 11, I11 and IV. The first metatarsal is reduced to
a very short wedge-like bone located distally, thus having no contact with the
tarsus. I t is closely appressed to, but not fused with, the postero-medial surface
of the second metatarsal. Until discovery of the Eichstatt specimen, no vestige
of metatarsal V had been recognized, but in the left metatarsus (on the main
slab) of that specimen there is a very thin (less than 0.5 mm) splinter of bone
about 6.5 mm long, extending from the tarsus down the posterior surface of
the fourth metatarsal (Fig. 16A). The shaft tapers very slightly distally and the
proximal end is somewhat expanded into what appears to be an articular head.
If this actually is a remnant of the fifth metatarsal, as Wellnhofer (1974) also
believes, then the metatarsal condition in Archaeopteryx is much more similar
to that in a variety of theropods, such as Coelophysis, Composognathtts
(Fig. 16B), Deinonychrts, Velociraptor, and Stenonychosuurus, than it is to
either modern birds or pseudosuchians.
ARCHAEOPTERYX AND THE OKICIN O F BIRDS 119
120 J. H. OSTKOM

Figure 16
 A R C H A E O P T E R Y X A N D T H E ORIGIN OF BIRDS 121

Over the years there have been different opinions as to whether the three
main metatarsals of Archaeopteryx were fused or not (Owen, 1863; Marsh,
1881b; Dames, 1884; Petronievics, 1925; Heilmann, 1926; de Beer, 1954b;
Heller, 1959). It is possible that the metatarsus is not fused in all of the
specimens, although evidence against this is the fact that there is no physical
separation or displacement of any of the three main metatarsals preserved in
any of the specimens of Archaeopteryx. Moreover, the Maxberg specimen, as
reported by Heller (1959), reveals no discernible physical separation or suture
between the proximal ends of the three main metatarsals. X-rays indicate
fusion of these elements proximally, but not distally (Heller, 1959: pl. 13-1).
Wellnhofer (1974) concluded that the metatarsus was not fused in the new
Eichstatt specimen, but that specimen is almost certainly an immature
individual. Fusion of the metatarsals, even if only partial and variable, is the
most significant difference between the metatarsus of Archaeopteryx and that
of most theropods. With the exception of Syntarsus (Raath, 1969) and
Ceratosaurus (where it may be pathological), fusion of the metatarsus is not
known in theropods. (Neither is it known in ornithischians or pseudosuchians.)
Despite the bird-like fusion (even if incomplete), the metatarsus of
Archaeopteryx is distinctly non-avian in another feature. There is no sign
whatever of the prominent posterior crests of the hypotarsus (entocalcaneal
and ectocalcaneal ridges) on the proximal posterior surfaces. This condition is
likewise characteristic of all theropod metatarsi. Considering the prominence of
the hypotarsus in modern birds, we can presume that the condition in
Archaeopteryx and theropods reflects relatively poor leverage for the M.
gastrocnemius and probably is a primitive condition.
Tarsus. The tarsus is not clearly preserved or fully discernible in any of the
specimens of Archaeopteryx, but it ‘Ippears to consist of two proximal
elements, astragalus and calcaneum, which are co-ossified with the tibia and
fibula, and two or perhaps three distal tarsals which are fused to the
metatarsus. Thus Archaeopteryx would appear to have had at least an incipient
tarsometatarsus, although not yet fully avian. Consequently, the ankle joint is a
mesotarsal joint, as in modern birds, and also as in theropods and ornithopods.
The astragalo-calcaneum is recognizable in all but the Teyler specimen, but is
most distinct in the London and Berlin specimens. Both of the last, and the
Eichstatt specimen, clearly show a well developed ascending process of the
astragalus closely applied to the anterior surface of the tibia1 shaft (see Fig.
17A, B). This same feature is present in all adequately known theropods,
except Syntarsus (Raath, 1969). It also occurs in very early developmental
stages of many birds, but is not recognizable in mature individuals. An anterior
ascending astragalar process is not known in any ornithopod, despite their
having a mesotarsal joint, nor is it known in any pseudosuchian.
The Eichstatt specimen provides the best evidence concerning the distal
tarsals, although the details are not beyond challenge. The left ankle, preserved

Figure 16. Left metatarsus and tarsus of the Eichstatt specimen o f A r c h a e o p t e r y x (A) showing
the splint-like fifth metatarsal, compared with the same features of the right foot of
Compsognathus longipes (B). Scale divisions = 0.5 rnm. Mtt. Metatarsal.
122 J . H. OSTROM

Figure 1 7
 ARCHAEOPTERYX AND THE ORIGlN OF BIRDS 123

on the main slab, shows two distinct cap-like distal tarsals tightly appressed
against the proximal ends of metatarsals I1 and 111. In the Maxberg specimen,
metatarsals 111 and IV are capped by a distal tarsal, whether by one or two
cannot be determined. Probably, this represents the same element that caps
metatarsal I11 in the Eichstatt specimen. From this we can conclude that the
tarsus of Archaeopteryx had a minimum of at least two distal tarsals and
perhaps as many as three. Wellnhofer (1974) suggests that a third tarsal may
cap the fifth metatarsal. Perhaps the most important feature, however, is that
some distal tarsals, whatever their number, were at least partly fused to the
metatarsus, as is clearly shown by the Maxberg and Eichstatt specimens.
The partial fusion of tarsals to metatarsals, and probably of the main
metatarsals to each other, approaches the condition of the modern avian
tarsometatarsus more than that of any other group. In theropods, distal tarsals
are preserved in Syntarsus, two of which are fused with the second and third
metatarsals. Only two distal tarsals (unfused) are known in Deinonychus
(capping metatarsals 11, 111 and IV) and Allosaurus (capping metatarsals 111 and
IV only). Struthiomimus has three distal tarsals applied to the proximal ends of
metatarsals I1 and IV.
No final judgement can be made at this time as to the exact nature of the
distal tarsalia of Archaeopteryx, but they appear to approach the theropod
condition more closely than that of pseudosuchians. This is consistent with the
theropod-like foot, the nature of the proximal tarsals, and the mesotarsal joint.
Tibia and fibula. These elements are perhaps less distinctive than other
elements of the hindlimb, but nevertheless collectively they are intermediate
between those of theropods and later birds. Both bones are relatively more
slender than in theropods, even the very small forms like Cornpsognathus and
Microvenator, but are more robust than in most later birds. The fibula is
complete, extending from the knee to the tarsus, and is not fused to the tibia,
although it is closely applied to the tibia throughout its length, reminiscent of
the condition in some theropods (Compsognathus, Deinonychus, Struthio-
mimus). In most modern birds the fibula does not reach the tarsus. and is rarely
fused to the tibia except distally. In their relative lengths these two bones are
more bird-like than theropod-like, being about 3 5% longer than the femur.
Among theropods, only in ornithomimids, Microvenator, Compsognathus( ?),
Saurornithoides and Deinonychus (Ostrom, in press) is the length of the tibia
known to exceed that of the femur, but never by more than 15%, whereas this
condition is virtually universal among modern birds. The tibia of
Archaeopteryx bears a prominent internal cnemial crest, as in both birds and
theropods, but unlike modern birds it has no external cnemial crest.
Femur. The femur o f Archaeopteryx, like the epipodials, appears to be
intermediate in its morphology between those of theropods and modern birds.

Figure 17. Comparison of the tarsus in Archaeopteryx and two theropods. A, Berlin specimen
of Archaeopteryx, left tarsus in antero-medial aspect; B, London specimen of A rchaeopreryx.
left tarsus in antero-medial view; C, Microvenator celer (A.M.N.H. 3041), left tibia and
astragalus in antero-lateral aspect; D. Deinonychus anfirrhopus (Y.P.M. 5 2 2 6 ) , left astragalus
and calcaneum in anterior view. Arrows mark the prominent ascending process of the
astragalus. a feature found only in theropods. Archaeopteryx and early stages of modern bird
ontogeny. Scale divisions in A and B = 0 . 5 mm; in C and D = 1.0 mm.
9
124 J. H. OSTROM
Although shorter than the tibia, it is relatively longer and more slender
compared with that of many birds and all larger theropods. I t is also longer and
more slender than that of some small theropods, such as Compsognathus.
Perhaps its most distinctive character is the pronounced antero-posterior,
non-sigmoidal curvature that is well preserved in all specimens (Fig. 18). Similar
curvature occurs in most birds and all small theropods (Compsognathus,
Coelurus. Ornitholestes, Microvenator and struthiomimids), but not in the
larger forms. This may be a reflection of normal femur position-nearly vertical
in the heavy bipeds such as Allosaunts, versus an extended, nearly horizontal,
parasagittal attitude in the smaller theropods, as in living birds.
As for structural details, the head is sharply offset from the shaft at about
90°, but is angled slightly forward of the transverse plane. The Berlin
(underside) and Eichstatt (main slab) specimens seem to show that the head is
nearly hemispherical in form. Lateral to the head, two distinct but poorly
defined prominences are well preserved in both the London and Berlin
specimens (Fig. 18D). The larger and uppermost of these prominences lies just
lateral to, and slightly behind the head and may correspond to the trochanter
major of modern birds. On the other hand, it corresponds better to the “greater
trochanter” of theropod femora (see the femur of Microvenator, Fig. 18D, E).
Situated where it is, below, as well as behind and lateral to the head, there
seems to be no way in which this structure could have articulated against an
“anti-trochanter” of the ilium, as in modern birds. Hence we may presume that
in Archaeopteryx it served exclusively for insertion of the “gluteal” muscles.
Below, lateral and anterior to this “greater trochanter” is a smaller and less
well defined prominence or swelling. No comparable feature is located in this
area in modern birds (unless it is the anterior part of the trochanter major), but
theropod femora often bear a small to moderate process here, sometimes as a
high flange and sometimes as a low knob. This feature is commonly referred to
as the “lesser trochanter”. Neither of these “trochanters” is the true
homologue of mammalian trochanters, and they may not be homologous with
theropod “trochanters” either, but the similar location in Archaeopteryx and
known theropods is suggestive, to say the least.
A distinct knob also occurs below and posterior to the “lesser trochanter” in
Archaeopteryx, on the external posterior surface of the shaft. A similar feature
occurs at this site in Microvenator and Deinonychus (Ostrom, in press) and
various other theropods. The functional significance of this structure (here
termed the posterior trochanter), is not known.
Hindlimb summation. As with the forelimb, each of the hindlimb elements
in Archaeopteryx has striking morphological resemblances to the same
elements in various theropods. Considered as a whole, the hindlimb is perhaps
more bird-like than theropod-like, particularly because of the slender form and
the respective proportions of the several segments. Compared with total
forelimb length, hindlimb length is also more avian than theropodian, but is
closer to the latter than to most pseudosuchians (see Fig. 3 1).

Pelvic girdle
Reference has already been made to the importance attributed by past
workers to the pelvis, particularly because of the apparent bird-like orientation
of the pubis as it is preserved in the Berlin specimen. As I demonstrated earlier,
 ARCHAEOPTERYX AND T H E ORIGIN OF BIRDS 125

t F

tm
P’
Figure 18. Right femur, Berlin specimen of Archaeopteryx (A) compared with the left femur of
Microwenator celer in lateral (B) and anterior ( C ) views. Notice the simple forr-aft curvature in
both. D. E and F contrast the proximal ends in lateral (upper series) and proximal (lower series)
views of the right femora of Archaeopteryx (D), Microwenator (E), and a modern carinate,
Catharres (F). Scale divisions in A = 0 . 5 mm. in C and D = 1.0 mm. Vertical scale lines =
1 0 mm. gt. Greater trochanter; It. lesser trochanter; tm. trochanter major; pt. posterior
trochanter. See Fig. 3 0 .
126 J. H. OSTROM

there is considerable evidence (see pp. 102-9) that establishes beyond any doubt
that the pubic position preserved in that specimen is not the natural position.
Hence the supposed similarity to the ornithischian pelvis cannot be verified in
this or in any of the other specimens of Archaeopteryx. The original pubic
position cannot be determined with certainty either, but it appears probable
that in the Eichstatt specimen the preserved position approximates the natural
position. This orientation is intermediate between the modern avian condition
and that of theropods.
Pubis. The pubes are best preserved in the Eichstatt, Berlin and London
specimens, with only fragments preserved in the other two. The London
specimen establishes the presence of a long symphysis uniting the pubes distally
(Fig. 20A) and extending over nearly half their total length. A pubic symphysis
is a distinctly reptilian condition, in contrast to birds where the pubes are fused
only in the ostrich.
The pubis of Arctiaeopterj)x is very long, almost as long as the femur, with a
narrow cylindrical shaft. The distal extremity is enlarged in the sagittal plane
into a backwardly directed foot-like expansion (Fig. 19D, E). The London
specimen seems to show this feature expanded transversely, as well as
longitudinally, but such is definitely not the case in the Berlin, Eichstatt or
Teyler specimens. The size and shape of the distal expansion vary somewhat
among these last three specimens, perhaps because of varying degrees of
preservation of associated cartilaginous tissue (see this region in the Eichstatt
counterpart slab, Fig. 19E). Proximally the shaft expands moderately in an
anterior-posterior direction. The Berlin specimen seems to show a sharp angular
forward projection (Fig. 20B) just below the pubic peduncle of the ilium.
However, rhe Eichstatt specimen clearly lacks any such projection, and in view
of the dislocation of the pubis in the Berlin specimen, that feature is perhaps
best considered as an artifact. Indeed, if the pubis is rotated forward, as I have
suggested (see Fig. 8C), an orientation approaching the condition preserved in
the Eichstatt, Maxberg and Teyler specimens results, and this anterior
projection of the pubic peduncle disappears, becoming part of the junction
between the pubis and the ilium.
The morphology of the pubis of Archaeopteryx is obviously similar to the
pubic form in a variety of theropods, especially in its long narrow shaft, the
foot-like distal expansion, the long symphysis and the great length relative to
that of the femur. Compare this morphology with that of Struthiomimus,
Coelurus, Compsognathus, Microvenator or, in fact, that of any theropod large
or small (Fig. 19A, B, C, F). Virtually every theropod known from adequate
material possesses this distinctive form of the pubis, there being only small
differences in the shape or size of the distal expansion. To the best of my
knowledge, this type of pubis is unique t o theropods and Archaeopteryx.
Ilium. The ilium is not complete in any of the present specimens of
Archaeopteryx, but a composite reconstruction is possible, using the Berlin,
Eichstatt and London specimens (see Fig. 20). This reconstruction indicates a
long, low blade with a gently convex dorsal border, a short and sharply tapered
posterior process and a longer and more rounded anterior process. The London
specimen suggests the presence of a slight downward expansion of the inferior
margin of the anterior process a t about mid-length, but this is not verified by
the other specimens. The acetabulum, which is open as in all higher archosaurs,
 ARCHAEOPTERYX AND THE ORIGIN OF BIRDS 127

Figure 19. Comparison of pubic morphology in some theropods (Srrurhiomimus, A; ~Ilosourus,

B; Gorgosaurus, C ; and Microvenator, F) with that of the Berlin (D)and Eichstatt ( E )
specimens of Archaeopteryx. Long narrow pubic shafts and prominent distal expansions
(arrows in D and E) are features that are known only in the pubes of theropods and
Archaeopteryx. Horizontal scale lines in A. B and C = 20 cm; scale divisions in D = 0.5 mm; E
and F = 1.0 mm. Compare with Fig. 32.
12x J. tl. OSTROM

Figure 20
 ARCHAEOPTERYX AND THE ORIGIN OF BIRDS 129

is situated behind the mid-point of the ilium. The pubic peduncle is much more
robust than the ischiadic peduncle, a typical saurischian condition, but there is
no indication of any expansion that might be equated with a pectineal process.
(The anterior “projection” of the pubis in the Berlin specimen, mentioned
above, might be construed as a pectineal process, but, as I have shown, this
feature is probably an artifact of the dislocation of the pubis and not a real
process at all.) There is n o anti-trochanter.
In its general shape, the ilium of Arc/zaeoptc’r~is.like the pubis, is strongly
reminiscent of those of theropods. ’The most conspicuous differences are in the
relative lengths of the anterior and posterior processes (the posterior process is
generally the longer of the two in theropods), the posterior position of the
acetabulum (a reflection of the short posterior process) and the lower total
height of the ilium in Archacvptcrjix. Nevertheless, the theropod ilium is much
closer to that of Arclzaeopterys than is that of any other reptile-or of any
modern bird.
Ischiiim. This bone is best revealed, although incompletely, in the Eichstatt
specimen. The form is unique (Fig. 20), but it is confirmed by the poorly
preserved but similar ischium of the London specimen. There appears t o have
been a rather stout peduncle buttressed against the posterior proximal portion
of the pubis (this part perhaps corresponds to the pubic side of the “fracture”
line between the pubis and ilium in the Berlin specimen). Along the posterior
border there are two sharp triangular projections, one proximal just beneath
the iliac articulation and the other near mid-length. The most distinctive
feature is the distal extremity which is bifurcated into a narrow, curved
posterior process and a more robust anterior projection. The functional
significance of this strange configuration is not known, especially since nothing
comparable has been recognized in any other taxon. This unique morphology
may be related to one other surprising aspect of the ischium in Archaeopteryx:
its greatly abbreviated length as compared with that of the pubis, and the much
greater relative length of the ischium in all theropods (but see below),
pseudosuchians and modern birds. For example, there is no evidence on the
proximo-ventral border of an obturator process, as occurs in theropods. Is it
possible that the more anterior of the two processes at the distal extremity
might be the obturator process on a greatly shortened ischium? Whatever the
ancestry of Archaeopteryx, the ischium appears t o have been greatly shortened
and that may obscure the identity of the several seemingly unique processes.
Although the morphology is different, it is interesting that the ischium of
Deirzonychus also is extremely short relative to the pubis (Ostrom, in press).
None of the specimens of Archaeopteryx provides any information about
the ischiadic symphysis. Presumably this primitive condition was retained, in
view of the very long pubic symphysis that is visible in the London specimen.
However, the peculiar form of the ischium, especially of the distal processes
and the posterior projection at mid-length, and the short length suggest that the

Figure 20. Morphology of the ilium and ischium in Archaeopteryx, as preserved in the London
(A1 and 2). Berlin (B1 and 2 ) and Eichstatt (C1 and 2 ) specimens. Compare the shape of the
ilium as preserved in these specimens with those of some theropods. as shown in Fig. 19A. B
and C. and certain thecodontians (Fig. 32). Scale divisions in A = 0 . 5 mm; in B, C = 1.0 mm.
isc, Ischium; It, left; pu, pubis; r t . right.
130 J. H. OSTROM

Figure 21
 ARCHAEOPTERYX AND THE ORIGIN O F BIRDS 131

ischiadic symphysis may have been reduced or entirely lost. The latter, of
course, is an avian condition unknown in theropods.

Skit11 arid jaws

Heilmann (1926) presented a rather detailed reconstruction of the skull and
jaws of Archacoprer.v.u based entirely on the Berlin specimen (see figs 4, 5c,
105-2, 118-4, and 133c of Heilmann, 1926). Despite the details shown there,
the actual specimen does not permit such detailed and precise conclusions. I t is
badly crushed and the bones are extensively fractured, chipped and
distorted-to the extent that very few cranial or mandibular sutures are
unmistakably identifiable (Fig. 2 1A). Heilmann’s reconstruction has been
republished by many authors and subsequent interpretations and hypotheses
have been based on it (for example, see Bock, 1964). Quite probably, some
authors have been unaware of the inadequate basis of Heilmann’s
reconstruction, and understandably so unless they have had the opportunity to
examine the Berlin specimen itself. Bock (1964) pointed out the damaged
condition of the Berlin skull and warned about the false “authenticity” that
had resulted from the repeated publication of Heilmann’s interpretation of that
skull. Nevertheless, Bock accepted that reconstruction as “a reasonable one”.
Fortunately, the Eichstatt specimen now provides a comparative basis for
evaluating and correcting past reconstructions of the Berlin skull (Fig. 2 1B).
Prior t o examining the Eichstatt specimen, I had concluded, after many
hours of microscopic inspection of the Berlin specimen, that the only
indisputable features preserved there are the following:
(a) The skull is triangular in profile with a sharply tapered snout and a
relatively deep and expanded temporal region.
(b) Only three major skull openings are visible in lateral aspect: a narrow,
elliptical and obliquely oriented external naris; an intermediate-sized triangular
antorbital fenestra; and a relatively enormous circular orbit containing a
sclerotic ring composed of an indeterminate number of plates. The temporal
region is imperfectly preserved, although clearly inflated, so despite the general
conformation of the preserved regions which suggests the presence of one or
more temporal fenestrae, the existence of the latter cannot be established from
this specimen-all previous statements t o the contrary notwithstanding.
(c) Teeth are present, apparently set in sockets, more or less isodont and of
inflated conical form. The tooth row apparently did not extend behind the
midpoint of the antorbital fenestra.
(d) The mandible is long and very shallow, and bears teeth which, like those
in the maxilla, also seem to be thecodont. There seems to be a relatively long
retroarticular process and there may be an external mandibular fenestra, as
suggested by Heilmann (1926), but this last is far from certain (see later
comments).

Figure 21. Skulls of the Berlin Archueopreryx ( A ) and the new Eichstatt Archaeopteryx ( B ) .
Line drawings below (from Wellnhofer. 1974) interpret the details preserved in the Eichstatr
skull. For a reconstruction of the Eichstatt skull, see Fig. 33C. Scale divisions in A = 0 . 5 mm; in
B = 1.0 mm. Compare the skull morphology preserved in these two specimens with that
illustrated in Fig. 22.
132 J. H. OSTKOM

(e) The quadrate-squamosal region is badly damaged, but judging from the
backward extension of the parietal area relative to the preserved position of the
jaw articulation, and the coincidence of the anterior extremities of upper and
lower jaws, the suspensorium was probably inclined forward (descended
anteriorly).
(f) Associated with the lower jaw are two delicate, rod-like bones, 0.5 mm
in diameter, that are probably part of the hyoid apparatus.

Without dwelling on the unverifiable positions and courses of the cranial

sutures reconstructed by Heilmann, I should call attention to just one recent
interpretation that has resulted from Heilmann’s restoration. This is the
suggestion by Cracraft (1970) that the mandible of Arc.liuc.opter,vx provides an
example of mosaic evolution. Cracraft’s point is that Arcliueopterys (according
to Heilmann’s reconstruction) possessed an external mandibular fenestra
bordered below by the dentary, as in birds, rather than by the angular as in
reptiles. The toothed mandible thus features both reptilian and avian
characters. Much as I would like to accept this interpretation, the highly
fractured condition of the lower jaw bone (or bones) that borders the supposed
mandibular fenestra, either below or above, makes it impossible to certify their
identifications. In fact, the fractured upper margins of the supposed fenestra
leave considerable doubt as to the very existence of a “fenestra”-a doubt
which has not been removed by the new Eichstatt specimen.
The Eichstatt specimen (Wellnhofer, 1974) includes a much more complete
and better preserved skull which undoubtedly will permit more reliable
interpretations. This skull, together with Wellnhofer’s interpretation of it, and
the Berlin skull are illustrated in Fig. 21. Here it can be seen that the major
features listed above are verified by the new specimen. Wellnhofer’s study
(1974) has already produced a number of new details, as well as confirming
some of Heilmann’s interpretations. Most significant for the present study, the
two skulls together establish the presence of a variety of features that are also
known (though not exclusively in all cases) in some small theropods such as
O mit k oles tes, Compsogt ia t Ii us. Velocirup tor. per h ap s Sutr ro rn ittioides, an d in
some struthiomimids (Fig. 22). These features are:

(a) A sharply tapered snout.

(b) Long elliptical external naris bounded almost exclusively above and
below by the premaxilla.
(c) A large triangular antorbital fossa which contains two small anterior
openings and a large triangular posterior fenestra.
(d) A slender, nearly vertical preorbital bar separating the antorbital fossa
and the orbit.
( e ) A large circular orbit which contains a large sclerotic ring.
(f) A thin straight jugal bar.
(g) A stout quadrate of moderate length which is inclined forward (i.e.,
descends anteriorly).
(h) A lower jaw which is unusually shallow and has a conspicuous
downward bend behind the tooth row.
(i) A long retroarticular process.
 ARCHAEOPTERYX AND T H E ORIGIN 01: BIRDS 13:

I
.....
/
......
..........
............ . A

3 :K:;
I . . . . . .. .. .. .. .. .. .. .. ..

y
&
~

,I i . . . . . . . . . .
. . . . . . . . . .
. . .. . .. . .
. . .. .. .
. .......... ......
.... .._.

........

Figure 22. Skulls of A, Compsognarhus longipes (partly after Heilmann, 1926); B,

Sauromithoides mongoliensis (after Russell, 1969); C. Gallimimus bullatus (from Osmblska.
Roniewicz and Barsbold, 1972). Horizontal scales in A and B = 10 mm. Compare these skulls
with those illustrated in Figs 21. 3 3 and 34.
134 J. H. OSTROM

Important features that have not yet been recognized or confirmed in either
specimen are:
(a) The positive existence of either temporal fenestra, although the quadrate
apparently did not cover the entire temporal region and the general
conformation of the temporal region suggests the presence of fenestrae.
(b) Positive existence of an external mandibular fenestra.
(c) A complete bony postorbital bar separating the orbit from the temporal
region.
The last item is particularly important because it bears on the questions of
kinesis and a streptostylic quadrate, both of which are considered critical
conditions in bird ancestry (Versluys, 1912; Simonetta, 1960; Beecher, 1962;
Bock, 1964). Bock, for example, has theorized that Archaeopteryx was
probably both mesokinetic and streptostylic. The Eichstatt specimen seems to
support Bock’s speculations on both counts. Notice especially the distinct
discontinuity between the parietals and frontals in that skull (Fig. 2 1 B).
Beecher (1962), on the other hand, believed that there was no evidence (and
there was none in 1962) for mesokinesis and that for this reason
Archaeopteryx must have been a side branch off the main path of avian
evolution. ( I t is not clear to me why avian kinesis could not have developed
after Archac~opteryx.)Wellnhofer (1974) is convinced that the Eichstatt skull
was both kinetic (possibly mesokinetic) and streptostylic.
As for theropod kinesis, Versluys (1910, 1912) believed that most dinosaurs,
including theropods, were kinetic. In a recent study, Colbert & Russell (1969)
postulated that the small theropod Dromaeosaztrus probably had a mesokinetic
skull. On circumstantial evidence I suggested that the skull of Deinonychits
might also have been kinetic (Ostrom, 1969). In any case, as Bock (1964)
observed, there is no direct proof of mesokinesis in Archaeopteryx (that is still
true), and this holds for all theropods as well.
For some, the possible absence of an external mandibular fenestra in the
Eichstatt specimen may come as a disturbing surprise. After all, most modern
birds and nearly all archosaurs, including thecodonts and theropods, possess
this fenestra. I cannot testify to its universal occurrence in pseudosuchians, but
it is variable in theropods, being absent in Ornittiolestes, Velociraptor and
perhaps Compsognathus. It is also not universally present in modern birds! Its
presence or absence in Archaeopteryx. however, is not crucial for the simple
reason that the question is beyond resolution-at least until such time as the
reverse (left) side of the new Eichstatt specimen can be prepared. The region of
the external mandibular fenestra is not intact on the right side of that
specimen. The posterior parts of the dentary and the adjacent regions of the
surangular and angular were apparently lost when the Eichstatt slabs were split,
so that the inner surface of the splenial is exposed (Wellnhofer, 1974), rather
than the external mandibular surface.
Some of the features listed above are not exclusive to Archaeopteryx and
theropods and thus cannot be used as arguments in favor of a theropod
ancestry (i.e., sclerotic ring, tapered snout, circular orbit), but they d o add to
the general “theropod-like” appearance of Archaeopteryx. On the other hand,
there are a number of features that are not known in pseudosuchians but which
are typical of theropods, such as elliptical naris, triple antorbital fenestrae, the
 ARCHAEOPTERYX AND THE ORIGIN O F BIRDS 135

anterior inclination of the quadrate, long retroarticular process, and shallow

mandible with a downward bend*. These clearly argue in favor of theropod
affinities and militate against a close relationship with pseudosuchians.
Another important cranial feature that by itself does not favour either
theropod or pseudosuchian ancestry is the elevated posterior position of the
occipital condyle and foramen magnum; this can be inferred from the position
of the cervical series, well above the dorsal end of the quadrate (see the Berlin
and Eichstatt skulls, Fig. 21). This primitive condition is characteristic of both
pseudosuchians and rheropods, in contrast to all later birds where the occipital
condyle and foramen magnum are at the base of the skull, well below the level
of the upper extremity of the quadrate. In this feature, Arclzacopter.vx was far
from avian.

Vertebral column
Little precise anatomical information pertinent to the question under
consideration here is obtainable from the vertebrae preserved in the various
specimens of Archaeopteryx. In most respects the column appears more or less
primitive. For example, it is generally agreed that the vertebrae lack the
saddle-shaped articulations of the centrum that are characteristic of modern
birds. The two or three surfaces of centra that are exposed in the London
specimen seem to be slightly concave and several authors accordingly have
reported the vertebrae of Archaeopteryx to be amphicoelous. X-rays of the
Maxberg and Berlin specimens seem t o substantiate this condition for some
parts of the column, but the question is still open for other vertebral regions. If
the vertebrae of the entire column were amphicoelous, this would be consistent
with either theropod or thecodontian affinities and would prove nothing about
bird origins. One possibly noteworthy feature is the apparent presence of small
pleurocoels on some of the posterior dorsal vertebrae in both the London and
Berlin specimens. Such pleurocoels are not known in pseudosuchians or
ornithischians, but they are a common feature among theropods. The caudal
vertebrae of Archaeopteryx, as is revealed most clearly in the Eichstatt
specimen, are typical of long-tailed reptiles (theropods included) both in their
form and in their progressive change in length and morphology along the tail.
Of greatest interest, though, is the peculiar elongated form of the zygopophyses
of the last 1 5 to 16 vertebrae. While not so extreme as in the dromaeosaurids
Deinonychus or Velociruptor, or in the rhamphorhynchoid pterosaurs, their
form is very reminiscent of the condition in struthiomimid theropods.
Similarly, the chevrons behind the seventh caudal are also greatly elongated
antero-posteriorly and severely flattened dorso-ventrally, as in both
struthiomimids and dromaeosaurids. To the best of my knowledge, these
conditions are not known in any pseudosuchian, or in any other reptile other
than the theropods cited and rhamphorhynchoids.
According to my own count, which agrees with that of Dames (1884), the
vertebral count of the Berlin specimen shows a minimum of 22 presacral
vertebrae including an indistinguishable atlas. Twelve of these are clearly
Another, less direct. but very interesting piece of evidence that hears on theropod-avian
relationships as advocated here is the similarity of the intramandibular joint of Hesperornis and
Ichrhyomis (Gingerich, 1973; Gregory, 1 9 5 1 ) to those of Deinonychus and Dromaeosaurus and the
similarity of the surangular-articular relationships in Hesperornis and Deinonychus.
136 J. El. OSI’KOM

recognizable as dorsals or trunk vertebrae, according to Dames, but I believe at

least 13 and possibly 14 dorsals are present. Including a small undected atlas,
the cervical count is then 8 or 9. The vertebrae of the London specimen are
somewhat displaced and an unknown number are missing. According t o de Beer
(1954a,b), 20 precaudals are preserved in a continuous series. Five of these are
definitely sacrals (de Beer included an unfused sixth, “T-15”). Of the remaining
15, all appear to be trunk elements, but the first is so poorly exposed and
preserved that its identification is extremely doubtful and several others (“T-6”
and “T-7”. by de Beer’s designation) similarly cannot be positively identified
even as vertebrae. The Eichstatt specimen shows 22 distinct presacrals, eight of
which appear to be cervicals. Adding one more for the undetected atlas, the
Eichstatt vertebral column consists of nine cervicals and 14 dorsals. Wellnhofer
(1974) recognized a fragment of the neural arch of the atlas and independently
arrived at this same presacral count.
Dames ( 1884) estimated that seven sacral vertebrae are concealed beneath
the right ilium of the Berlin specimen, but if the length of the last exposed
dorsal is used as an index of sacral vertebral length, no more than six, and
probably only five segments lie beneath the ilium. X-rays clearly show five, but
the posterior end of the fifth is indistinct, so there might possibly have been a
sixth even though there seems to be insufficient space for it. The London
specimen, as already noted, seems to have five also, but the preservation is very
poor in that region. The Eichstatt specimen clearly has five sacrals by my
count-a number verified by Wellnhofer (1974). The caudal series in the
London and Berlin specimens have been reported at 20 segments in each, but I
think the Berlin specimen may have 21. The Eichstatt specimen positively
includes no less than 22 caudal vertebrae. Wellnhofer counted 23. Thus, from
these three skeletons, the vertebral formula of Archaeopteryx seems to be 9
cervicals, 14 dorsals, 5 (or 6?) sacrals and 20 to 23 caudals. A comparison of
the vertebral counts in the various relevant taxa is as follows:

Archaeopteryx Coelurosaurs Pseudosuchians Ornithopods

Ccrvicals 9 9-10 7-8 9-1 3

(inc. atlas)
Dorsals 14 13-14 13-18 14-17
(typically 17)
Sacrals 5 4-5 2 -4 4-8
(typically 2)
Caudals 20-2 3 30-40 30-40 40-60

While not conclusive, there can be little disagreement that the vertebral count
in Archaeopteryx conforms most closely to that of coelurosaurian theropods.
The vertebral count per se cannot be considered as strong evidence one way
or the other, regarding the question of ancestry. However, another feature of
the column that may be important is the nature of the cervical series. Both the
Berlin and Eichstatt specimens show that the neck was both long and flexible.
The strongly arched cervical series in both specimens indicates a high degree of
flexibility and its great length sharply delineates the neck from the trunk
region. The resulting picture is that of an animal with a nearly horizontal trunk
 ARCHAEOPTERYX A N D T H E ORIGIN OF BIRDS 137

and a well-defined, mobile, arched neck, supporting an elevated head. This

image is reinforced by the elongated, angled form of the cervicals, as preserved
in the Eichstatt specimen, which also suggests high flexibility. A long arched
cervical series is one of the distinctive characters of “coelurosaurian”, as
opposed to “carnosaurian”, theropods. Notice that this coelurosaurian-like
neck extended back from the rear of the skull in ArchacJoptcJr~~.u-as it does in
coelurosaurs, rather than from beneath as in later birds. The pseudosuchian
neck also extends from the rear of the cranium, but here the neck consistently
is short and poorly differentiated from the trunk in all presently known forms.
Also, there is no evidence that the pseudosuchian neck was particularly
flexible.
Quite probably the long flexible neck in Archaeopter)).u and coelurosaurs
was linked functionally with an active, obligatory bipedal habit. High mobility
of the neck and head would be advantageous in locating and catching prey, but
it also could have contributed to dynamic stability in a biped by producing
small, but quick, shifts of weight in front of the center of gravity, countered by
tail movements on the opposite side of the bipedal pedestal. In this sense, the
short, relatively inflexible neck of pseudosuchians is consistent with our
traditional conception of the most advanced pseudosuchians being only
facultative bipeds, in which a bipedal posture could be maintained only while
in rapid motion, as a result of inertia. In coelurosaurs, the cervical series
constitutes 45% to 55% of the presacral column, whereas in pseudosuchians it
s e e m to range from 25% to 35%. In Archaeopteryx it equals 45%.

Other skeletal elements

Sternum. De Beer (1954b) identified a fragment of the London specimen as
the sternum. The object is so poorly preserved that it defies accurate
description, let alone positive identification. By eliminating all the other
missing elements of the skeleton, de Beer concluded that this fragment must be
the sternum, figuring it as a narrow rectangular transverse bone. De Beer
recounted past interpretations and speculated that this was the object which
Marsh (1 881) claimed was “a well-ossified, broad sternum”. Whatever Marsh
saw is unknown now, for he did not figure it. Of the several other missing
elements dismissed by de Beer, it could just as easily be one or two badly
preserved vertebrae. The X-ray published by de Beer (pl. VI-2) shows it to be a
relatively dense ossification, which seems inconsistent with the thin plate-like
form expected of a sternum. In view of this evidence and the condition in the
other specimens which have no identifiable sterna, I believe that this object
probably represents one or more vertebrae.
If the London sternum is in doubt, there can be no doubt about the
condition in the Berlin and Eichstatt specimens. Dames (1897), after
preparation of the underside of the main slab of the Berlin specimen, claimed
that the sternum was present, at least in part (see his Fig. 1). Thanks to X-rays
kindly supplied to me by Dr H. Jaeger of the Humboldt Museum fur
Naturkunde, the bone which Dames believed to be the sternum can now be
seen to be the right coracoid oriented upright and still articulated with the right
scapula. The left coracoid, which was correctly identified by Dames, has
collapsed under sediment compaction and lies in the plane of the slab. Other
fragments occur in this region, but none appears to represent any part of the
138 J . H. OSTROM

sternum. The Eichstatt specimen, because it is small and thus probably

immature, may not settle the question once and for all, but it contains no
evidence whatever of an ossified sternum. The same is true of the much larger
Maxberg specimen as well.
In view of the above, it must be concluded that there was no ossified
sternum in Archaeopteryx. This has far-reaching implications for the flying
ability of Archaeopteryx, as I have discussed elsewhere (Ostrom, 1974a). I t has
less significance, perhaps, for the question of the origins of Archaeopteryx.
With the possible exception of one specimen of Velociraptor (Kielan-
Jaworowska & Barsbold, 1972; pl. 11-2), ossified sterna apparently are not
known in theropods, despite the fact that they do occur in other saurischians.
More surprising though is the fact that the sternum is also unknown in
thecodontians! By that token, following Heilmann’s example, we should
perhaps exclude pseudosuchians from the ancestry of all later archosaurs, since
sauropods, ornithischians, pterosaurs and crocodilians (as well as modern
birds!) all possess sterna. This would appear to be doubly justified since the
sternum is an endochondral element. However, it is obvious that we should not
make any such rash conclusion on negative evidence only.
Clavicle- furcula question. The London specimen of Archaeopteryx includes
a symmetrical “boomerang”-shaped bone (Fig. 2 3 ) which all previous
investigators have identified as a furcula. I t is situated in the anterior part of
the trunk region between, but not in contact with, the two scapulae. I t is a
robust element with nearly uniform dorso-ventral breadth (about 4.0 mm)
throughout. The exposed surface is convex. A similar bone is partially
preserved in the two slabs of the Maxberg specimen. The dimensions are
approximately the same, but impressions indicate that one side, probably the
posterior surface, is concave. In neither specimen is there any indication of

Figure 23. The furcula (arrow) in the London specimen of Archacopferyx. A similar but
fragmentary furcula is preserved in the Maxberg specimen, and several fragments in the Berlin
specimen may represent parts of the same element. Scale divisions = 0 . 5 mm.
 ARCHAEOPTERYX A N D T H E ORIGIN O F BIRDS 139

what might be considered a hypocleideum. Compared with most modern

carinates, the angle between the rami of this furcula is quite broad,
approximately 80’. In the Berlin specimen, fragments of bone adjacent to the
coracoids have been considered as part of the furcula by several previous
workers. I am inclined to agree with this identification, although it cannot be
proved. In view of its presence in these three specimens, it is something of a
surprise to find that there is no sign of a furcula in the very well preserved
Eichstatt specimen. Its apparent absence may be an artifact of preservation, but
a more likely explanation is that, because of the small size and probable
immaturity of this specimen, the furcula had not yet ossified. (Is its absence
here any less or more significant than its supposed absence in theropods?).
The presence of what appears to be a well developed furcula in the London
and Maxberg specimens confirms the avian status of Arclzaeopferyx indicated
by the feather impressions. I t also has critical significance regarding the origin
of Archaeopteryx and the specific ancestry of birds. Inasmuch as the furcula is
widely accepted as the coalesced clavicles (but see p. 166), it follows that the
immediate ancestor of Arclzaeopterys must have possessed either paired or
fused clavicles. Broom (191 3) did not specify his reasons, but in all probability
the supposed absence of clavicles in all theropods then known is what led him
to conclude that dinosaurs were too specialized, but that pseudosuchians were
“still primitive enough” to include bird ancestors. Heilmann (1926) presented
an impressive comparison of the numerous similarities between birds and
Archaeopteryx on the one hand and coelurosaurian theropods on the other.
And after comparison with other potential ancestral groups, he concluded:
“From this it would seem a rather obvious conclusion that it is
amongst the Coelurosaurs that Ge are t o 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 t o prove these saurians could
not possibly be the ancestors of the birds.” (Heilmann, 1926: 183)
On the basis of that single piece of negative evidence, Heilmann completely
dismissed an impressive array of data and thereby effectively stilled all but a
few advocates of a coelurosaurian ancestry of birds.
Although this will be discussed at length later, it is important to point out
here that the only evidence that has been advanced so far that is contrary to a
theropod ancestry of Arclzaeopteryx is the supposed absence of clavicles in
theropods. This is negative evidence only and therefore inconclusive (like the
absence of a sternum in all known thecodonts). Unless found in perfect
articulation with the scapulocoracoid, the clavicle could easily be mistaken for
a rib fragment. But even more significant is the discovery by Camp (1936) of
the unmistakable presence of a clavicle in articulation with the scapula and
coracoid in the small Triassic coelurosaur Segisaurus. Also, Osborn (1924)
reported the presence of what appears t o be a fused clavicle and interclavicle in
the Cretaceous Mongolian coelurosaur Oviraptor. A remarkable specimen of
Vefociraptor reported by Kielan-Jaworowska & Barsbold (1972) also possesses
what appears to be a clavicle in natural articulation. From these few specimens,
it would seem that the clavicle is not lacking in all theropods and I suspect that
a careful search would uncover others.
Gastralia. Gastralia, or dermal abdominal ribs, are present in all five skeletal
10
140 J . tl. OSTROM

specimens of Arclzuroptrryx, but they are best preserved in the Berlin and
Eichstatt specimens. They are delicate hair-like bones, at least some of which
are “V” or chevron-shaped. Their exact arrangement cannot be reconstructed
with certainty, but they appear to have been rather widely spaced extending up
and backward at a wide angle to the dorsal ribs, very much like the widely
spaced arrangement preserved in Struthiornimiis ultiis (see pl. 24, Osborn,
19 17). Gastralia are present in a variety of reptiles including pseudosuchians
and theropods. Their presence in all of the specimens of Arcliawpfcirj2xmay be
considered as evidence of reptilian origins, but it does not necessarily support
either theropod or pseudosuchian affinities, although the resemblance is closer
to the theropod pattern. I t does, however, add evidence against the
ornithischian affinities of Archaeopferj*x since gastralia apparently are absent
in all ornithopods*.
The presence of gastralia in Archueopterj1.r and their absence in all modern
birds presumably correlate with the unossified sternum that supposedly existed
in the former and with the greatly enlarged and well-ossified sterna that are
present in all the latter. In fact, the gastralia provide positive evidence that the
sternum, whatever its condition in Archaeopteryx. must have been relatively
small. Obviously, the absence of gastralia in modern birds has no significance
relative to the avian affinities of Archueoptrrj*s.

Sit m mury of’tti rropod e videti ce

In summary, a considerable amount of evidence points to a close phyletic
relationship between theropods and Archaeopteryx. The only derived
characters for birds that are present in Archaeopteryx are the furcula and the
evidence of feathers. By contrast, the specimens of Arclzueopteryx possess a
large number of derived characters for coelurosaurian theropods: tridactyl
manus and metacarpus design, construction of the carpus, elongated forelimb,
morphology of the scapulo-coracoid, construction of the pes and metatarsus
with a short elevated hallux, mesotarsal joint with an anterior ascending process
of the astragalus, morphology and orientation of the pubis, morphology of the
ilium, obligate bipedal posture. These must be considered as prima jucie
evidence of very close phylogenetic relationship between Archaeopteryx and
coelurosaurian theropods. To date, the only evidence that has been offered
against a theropod ancestry of birds, the supposed absence of clavicles in the
theropods, is now known to be false. Consequently, no longer is there any
logical reason to dismiss the Theropoda as the most probable immediate
ancestral stock of Archaeopteryx and higher birds.

Tlie psetr dosuchiarr evidence

Most advocates of a pseudosuchian ancestry of birds have followed
Heilmann’s (1926) example and discussed this with specific reference to
Eupurkeria and/or Ornithosuclins (see for example Tucker, 1938b; Simpson,
Cilmore (1924b) illustrated several isolated splint-like bones of Sregoceras (=Troodon), which he
interpreted as abdominal ribs, b u t I believe them to be ossified caudal tendons. They were associated with
such tendons (which are characteristic of nearly all ornithopods) and they arc almost identical in form to
ossified caudal tendons preserved in place in the pachycephalosaurid Homalocephale described by
Maryhska & Osmblska (1974).
 ARCHAEOPTERYX A N D THE ORIGIN OF BIRDS 141

1946; de Beer, 1954b; Swinton, 1960, 1964; and Brodkorb, 1971), implying at
least that these taxa were representative of the supposed pseudosuchian stage in
bird ancestry. At the moment, though, the term Pseudosuchia means different
things to different people. This is due in large measure to the many important
thecodontian discoveries made in South America, Africa and elsewhere in
recent years and the fact that no general consensus has been reached as yet
concerning the relationships between these and previously known forms such as
Eiiparkeria, Orriithosirchirs, S?~hi.riosirchrrs and others. Despite the fact that
most authorities had come to accept the idea that all higher archosaurs and
birds were probably descendant from pseudosuchian thecodontians, very little
was actually known about pseudosuchians other than those mentioned above,
and a few others. Acceptance of that theory is understandable though, because
Broom (1913) and Heilmann (1926) both observed that the “pseudosuchians”
then known were similar in some ways to Arc.haeopterj~.uor later archosaurs,
yet were still primitive in many other features. Today, the picture is
complicated by various new discoveries and it may be some time before the full
significance and relationships of all the new specimens are understood and can
be expressed in a “consensus” classification. For that reason, this section
(which as a result may be somewhat premature) will deal with pseudosuchians
in general (and occasionally with non-pseudosuchian thecodontians), rather
than with a particular kind, to show that the basic coelurosaurian features of
Archacopterjix are not present in these reptiles as they are presently known. 1
hope t o demonstrate that the relationship between Archaeoptiv-y.u and
pseudosuchians is remote.
In recent years, several important events have occurred which have
contributed further to the present confusion over what is or is not
pseudosuchian. In 1964, A. D. Walker came to the conclusion that
Oriiithosrtchits (long recognized as an advanced pseudosuchian and often
referred to as the classic example of the pseudosuchian state) was not a
pseudosuchian at all, but a primitive carnosaurian theropod; this conclusion is
not generally accepted. Later, Walker (1970) proposed a revision of the
traditional Order Crocodilia, recognizing a new Order Crocodylomorpha
composed of two suborders, the Paracrocodylia and Crocodylia. The latter
included all proper crocodilians plus various long-recognized primitive forms
(Protosuchia). To the Paracrocodylia Walker assigned a variety of taxa which
have certain crocodilian tendencies, but which seem to be removed from the
main crocodilian lineage. Among these, Walker included Sphenosiichus from
the Late Triassic of South Africa, which over the years has usually been
classified as a pseudosuchian, even though its affinities with crocodilians have
long been recognized (von Huene, 1925; Broom, 1927).
Bonaparte (1971a), in describing some of the new Triassic reptiles from
Argentina, presented a new classification of all currently known thecodontians
in which he recognized three pseudosuchian infraorders: Ornithosuchia
(including Ornithosuchus and Euparkeria), Sphenosuchia (including with
Sphenosuchus one of the new Argentine forms, Pseu~hesperosuchus,and also
Triassolestes) and Proterochampsia (including Cerritosaunis and the
proterochampsids). Subsequently, Romer ( 1 9 7 2 ~ published
) his first attempt at
classifying the new and the old thecodontians, placing Eiiparkeria and
Ornithosuchus in two, of three, separate families of the Pseudosuchia. He
142 J. H. OSTKOM

placed Sphenosuchus (and YPedeticosaurus and Hemiprotosuchus) in the

crocodilian suborder Protosuchia, which consisted of two families,
Protosuchidae and Sphenosuchidae. Also included incertae sedis in the
Protosuchia was Bonaparte’s Pseiidhesperosuchus.zus. Romer regarded
Cerritosaurus and other proterochampsids as proterosuchians rather than
pseudosuchians.
Except for shuffling some possibly relevant taxa (not all have been
mentioned here) in and out of the potentially ancestral “Pseudosuchia”, these
events would seem to have little bearing on the question at hand. That would
be true except for two other papers by Walker (1972, 1974), in which he
argues that Sphenosuchus, in addition to being close to crocodilian origins, is
also close to the origin of birds.
Some authors may still prefer to consider Sphenosuchus as a thecodontian,
perhaps even a pseudosuchian. But in deference to Walker’s theory, I will not
include Sphenosuchus in the following discussion dealing with the evidence for
a pseudosuchian origin of birds. This should not be taken to mean that I either
accept or reject Walker’s taxonomic assignments. Romer (1972c), has
summarized his views on the alignment of the Pseudosuchia, but these have not
received general acceptance either. Romer’s classification is as follows:
Order Thecodontia
Suborder Pseudosuchia
Family Euparkeridae: Euparkeria (Brownidla), ? Wangisuchus.
Family Ornithosuchidae: Orn ith osu ch us ( YDasygnathus.
Dasygnathoides), Gracilisuchus. Venaticosuchus, Riojasirchus,
?Parringtonia, ?Dyoplax.
Family Scleromochlidae: Scleromochlus, ?Lagerpeton.
Pseudosuchia, presumably representing a number of distinct families:
Lagosii ch us, Hesperosu chus, Lew isuchus, Sal toposuch us,
Strigosuchus, Dibo throsuchus. Teleocrater, Erpetosuch u s
(Herpetostich u s ) , Triassolestes *.
I also consider Rauisuchus. Ticinosuchus and Hesperosuchus as probable
pseudosuchians.
Yet another alignment of thecodontians is that of Charig, Krebs, Sus &
Westphal (in press), which I have adopted here (see Appendix).
Despite these uncertainties in classification, there is widespread agreement
that “pseudosuchian type” thecodontians are more primitive in almost all
characters than are all other non-thecodontian archosaurs and birds. And,
although Krebs (1963, 1965, 1974) does not agree, it is generally held that as a
group, they probably did give rise to all non-thecodont archosaurs and birds.
But the question that still remains: Is there any reason to believe that
Archaeopteryx arose directly from a pseudosuchian rather than from a
coelurosaur? In the absence of any contrary evidence from coelurosaurs, we
must determine if there is any concrete evidence in presently known

* Charig & Reig ( 1 970) considered Saurosuchus, Prestosuchus, Mandasuchus and Ticinosuchus to be
pseudosuchians. Krebs (1965) had originally assigned Ticinosuchus to the Pseudosuchia (family
Rauisuchidae) and in 1973 offered new tarsal evidence for a pseudosuchian placement of Rauisuchus.
Although Romer ( 1 9 7 2 ~ )placed Ticinosuchus (and the other taxa above) in the Proterosuchia
(Prestosuchidae), Ticinosuchus is included in the following discussion of pseudosuchian evidence.
 ARCHAEOPTERYX AND THE ORIGIN O F BIRDS 143

pseudosuchians t o substantiate a closer relationship between birds and

pseudosuchians than between birds and coelurosaurian theropods.

Manus and forelimb

The pseudosuchian manus and forelimb are primitive and unspecialized in
virtually every respect. Accordingly, it is conceivable that these structures
could have been modified t o the specialized condition of Archaeopteryx, just
as they are supposed to have given rise to the equally specialized manus and
forelimb of coelurosaurs. I t must be emphasized, however, that there is nothing
about the manus or forelimb, insofar as they are presently known in any
pseudosuchian, that even remotely resembles features in either Archueopteryx
or any theropod. Independent derivation of two such strikingly similar
appendages from a dissimilar, albeit primitive, ancestor is at least twice as
improbable as a common derivation from a single lineage.
Manus. The pseudosuchian manus in all cases appears to have consisted of no
less than four digits, and probably five. In Euparkeria, the hand is incompletely
known, but Ewer (1965) reported parts of four metacarpals in two specimens.
Similarly, a complete manus is not known in existing specimens of
Ornithosuchus. Walker (1964), however, identified metacarpals I to IV and
presumed that a fifth was present. Colbert (1952) reported fragments of five
metacarpals in Hesperosuclzus and Krebs (1965) reconstructed five complete
digits in Ticinosuchus from disarticulated elements of both hands. As a
“typical” pseudosuchian, Ticinosuchus seems to provide the best available
evidence on the pseudosuchian manus. As pieced together by Krebs, the manus
consisted of five complete digits (Fig. 24) with a primitive formula (2-3-4-5-3).
The fingers are relatively short, apparently only a little longer than the
respective metacarpals. The proximal phalanx is the longest in each finger and
digit 111 is the longest finger.
This structure differs significantly from that of Archaeopteryx and most
coelurosaurs. Conversion into either would require the loss of digits IV and V,
great elongation of all remaining digits relative to their metacarpals, and
elongation of all penultimate phalanges relative to the proximal elements. While
digit V may have been reduced or lost in Euparkeria and Ornithosuchus and
perhaps other genera as well, there is n o certain evidence of this, nor is there
evidence of reduction of digit IV in any pseudosuchian.
Metacarpus. As noted above, five metacarpals are present in Ticinosuchus
and ffesperosuclius. Only four are known in Euparkeria and Ornithosuchus,
although Walker (1964) supposed a fifth t o be present. Bonaparte (1971b)
identified five metacarpals in Riojasuchus, although the second to fifth were all
incomplete. He described the first two as the more robust. In Euparkeria,
metacarpal I appears to be the shortest, suggestive of the conditions in
Archaeopteryx and many theropods, whereas V is the shortest in Ticinosuchus
while IV (V is not known) is the shortest in Ornithosuchus. Metacarpal 111 (not
11, as in Archaeopteryx and theropods) apparently is always the longest. None
of the five metacarpals is complete in the only specimen of Hesperosuctius, but
their slender construction, even of metacarpal I, suggests a greater relative
length for the metacarpus in this genus.
Available evidence indicates a short, primitive construction of the
pseudosuchian metacarpus, with five elements present in some and possible
144 J. H. OSTKOM

Figure 24. Comparison of some thecodontian hands with that of Archaeopteryx. A ,

Ticinosuchus (after Krehs, 1965); B , the aetosaur Stugonokpis (after Walker, 1961 ); C.
Archaeopteryx (Berlin specimen); D, Riojasuchus (after Bonaparte, 1971 b). All are right hands,
but not drawn to the same length. Vertical scales = 3 cm. Contrast the pronounced differences
from Archaeopteryx with the strong similarities illustrated in Fig. 9.

reduction to four in Euparkeria and perhaps Ornithosuchus. Ticinosuchus with

metacarpal V the shortest, coupled with Ornithositchits where IV is the shortest
and no evidence of a fifth exists, might be considered as indicative of a trend
among some pseudosuchians toward the three-fingered manus of
Archaeopteryx. However, Ornithosuchtts has an elongated metacarpal I,
whereas in Euparkeria the first is very short and theropod-like, but there is no
evidence that metacarpal IV is undergoing reduction. The metacarpus of
Archaeopteryx obviously could have been derived from any of these, but
structural similarities between the metacarpi of Archaeopteryx and the
Pseudosuchia are not obvious.
Carpus. The few carpals that are known in pseudosuchians are, for the most
part, rather indistinctive ossicles of uncertain identity. N o carpals are preserved
 ARCHAEOPTERYX AND THE ORIGIN OF BIRDS 145

in Euparkeria, but because of the space between the metacarpals and the
epipodials, Ewer (1965) supposed them t o have been cartilaginous. Von Huene
(1914) recognized three carpals in Ornithosuchus, and, more recently, Walker
(1964) identified a fourth. Walker was not able t o identify which carpals were
represented. Krebs (1965) found only two elements in Ticiriosirchus which he
labelled the radiale and ulnare. Bonaparte (1971b) reported three large and
massive carpals (radiale, ulnare and an intermedium) in Riojasuchus.
None of these various patterns shows any particular resemblance to the
carpus of Archaeopteryx (see Fig. 24). In that light, there is nothing to be
gained from theorizing about transitional stages from one to the other. The
Archaeopteryx pattern could have been derived from one of these, but at the
moment there is n o evidence of that.
Radius and ulna. The epipodials of pseudosuchians are not particularly
distinctive, being more or less straight and cylindrical and of varying
robustness. Those of Hesperosuchus are long and quite slender, while those of
Riojasuclrus and Ticinosuchus are shorter and more massive. A moderate to
prominent olecranon on the ulna is present in all. In Hesperosuchus and
Ornirhosirchus, the radius and ulna appear to have been positioned close
together, while in Ticinosuchu~ and Riojasuchus there was a significant
intermembral space. Only in Hesperosuchus does the epipodial form approach
the very long and slender form of the radius and ulna in Archaeopteryx.
Humerus. The pseudosuchian humerus is considerably more robust and
much shorter relative to the dimensions of the animal as a whole, than in either
Archaeopteryx or coelurosaurs. The shaft is essentially straight but flares
prominently at both ends and thus has a narrow-waisted shaft profile. The
deltopectoral crest projects prominently from ths shaft, but is rather short and
limited to the proximal quarter of humeral length. Considerable variation exists
in the relative massiveness of the pseudosuchian humerus (Fig. 25). That of
Hesperosuchus is very slight, even delicate, as compared with that of
Ticinosuchus or Riojasuchus. Apart from being straighter and more robust, the
most conspicuous difference between typical pseudosuchian humeri and that of

A B C D E F

Figure 25. Comparison o f various thecodontian humeri with that of Archaeopteryx. A , The
aetosaur Srugonolepis (after Walker, 1961); B, Riojasuchus (after Bonaparte, 1971h); C,
Eupurkeria (after Ewer, 1965); D, Ticinosuchus (after Krehs, 1965); E. Hesperosuchus (after
Colbert, 1952); F, Archaeopteryx (Berlin specimen). All are left humeri viewed in ventral
aspect and drawn to unit length. Vertical scales = 1 0 cm. Compare these humeri with those of
Fig. 1 1 .
146 J. H. OSTROM

Archaeopteryx is the much shorter length of the shaft in pseudosuchians distal

to the deltopectoral crest and the more massive distal extremity.
Forelimb summation. The pseudosuchian forelimb is relatively much shorter
and more massive than that of Archaeopteryx or coelurosaurs (Fig. 26), and
the hand constitutes only a small fraction of forelimb length. Total forelimb
length ranges from about 40% to 50% of presacral vertebral length in most
pseudosuchians and reaches a maximum of about 60% in Hesperosuchus.
compared with about 7 5% in Ornitholestes and 120% in Archaeopteryx.
Considerable variation exists among pseudosuchians in the ratio of humerus to
epipodial lengths, with the humerus length exceeding that of the radius by
about 10% in Euparkeria and Hesperosuchus, and by 20% in Ornithosuchus,
whereas the radius is 5% longer than the humerus in Ticinosuchus and the two
elements are equal in Scleromochlus. While the ratio in Eutiarkeria and

Figure 26. The right forelimb skeleton of Archaeopteryx (A) contrasted with that of several
thecodontians; Ornithosuchus (B); Ticinosuchus (C);and Stagonolepis (D).For convenient
comparison, all humeri'are drawn to the same length to show differences in component
proportions and robustness. Vertical scales = 5 cm. Compare with Fig. 12.
 ARCHAEOPTERYX AND THE ORIGIN OF BIRDS 147

Hesperosuchus approximates that of Archaeopteryx. the proportionately much

shorter total limb length is in strong contrast. Likewise, the short massive hand
is completely different in design and proportions from that of Archueoptcr.v.w.
In summary, the forelimb and hand as known in pseudosuchians are
sufficiently primitive in all features for both the Archaeopteryx and
coelurosaurian conditions to have been derived from them, but there are no
strong similarities to either of the latter.

Pectoral girdle
The scapulo-coracoid, known in a variety of pseudosuchians, appears to have
been fairly uniform in design and bears only superficial resemblance to that of
Archaeopteryx and theropods. In some specimens, a well-ossified clavicle and
interclavicle have been noted.
Scapula. The scapula consists of a narrow to moderately wide blade which
flares to a variable extent at its posterior extremity (Fig. 27). Proximally there
is a very large and long acromial expansion projecting away from the glenoid.
The blade appears t o have been oriented at nearly right angles to the vertebral
column or inclined slightly backward, instead of nearly paralleling the vertebral

Figure 27. Outline drawings of the left scapulo-coracoids of various thecodontians compared
with that of Archaeopteryx ( G ) . All figures are viewed normal to the scapula with
corresponding distortion of the coracoid outlines. Notice the broad, flaring form of the
posterior extremity which is typical of the thecotlontian scapula. A, The aetosaur Stagonolepis
(after Walker, 1961); B. Ticinosuchus (after Krehs. 1965); C, Riojasuchus (after Bonaparte,
1971b); D. Hesperosuchus (after Colbert, 1952); E. Euparkeria (after Ewer, 1965); F.
Ornithmuchus (after Walker, 1964); G, Archaeopteryx (reconstructed from the Berlin and
London specimens). Scapulae are all drawn t o the same length. Vertical scales = 2 cm. Contrast
these data with those illustrated in Fig. 13.
148 J. H. OSTROM

column as seems to have been the case in Archaeopter.y.x (see, for example, the
Berlin specimen). The glenoid region is robust and lies well behind the blade
axis.
Coracoid. The coracoid of pseudosuchians is crescentic in shape with a
strongly curved antero-ventral margin. The coracoid part of the glenoid is much
larger than the scapular part and extends well behind it. Below that is a long
posterior extension of the venrral coracoid edge which reaches far behind the
glenoid. A faint to strong ridge is present in Ornithosuchus, extending from the
glenoid rim down and back to the end of this posterior extremity. Some part of
this ridge may correspond to the “biceps tubercle” of Archaeopteryx and
coelurosaurs, but its form and location are significantly different. Perhaps a
better comparison can be made with what appears to be an external swelling
between the glenoid and the supracoracoid foramen in Euparkeriu, as figured
by Ewer (1965, fig. 9).
Clavicle and in terclavicle. These dermal elements of the pectoral girdle have
been identified in several pseudosuchians, namely in specimens of Euparkcria,
O m ithosu ch us, Sal toposuch us, Ticinosu ch us and perhaps more. The clavicle
appears to have been a thin, curved rod-like element adjacent to the anterior
margin of the coracoid and the acromial border of the scapula. As
reconstructed by Ewer ( 1 9 6 9 , the interclavicle in Euparkeria is a thin spatulate
bone that was situated between the medial or inferior borders of the two
coracoids. No interclavicle has been recognized as such in Archaeopreryx, but
Osborn (1924) recognized an interclavicle in Oviraptor, as did Camp (1936) in
Segisaurus. Clavicles have also been identified in both of these genera and may
also be present in a specimen of Vdociruptor, as was noted above. There is no
obvious similarity between known pseudosuchian clavicles and the furcula
preserved in two of the Archaeopteryx specimens.
Sternum. As noted earlier, the sternum is not known in any pseudosuchian.
We may safely assume that it was present in a cartilaginous state, since it
occurs, usually in an ossified state, in all other major archosaurian groups,
including birds, of course, although apparently not in Archaeopteryx or most
theropods.
Hitidlimb and pes
The pseudosuchian pes and hindlimb are much better known than the
forelimbs, with well preserved remains available in Euparkeria. Ornithosuchus,
Gracilisuchus, R iojasuch us, Sclero mo ch lus, Hesperosu ch us and Ticinosu ch us,
plus the potentially very important new South American forms Lagosuchus and
Lagerpeton. For the most part, the rear appendage is primitive for an
archosaur, although clearly advanced over the lepidosaurian condition, and is
potentially a good structural precursor to the condition in Archaeopteryx and
that of all theropods. There are some difficulties, however, concerning the
evolutionary significance of the typical pseudosuchian tarsus. These difficulties
have been most clearly elucidated by Krebs (1963, 1965, 1974). The recent
discoveries of the South American genera Lagosuchus and Lagerpeton (Romer,
1971, 1972b) may provide important evidence for resolving this problem.
Pes. Where adequately known, the pseudosuchian foot is composed of at
least four and usually five digits with a primitive phalangeal formula of
2-3-4-5-(3), (see Fig. 28). Digit V is present in Euparkeria, Ornithosuchus,
 ARCHAEOPTERYX AND T H E ORIGIN O F RlKDS 149

Hesperosuchus, Riojasuchus and Ticinosuchus, but is reduced to a splint-like

vestige of the metatarsal in Lagosuchus, Gracilisuchus, Lugcrpeton and
apparently (Woodward, 1907) in Sclermochlzts. In Riojamclius. the fifth toe
may have been reduced to two phalanges. Except for Lagcrpeton. where the
fourth toe is the longest, and 111, I1 and 1 are successively shorter, the third toe
is the longest in all known pseudosuchians. The fifth (when present) and first
digits are the shortest and subequal in length, and the second is generally
slightly shorter than the fourth. In all instances, the proximal phalanx is the
longest in each toe. The foot is relatively short and broad in Euparkeria and
Riojusuchus, as compared with those of Ornithosrichus and Hcsperosiichus. By
contrast, those of Lagerpcton and Lagosuchus are surprisingly long and slender.
The reduced feet of Lagosuchus and Gracilisirchus are of special interest
because with the loss or reduction of the fifth toe, together with the slender
and nearly symmetrical construction, they approach the condition in
Archaeopteryx and various coelurosaurs much more closely than does any
other pseudosuchian. The unreversed hallux is the most obvious difference
from Archaeopteryx.
Metatarsus. Five metatarsals are present in all adequately known
pseudosuchians (Fig. 28) but the fifth is reduced to a splint-like vestige in
Lagosuchus, Gracilisirchus and Lagcv-peton and perhaps in Scleromochlus.
Eiiparkeria and Riojasuchus are judged to have the most primitive metatarsi in
that these are relatively short, none of the elements seems to be reduced and
the fifth metatarsal is distinctly hook-shaped, as in lepidosaurs. The metatarsus
of Ticirzosuchus is similar. In Etiparkeria the metatarsals are quite short,
approximating the length of the corresponding toe*. The metatarsals are longer
than the digits in Hesperosuchus, and very much longer in Lagosuchus,
Lagerpeton and perhaps Gracilisuchus where the entire foot including the
metatarsus is very long and narrow. Where the fifth metatarsal is not reduced,
the metatarsus is nearly symmetrical, with 111 the longest, I1 and IV of shorter
subequal lengths and I and V the shortest and also subequal. Exceptions to this
form are found in Riojasuchus, where the fourth metatarsal is about the same
length as the third, and in the incompletely known Lewisuchus where the
second and third metatarsals are of equal length. Lagerpeton has the most
specialized design, the metatarsus being asymmetrical with metatarsal IV the
longest and 111, I1 and I successively shorter.
The reduced metatarsus with a splint-like vestige of the fifth is reminiscent
of coelurosaurian metatarsi, like those of Coelophysis, Compsognathus,
Ornitholestes and others. The metatarsus of the Eichstatt specimen of
Archaeopteryx is also similar in this respect. But the reduced metatarsi of the
new South American forms, and of pseudosuchians in general, are distinct from
that of Archaeopteryx and all theropods in that the first metatarsal is never
reduced (in fact it is very long in every pseudosuchian except Lagerpeton), nor
is it positioned behind metatarsal 11. The avian and theropod metatarsi are
easily derivable from the generalized pseudosuchian condition, but, except for
reduction of the fifth toe in some, there is no obvious trend toward this
condition in known pseudosuchian material.
Charig (1972) has pointed out that, in most instances, greater relative lengths of metatarsals
compared with digit lengths is probably a consequence of phalangeal shortening rather than of metatarsal
elongation.
150 J. H. OSTROM

D E
‘A

‘i
Figure 28. Outline drawings of the right pes. metatarsus and tarsus of various pseudosuchians
compared with those of Archaeopteryx (F) and the aetosaur Stagonolepis. All views are dorsal
(anterior) and are to the same length to eliminate size differences and facilitate comparison. A,
Stagonolepis (after Walker, 1961); B, Ticinosuchus (after Krehs, 1965); C. Euparkeria (after
Ewer, 196.5); D, Lagerpeton (after Romer, 1971); E. Lagosuchus (after Romer, 1971); F,
Archaeopteryx. Vertical scales = 2 cm. Compare with Fig. 1 5 .

Tarsus. The pseudosuchian tarsus, where adequately known, consists of two

large proximal elements, the astragalus and calcaneum, and two smaller distal
tarsals that are usually identified as the third and fourth. The distal elements
appear to have been positioned at the proximal ends of the third and fourth
(and sometimes the fifth) metatarsals. A slightly different arrangement
apparently existed in Gracilisuchus in which the calcaneum articulated directly
with the fourth and fifth metatarsals and the two distal tarsals occupied
positions between the astragalus and the first three metatarsals, according to
Romer’s ( 1972a) reconstruction. In nearly all pseudosuchians, the calcaneum
bears a prominent “heel” or backwardly projecting tuber, which like that of
crocodilians probably provided leverage for extensor muscles of the foot.
Figure 29 compares the proximal tarsals of Ticinosuchus with those of
Deinonychus and Archaeopteryx. Notice that there is n o calcaneal tuber in the
last two and no ascending process of the astragalus in the first.
As shown, the typical pseudosuchian tarsus differs markedly from that in
 ARCHAEOPTERYX AND T H E ORIGIN OF BIRDS 151

A B

as as

D E
.-. --.

ca

Figure 29. Pseudosuchian proximal tarsals ( A and B) contrasted with those of A rchaeoptervx
(C) and a small theropod dinosaur ( D and E). A and B, anterior and posterior views,
respectively, of the astragalus and calcaneum of Ticinosuchus, as reconstructed by Krebs
(1965). C , Anterior view of the proximal tarsals as reconstructed by the author from the Berlin.
London and Eichstitt specimens of Archaeopteryx (see Fig. 17). D and E, anterior and
posterior views. respectively, of the same proximal tarsals in Deinonycltus untirrliopus (Y.P.M.
5226). Of special importance is the prominent dorsal o r ascending process of the astragalus in
Archaeopteryx and theropods, a blade-like dorsal flange that overlaps the lower anterior surface
of the tibia. Also important is the prominent posterior tuber of the pseudosuchian calcaneum, a
feature that is unknown in theropods and Archaeopteryx. Vertical scales in C = 2 mrn; in A , B.
D and E = 2 cm. as, Astragalus; ca, calcaneum; tu, calcaneal tuber.

Archaeopteryx and theropods, in which the tarsus forms a simple roller-bearing

or hinge-like mesotarsal joint between the foot and the crus, with the two
proximal tarsals firmly joined t o the crus and the distal tarsals united with the
metatarsals. As Charig (1972) and others have noted, the mesotarsal joint is
correlated with digitigrade posture, upright parasagittal limb orientation and
sometimes with bipedality. The typical condition in pseudosuchians is a
crurotarsal joint in which the calcaneum is structurally part of the foot and the
astragalus is part of the crus, as in modern crocodilians. This organization is
associated with a plantigrade condition and semi-erect or sprawling posture.
Contrary to the mesotarsal joint where nearly all of the movement takes place
152 J. 11. OSTKOM

between the distal tarsals on one side and the astragalus and calcaneum on the
other, in the pseudosuchian tarsus the principal movement takes place at a
complex ball-andsocket articulation between the astragalus and calcaneum.
The function of the calcaneal tuber is clearly to provide increased leverage for
extension (backward thrust) of the plantigrade foot. In the mesotarsal ankle,
this thrust is provided to the digitigrade foot through the digital flexors, hence
no calcaneal tuber is necessary.
Krebs (1963, 1965, 1974) has argued persuasively that the mesotarsal
condition cannot have been derived from the crocodiloid or crurotarsal
condition typical of pseudosuchians. His principal argument: it is difficult to
visualize the transfer of the calcaneum from the pes to the crus without
disrupting the primary ankle functions of flexion and extension. Charig (1972)
acknowledged the difficulties cited by Krebs, but he did not consider them
insurmountable. He postulated that the mesotarsal joint might have evolved by
way of a small, unknown pseudosuchian in which “because of its lightness, the
development of a complex ‘crocodiloid’ tarsus with a massive calcaneal tuber
was unnecessary”.
I t now appears that Charig may have been correct and that Krebs’ arguments
are purely academic. Several specimens of two new Argentine pseudosuchians,
Lagerpetmi and Lagosuchus (Romer, 1971, 1972b), both small, have what
appear to be fully developed mesotarsal joints. In both, the astragalus and
calcaneum are united and closely applied to the tibia and fibula. Two distal
tarsals are present, capping metatarsals 111 and IV in Lagerpeton and I to IV in
Lagosuchus. Of special interest in Lagosuchus is a small bony process that
projects backward from the fibular region of the astragalo-calcaneum. If this
structure does not represent a vestige of the calcaneal tuber, it certainly is
remarkably coincidental in its position and construction. N o such feature is
preserved in Lagerpetorz. In Lagerpeton, an ascending process of the astragalus
occurs on the flexor surface of the tibia, rather than on the extensor or anterior
surface.
Neither of these genera can be directly ancestral to later dinosaurs or birds
because of the specializations of the astragalus in Lagerpeton and the peculiar
form of the ilium in Lagosuchus, but they d o indicate that some
pseudosuchians possessed a mesotarsal ankle joint, thus obviating the problem
raised by Krebs of converting the much more common pseudosuchian
crurotarsal joint. The tarsus of Lagosuclzus is even more interesting for the
question at issue here, because, as was noted above, the pes and metatarsus
also approach the pattern of Archaeopteryx and coelurosaurs much more
closely than do those of any other known pseudosuchian. As Fig. 28 shows, the
toes and metatarsals are long and slender and the relative lengths of all are
similar to those of Archaeopteryx. The most significant difference is the
unreversed hallux in Lagosuchus.
Tibia and fibula. These elements provide little evidence for resolving the
question of the origin of Archaeopteryx one way or the other. Proportions and
relative lengths differ slightly among pseudosuchians, but the morphology
typically is that of nearly straight, cylindrical bones with expanded extremities.
A cnemial crest is present in some but is never prominent (ridge would be a more
appropriate term). In Riojasuchus and Ticinosuchus there is a prominent
anterior expansion near mid-shaft on the fibula. Although incompletely
 A R C H A E O P T E R Y X AND THE ORIGIN OF BIRDS 153

preserved, the fibulae of Eupurkeria and Ornitliosuchus d o not seem t o possess

a similar feature, nor has it been recognized in other pseudosuchians that I am
aware of. In the new South American forms Lagosirclurs, Lagerpeton and
Gracilisucliirs the tibia and fibula are long and slender. In 1:’irparkcriu.
Orriitliosirchus, Ticiriosirclitts and Kiojusirchtrs they are relatively shorter and
more massive. The fibula is never splinter-like as in Arcliaeo~,trri)x.
Fmziir. The pseudosuchian femur is distinctly crocodilian in form, slightly
sigmoidal with a stout, compressed cylindrical shaft (Fig. 30). I t is distinctly
non-avian, unlike the femur of Archacwpter?’s. The proximal end bends
medially and anteriorly, the head being only slightly offset from the shaft with
no distinct neck. The head tends to be elliptical rather than hemispherical.
There is no sign whatever of a “greater trochanter”, but a conspicuous external
protuberance occurs slightly below the level of the femoral head in
Ornitiiosiichirs and Riojasticlius. This was labelled the “lesser trochanter” by
Walker (1964) and Bonaparte (1971b). A much less prominent swelling or ridge
is present in Hespcwmcliiis and Eupurkeria. and perhaps also in Lagerpeton,
according to Romer’s (1972b) restoration. A probable fourth trochanter is also
present in Riojasirchus. where it is very prominent, and in Ortiitliosrrcliir.~.As
Fig. 30 illustrates, there is very little resemblance between pseudosuchian
femora and that of Arch aeop tcry x.
Hitidlimb sirmmation. Individually, the hindlimb components in classical
pseudosu ch ians (e.g., Eupurkeria, O m ithosir cti us, Hesperosucli 11s) bear little
resemblance to those of Arcliacwptcrj3.u.The femur is distinctly crocodilian-like
and longer than the tibia, and the distal elements are relatively shorter and
much more massive. Taken as a whole, the hindlimb proportions also differ
greatly from Archaeopteryx (see Fig. 31). The new South American forms
Lugosiccliirs and Lugerpeton, and also Scleromochliis, however, do show
tendencies toward the conditions in Archueopteryx. The femur is straighter and

A B C D E

Figure 30. Comparison of right femora in lateral aspect, of various pseudosuchians with that of
Archaeopteryx (E). Notice that, in addition to being more slender, the femur of Archaeopteryx
displays a simple fore-aft curvature, in contrast to the sigmoidal curvature of pseudosuchian
femora. A , Lugerpeton (after Romer, 1971); B. Hesperosuchus (after Colbert. 1952); C ,
Ticinosuchus (after Krebs, 1965); D, Ornithosuchus (after Walker, 1964); E, Archaeopteryx
(see Fig. 18). All elements are drawn to the same length; relative scales are indicated by the
vertical scales which equal 2 cm.
154 J. H. OSTROM

B D F

I I
c

Figure 3 1 . Outline drawings of the left hindlimb skeleton of Archaeopteryx ( E ) contrasted with
those of various pseudosuchians (A-D) and that of a small theropod ( F ) . All figures are views of
the anterior aspect with femora drawn to the same length in order to minimize differences due
to size. A. Euparkeria; B, Riojasuchus; C, Ticinosuchus; D, Lagerpeton; E, Archaeopteryx; F,
Compsognathtrs. Vertical scales = 2 cm.

more slender, the fifth digit is reduced and the epipodials, metatarsus and foot
are elongated and slender. Furthermore, the hindlimb proportions of these
genera are quite close to those of the several specimens of Archaeopteryx, with
the tibia approximately 20% to 30% longer than the femur, compared with
30% to 40% in Archaeopteryx, and the metatarsus constituting a similar
fraction of total hindlimb length.

Pelvic girdle
Pelvic form seems to have been relatively conservative within the
Pseudosuchia. All possess comparatively low and short ilia with very short
anterior processes and a posterior process of long or only moderate length
(Fig. 32). The acetabulum is large and closed (except for an incipient
perforation in Ornithosuchus) and a long vertical suture joins the pubis and
ischium beneath the acetabulum. A robust ischium projects almost straight
backward and slightly downward. The pubis, only slightly less robust, projects
antero-ventrally in its proximal part, but then turns sharply downward distally.
This latter feature has been noted by Heilmann (1926) and others as possibly
indicative of an initial stage in the backward rotation of the pubis toward the
avian condition-a possibility that I reject in view of the total dissimilarity
between avian and pseudosuchian pubes. There is little resemblance between
 ARCHAEOPTERYX AND THE ORIGIN OF BIRDS 155

Figure 32. Thecodontian pelves compared with that of Archaeopteryx (F). All figures are of
left pelves in lateral view, but not to the same scale. A. Erythrosuchus (a proterosuchian); B,
Stagonolepis (an aetosaur); C . Ticinosuchus (a pseudosuchian) D, Euparkeria (a
pseudosuchian); E, Ornithasuchus (a pseudosuchian); F. Archaeopteryx (as reconstructed by
the author from all five skeletal specimens).

the pelvis (as a whole) of any known pseudosuchian and that of

Archaeopteryx.
Pubis. The pseudosuchian pubis is distinctly primitive in its morphology, as
is well illustrated by those preserved in Euparkeria, Ornithosuchus and
Ticinosuchus. From an extensive dorsal union with the ilium, a robust
longitudinal and nearly horizontal bony plate of the pubis curves downward
beneath and medial to the acetabulum into a broad transverse and nearly
vertical plate distally. There is no mid-length constriction into a long slender
cylindrical shaft as in coelurosaurs and Archaeopteryx. A long symphysis joins
the two pubic plates in front of and beneath the sacrum into a wide transverse
apron which extends up and backward to the inferior junction with the ischia.
In Euparkeria two distinct foramina are present below the acetabulum-an
obturator foramen below and a smaller thyroid foramen above. Only the
obturator foramen can be recognized in Ornithosuchus. Neither of these occurs
in Archaeopteryx, the obturator foramen being represented by an open notch
as in most theropods and birds. The broad transverse pubic apron is also
markedly different from the narrow longitudinal distal expansion of the pubes
in theropods and Archaeopteryx.
Ilium Most pseudosuchian ilia are constructed on a similar plan, being
relatively low and short vertical blades with nearly straight or gently convex
upper margins. The anterior process is very short or almost non-existent (see,
for example, Euparkeria, Fig. 32D), and rarely extends much in front of the
acetabulum (Ornithosuchus, Fig. 32E). The posterior blade, on the other hand,
is a moderate to lengthy extension behind the acetabulum (see Ticinosuchus,
Fig. 32C). In a general way, these relative proportions resemble those in a
I 1
156 J. H. OSTROM

variety of theropods, except that there the ilium is usually much deeper. This
contrasts sharply with the proportions in Archaeopteryx where the anterior
process is the longer. Another similarity between pseudosuchian and theropod
ilia is the sharply defined notch between the pubic peduncle and the anterior
process, immediately above the anterior rim of the acetabulum. No such notch
exists in any of the specimens of Arcliaeopteryx. Compared with
Arciiaeopteryx, the pseudosuchian acetabulum is enormous relative to the
ilium; the greater relative iliac length in Arcliaeopterjjx suggests much greater
leverage for the ilio-femoralis muscles along the longitudinal axis, which may
presumably be correlated with the upright posture and parasagittal limb
excursion. The typical brachyiliac condition of pseudosuchians could have
evolved into the more elongated iliac condition of birds, but there is no
indication of such a tendency among known specimens.
Unfortunately the pelves of Lagerpeton and Lagosirclius, which might be
expected to approach the avian condition more closely than those of other
pseudosuchians, are incompletely known. The ilium of each, however, is known
from two specimens. That of Lagerpeton, as figured by Romer (1972b),
appears to have been short and of the general pseudosuchian configuration,
with the posterior process much longer than the anterior process. The ilium of
Lagosircliirs is quite a different matter, but nevertheless it does not seem to
strengthen the evidence for close pseudosuchian-avian relationships. I t is of
peculiar form, reminiscent of a “problematical” bone reported by Colbert
(1952) in Hesperosirchtrs; it consists of two vertical blades, separated by a
broad trough, which join to form a hook-like posterior process. The “anterior”
process is a very short blunt nubbin that projects anterolaterally. So despite the
somewhat Archaeopfer~izc-likeconstruction of the hindlimb of Lagosir ciiiis, the
ilium of the latter has no resemblance whatever to that of Arcliaeopteryx or of
any known theropod.
ischiuni. The paired ischia of Orriitliosiichus and Euparkeria appear to have
been united over much of their length by a long sagittal symphysis. Thus the
pubo-ischiadic plate was primitively continuous, except that instead of being
nearly flat and horizontal, it bowed upward in the region of the pubo-ischiadic
suture. I t is not certain, but apparently the ischia of Archaeopteryx were not
joined medially, or at the very most had contact only near their extremities.
From the robust lower margin of the acetabulum, the pseudosuchian ischium
extends back and slightly downward as a broad, nearly horizontal transverse
plate. Newton (1894) observed an obturator process on the proximal inferior
margin in Ornitliostrcizus, but no comparable feature has been reported in other
pseudosuchian genera, as far as I know. An enlarged obturator process is
present in many theropods, but appears to have been absent in Arcliaeopfer.vx,
although the unique form and short length of its ischium may obscure
identification of the obturator process.
Clearly, there is no obvious similarity between the ischium of Arcliaeopteryx
and that of any pseudosuchian, where it is known. Unfortunately the ischium is
not known in the new South American genera.
Skull and jaws
Good cranial and mandibular materials exist for a number of pseudosuchian
genera, such as Euparkeria, Orriithosuchus, Riojasuchus, Venaticosuchus and
 ARCHAEOPTERYX AND T H E ORIGIN OF BIRDS 157

Gracilisuchus. Detailed studies of these specimens by Ewer (1965), Walker

(1964) Bonaparte (1971b) and Romer (1972a) provide a good picture of the
general nature of the pseudosuchian skull and jaws-a picture that perhaps is
better than we may ever have for Archaeopteryx even with the remarkable
Eichstatt specimen. As several authors have noted before, the pseudosuchian
skull is primitively archosaurian, featuring a number of characters that are
retained in later, more advanced archosaurs: the large antorbital fenestra,
sclerotic ring, large naris bounded by the premaxilla and nasal, and a diapsid
condition. All but the last are certainly present in Archaeopteryx. which may
have possessed the diapsid condition also. As Fig. 3 3 shows, however, the
general configuration of the pseudosuchian skull and jaws is quite different
from that of Archaeopteryx, with a number of cranial and mandibular features
that are not found in the latter. Some of these features are:
(a) There is a single antorbital fenestra, but no sign of smaller accessory
fenestrae at the front of the antorbital fossa.
(b) The orbit tends to be triangular and elevated, with both the preorbital
and postorbital bars converging to a junction beneath the orbit. In both
Ornithosuchus and Riojasuchus the lower temporal fenestra and the antorbital
fenestra extend beneath the orbit.

Figure 3 3 . Pseudosuchian skulls (restored) compared with that of Archaeopteryx (C). also
restored. All skulls are drawn to the same length, with relative sizes indicated by horizontal
scales which equal 1 0 mm. A. Ornithosuchus (after Walker, 1964); B, Riojusuchus (after
Bonaparte, 1971b); C, Archaeopteryx (after Wellnhofer, 1974); D, Euparkeriu (after Ewer,
1965); E. Gracilisuchus (after Ronier, 1972a).
 158 J. H. OSTROM

(c) The snout is short and the premaxillary portion tends to bend downward
in front of the mandibles. This is particularly evident in short-jawed genera such
as Ornithosu chus, Riojasuch us and Venaticosu ch US.
(d) The dentition is heterodont, consisting of relatively few teeth which are
unusually large, often tusk-like, compared to the size of the skull. A distinct
diastema is commonly present at the maxillary-premaxillary suture,
corresponding to the position of a very large lower tusk near the extremity of
the lower jaw.
'
(e) The suspensorium slopes (descends) backward and the articulation is
situated farther back than the occipital condyle. The quadrate is over-lapped
superficially by a large quadratojugal and squamosal which project forward
into the lower temporal fenestra, greatly constricting that opening in
Gracilisuchus and Ornithosuchus. A deep posterior notch (the otic notch?) is
formed by a posterior extension of the upper part of the squamosal.
(f) The mandible is short and deep, especially in those taxa with an
overhanging premaxilla. A large, long external mandibular fenestra is present in
all presently known pseudosuchians.
It cannot be stated that any of these features would exclude pseudosuchians
from the ancestry of Archaeopteryx, but neither d o they weigh in favour of
close relationships.
Vertebral column
Complete vertebral series are not known in many pseudosuchians, but the
typical column seems to consist of from 21 to 25 presacrals, two (rarely up to
fbbr) sacrals and a long caudal series of up to 5 5 segments, according to the
data reported for the genera listed below. In most specimens there is no clear
distinction between the cervical region and the dorsals, the division by various
authors often being arbitrary or based on the assumption of seven or nine
cervical vertebrae. The neck is never long, constituting only 30%or less of the
presacral length. Unlike Archaeopteryx and many coelurosaurs, there is no
indication of any high degree of flexibility in the neck region. All vertebrae are
amphicoelous, relatively short, and of simple construction without pleurocoels.
Vertebral lengths and neural spine heights are nearly uniform throughout much
of the precaudal series.
The most pronounced difference between the pseudosuchian vertebral
column and that of Archaeopteryx is the lesser degree of differentiation in the
presacral series and the shorter, apparently inflexible neck region in
pseudosuchians. A summary of pseudosuchian vertebral counts is as follows:

Cervicals + Dorsals Sacrals Caudals

Euparkeria 22 2 30-40
Ornithosuchus 24 3 25+
Scleromochlus 21 4 40-50
Gracilisuchus 23 2 ?
Ticinosuchus 24 or 25 2 55+
Riojasuchus 24 3 ?

Archaeopte yr 23 5 20-2 3
 ARCHAEOPTERYX AND THE ORIGIN O F BIRDS 159

Other skeletal elements

Gastralia. I t was noted above that gastralia are usually present in
pseudosuchians and perhaps the best example is the type specimen of
Euparkeria (see fig. 20, Ewer, 1965), where they appear to be completely
undisturbed. In this specimen the gastralia consist of numerous closely spaced
chevron-shaped or straight rod-like bones which form a long cuirass extending
from the anteriorly positioned interclavicle all the way back to the pelvic
region. This same close spacing was apparently true of the cuirass in
Ornithosuchus and Scleromochlus. The gastralia in the specimens of
Archaeopteryx have been slightly disarrayed, but the Berlin and Eichstatt
specimens show a similar chevron-like form. However, far fewer elements seem
to be involved, and, as preserved, they are much more widely spaced. Because it
is a primitive condition, the presence of gastralia alone has no significance in
relation to the origin of Archaeopteryx. Also, the significance of the smaller
number of gastralia and their wider spacing in Archaeopteryx is not known,
because the precise original arrangement cannot be established. However, it
does resemble the pattern preserved in some theropods like Stru thiomimus
(A.M.N.H. 5339; see Osborn, 1917: pl. 24) and Allosaurus (U.S.N.M. 4734),
more closely than that of pseudosuchians.
Scutes. Dermal ossifications are characteristic of various thecodontians, but
their presence in pseudosuchians cannot be considered a primitive state. No
dermal scutes are preserved in the most primitive thecodontians, nor in any of
the specimens of Archaeopteryx. Dermal ossifications have been reported in
theropods (Gilmore; 1920), but seem to be minor, or isolated occurrences.
Hence these elements are considered irrelevant to the question at issue, (until
such time as they are detected in a future specimen of Archaeopteryx).
Summary of pseudosuchian evidence
Beyond the generally accepted fact that pseudosuchians are indeed
“primitive enough” to have given rise to birds, as Broom and Heilmann both
observed, there are few reasons for postulating a close evolutionary relationship
between them. There are very few close anatomical resemblances between
Archaeopteryx and any pseudosuchian. In fact, only in one feature does any
pseudosuchian resemble Archaeopteryx more closely than does any theropod,
this being the tibia to femur ratio in Scleromochlus, Lagosuchus and
Lagerpeton, where the tibia is from 20%to 30%longer than the femur. Among
theropods, only in struthiomimids, Compsognathus, Microvenator and
Deinonychus* is the tibia longer than the femur but by only 10%to 15%. In all
other features, the closest resemblance to the morphology preserved in
Archaeopteryx is found in coelurosaurian theropods. On the basis of the degree
of anatomical similarity, Archaeopteryx must be considered much more closely
related to coelurosaurian theropods than to pseudosuchians.

The Sp henosu chus evidence

The question of bird origins has been complicated in recent years by a new
hypothesis put forward by Walker (1972, 1974) in which it is suggested that
* A new specimen of Deinonychus (now under study by the author) provides the only known femora
for this genus. The femur is nearly 10%shorter than the tibia (Ostrom, in press).
160 J . H. OSTROM

birds arose from an unknown Middle or Late Triassic reptilian stock (which he
refers to as “proavians”) which also gave rise to crocodilians. On the surface of
it, Walker’s common-ancestor hypothesis would seem to be only a rephrasing of
the long-held belief that thecodontians gave rise to crocodiles and birds (as well
as dinosaurs and pterosaurs). But his thesis is more complex than that. On the
basis of a single specimen of the South African Late Triassic thecodontian-like
reptile Sphenosuchus, Walker believes that birds and crocodiles shared an
immediate common ancestry above the thecodontian level of organization.
At first glance, Sphenosuchus appears to be a typical pseudosuchian not very
different from Euparkeria (Fig. 34), and various authors (e.g., von Huene,
1925; Broom, 1927; Romer, 1956) have so classified it. However,
Sphenosuchus also has certain crocodilian features, as d o several other Triassic
thecodontian forms (Pedeticosaurus, Erythrochampsa, Notochampsa). These
crocodile-like “thecodonts” have been something of an enigma for many years.
In 1966, Romer removed Sphenosuchus from the Pseudosuchia and placed it
with primitive crocodilians (Protosuchia), a placement that he maintained in
later years ( 1 9 7 2 ~ ) . Subsequently, Walker (1970) proposed his revised
classification of crocodilians in which he allied Sphenosuchus with other
aberrant, crocodile-like genera (Pedeticosaurus, Saltoposuchus and Hespero-
suchus). More recently, Bonaparte (1971b) allied the new Argentine form,
Pseudhesperosuchus, with Sphenosuchus and Hesperosuchus in his Infraorder

B I

Figure 34. Comparison of the restored skulls of Archaeopteryx (A) and Sphenosuchus acutus
(B). The Archaeopteryx restoration is by Wellnhofer (1974), based largely on the Eichstatt
specimen (see Fig. 218). The reconstruction of the skull of Sphenosuchus is by Walker (1972).
based on the only known specimen. Also included is a restoration (by Bonaparte, 1971b) of the
skull of Pseudhesperosuchus jachaleri (C), from the Upper Triassic of Argentina. It is included
here for comparative purposes because both Bonaparte and Walker consider
Pseudhesperosuchus to be a sphenosuchid, belonging t o a family which, according t o Walker,
should b e regarded as primitive crocodylomorphs. Bonaparte. however, places sphenosuchids in
the suborder Pseudosuchia. Horizontal scales = 10 mm.
 ARCHAEOPTERYX AND THE ORIGIN OF BIRDS 161

Sphenosuchia, but he considered these to be pseudosuchians, not

crocodylomorphs. In 1972(c), Romer summarized some of the more important
crocodilian-like features in the skull of Sphenosuchus, citing the forward
inclination of the quadrate, reduced quadratojugal, small antorbital fenestra,
loss of the postfrontal, fusion of the basipterygoid articulation, partial
development of a secondary palate, and, in the incomplete post-cranial
skeleton, the elongated form of the coracoid. There can be no question about
the crocodilian nature or tendency of these conditions, but some of them
(small or absent antorbital fenestra, absence of the postfrontal, and fusion of
the basipterygoid articulation) are widespread among archosaurs and are not
exclusively crocodilian. Of much greater importance here is Walker’s (1972,
1974) claim that other cranial features of Sphenosuchus are also present in
certain modern birds or closely resemble features thereof, and additional
characters of Sphenosuchus preview (anticipate) other conditions of later birds.
Dr Walker was kind enough t o show me what he believes to be evidence for
S~~henosuctius-avian affinities, much of which he has published in the two
papers cited above. In view of his announced intention to publish a
monographic study of Sphenosuchus. I will list here only the more important
details mentioned by Walker in those papers. I t should be kept in mind that
this listing is not necessarily the complete evidence recognized by Walker now,
inasmuch as his studies are still in progress. The major items cited by Walker
are :
(a) An anterior placement of the inner head of the quadrate, a condition
that is present but masked in modern birds because of expansion of the brain
case.
(b) An elongated cochlea of the inner ear, as in birds and crocodilians.
(c) A quadrate with all the essential features of the avian quadrate, including
orbital and pterygoid processes and a curved articular facet for the
quad r atoj ugal .
(d) An indication of streptostyly and kinesis (character “C” above) in the
juvenile (but not the adult) skull of Sphenosuchus, after the pattern of kinesis
and streptostyly in modern birds.
(e) An extensive system of air spaces within the cranium, from which the
pattern of pneumatic cavities in birds and crocodilians could be derived.
(f) A palatal configuration with a low position of the palatine-maxillary
contact and a system of ridges on the palatines close to the mid-line (a preview
of the avian hemipterygoids), all of which provide an ideal pattern from which
to derive the schizognathous bird palate.
(g) A crescentic shape of the occipital surface, with short downwardly and
backwardly directed paroccipital processes projecting well behind the quadrates
to form the posterior walls of the tympanic cavities, as in some modern birds.
(h) Contours of the occipital surface with its pattern of low ridges and
shallow depressions.
(i) A thin, transverse occipital crest with its paired dorso-lateral
culminations, resembling that of some living birds.
(j) A well developed sagittal crest between large upper temporal fenestrae
and a vertical orientation of the transverse (occipital) crest, in close
resemblance to those of the Great Northern Diver (Gavia imrner).
(k) The coracoid elongated like those of modern birds (and crocodilians).
162 J. H. OSTROM

The most important of these, apparently, is the evidence that Walker sees for
bird-like kinesis and streptostyly at some stage immediately preceding
Sphenosticlzzis (or in the juvenile Sphenostichus). For example, in addition to
the above features, Walker postulates the loss of the upper temporal arch and
the postorbital bar during the post-Sphenosuchus evolution of birds. This loss
presumably improved the kinetic mobility of the avian skull. Walker’s
interpretation is based on the upward and forward curvature of the upper
temporal bar in Sphenosuchus, which he believes agrees with the positions of
remnants of this bar that are retained in the modern bird skull.
Having myself seen some of Walker’s evidence in Sphenosuclzus, I am
satisfied that his descriptions of Sphenosiichus are accurate and that his
anatomical interpretations are reasonable. As for the similarities he cites
between Sphenosuchtrs and certain modern birds, the important question to be
answered is: Just what is their significance? I am concerned about the validity
of equating a few anatomical features in a primitive archosaur of Late Triassic
age(!) with similar anatomical features of a few modern birds (or modern
crocodiles, for that matter) and concluding that they represent real evidence
(homologous rather than homoplastic structures) of close evolutionary
relationship. Separated as they are by more than 200 million years, it would
seem to me that any modern bird is so far removed from any Late Triassic
6<
proavian” that the significance of the above similarities must be very
doubtful. For exactly these reasons, I remarked in an earlier section (p. 100)
that there was no value whatever in referring to anatomical conditions of
modern birds in our quest for the ancestral stock of Archaeopteryx. If we
accept that Archaeopteryx is close to the ancestral stock of birds, then modern
birds can tell us only what has happened since the Archaeopteryx stage, but
nothing about what happened before.
It is unfortunate that the Berlin and Eichstatt skulls d o not enable us to
establish whether any of these “Sphenosuchus-bird” conditions were also
present in Archaeopteryx-the earliest available stage of non-thecodontian
avian evolution. It is hoped that further preparation of the underside of the
Eichstatt specimen will be possible in the future and may reveal some of those
features. My own examinations of the Eichstatt specimen have led me to the
conclusion that the quadrate in Archaeopteryx probably was streptostylic, and
this means that there may have been some kinesis, as theorized by Bock (1964)
and Wellnhofer (1974). If these and other details cited above could be verified
in Archaeopteryx, it would do much to validate the significance that Walker
attributes to the similarities he finds between Sphenosuchus and some modern
birds.
Aside from the features noted by Walker, other published data on
Sphenosuchus (Haughton, 1915; von Huene, 1925; Broom, 1927) seem to
indicate the same dissimilarities with Archaeopteryx that were noted above in
pseudosuchians, namely:
(a) A single small antorbital fenestra with no indication of subsidiary
fenestrae.
(b) Heterodont dentition consisting of relatively large teeth. A wide
diastema at the premaxillary-maxillary suture, filled by a large tusk of the
lower jaw.
 ARCHAEOPTERYX AND THE ORIGIN OF BIRDS 163

Figure 35. The right scapulo-coracoid of Sphenosuchus mums (A) compared with that of
Archaeopteryx (B). Both specimens are viewed in lateral aspect but are not drawn to the same
scale. A is taken directly from Broom ( 1 9 2 7 ) . with no scale given. B is a reconstruction by the
author based largely on the London and Berlin specimens. Horizontal scales = 2 cm.

(c) Suspensorium, although steeper, still sloping (descends) backward; jaw

articulation still posterior to the occipital condyle.
(d) Mandible deep and slightly shorter than the upper jaw and bearing a
large external mandibular fenestra.
In the post-cranium, the scapula (Fig. 35). expands distally into a broad
flaring blade, quite unlike the narrow strap-like blade of Archaeopteryx. The
coracoid is elongated after the fashion of modern crocodilians, but in no way
does it resemble the peculiar elongated design of modern carinate birds. Any
suggestion that i t does so is entirely false. But, even more important, the great
relative length of the coracoid in Sphenosuchus cannot possibly have any
bearing on the elongation of the avian coracoid because in Archaeopteryx,
which is more recent in time, and presumably much higher up the avian ladder
than any sphenosuchid, the coracoid is relatively much shorter and more
quadrangular (see Fig. 14A, B).
In their general configurations, neither the skull and jaws, nor the
scapulo-coracoid of Sphenosuchus is remotely suggestive of affinities with
Archaeopteryx, despite the possible streptostyly and kinesis that may have
existed in both genera. Furthermore, the logic of a close relationship between
Sphenosuchus and ancestral birds is greatly diminished by the enormous time
gap between the Late Triassic Sphenosuchus and the living birds with which it
has been compared.

EVIDENCE SUPPOSEDLY CONTRARY TO

A THEROPOD ANCESTRY FOR ARCHAEOPTERYX

The pseudosuchian theory of bird origins is so widely accepted today that it

would be foolish not to expect some strong reactions opposing the theropod
164 J. H. OSTROM

ancestry proposed here. For this reason, it seems advisable for me to comment
on some of the counter-arguments and criticisms that have been raised in the
past, as well as others that are likely to come up in the future. Some of the
criticisms that can be anticipated are:
( 1 ) No theropod specimen of the proper geological age (pre-Kimmeridgian)
is known that possesses all the features required of an immediate ancestor of
Archaeopteryx.
(2) The anatomical evidence that I have presented represents a
“conglomeration of parts” taken from a variety of coelurosaurian theropods to
construct a makeshift and purely hypothetical bird ancestor.
( 3 ) Nearly all the coelurosaurian evidence cited by me postdates
Archaeopteryx, or at the very best is contemporaneous (Ornitholestes,
Compsognathus) and therefore cannot be related t o ancestral forms.
(4) The predominant trend among theropods was a shortening of the
forelimb, not elongation as would clearly be required in an Archaeopteryx
ancestry.
( 5 ) The bird-like orientation of the pubis in the Berlin specimen isnot like
that of coelurosaurian dinosaurs.
(6) The clavicle is absent in coelurosaurian dinosaurs.
(7) There is no ossified sternum known in theropods.
( 8 ) The bird-like features of some coelurosaurs are most logically explained
as parallelisms or convergent features.
Taking each of these in order, we must recognize the first criticism for
exactly what it is-negative evidence only. The fact that no “suitable”
coelurosaurian “pre-Archaeopteryx ” is known to us is neither surprising nor
significant. First of all, terrestrial vertebrate remains of Early and Middle
Jurassic age are extremely rare, as compared with those known from later
Jurassic or Late Triassic times. But even if an extensive fossil record of earlier
Jurassic vertebrate life were known to us, discovery of the immediate
antecedent of Archaeopteryx would be extremely improbable. The fact that
such an ancestor has not been found (or recognized) as yet does not establish
that it never existed. Obviously it did, whatever it was. We must be particularly
cautious about drawing absolute conclusions from negative evidence. For
example, an equally illogical alternative argument is that the absence of any
known “suitable” intermediate form between Late Triassic pseudosuchians and
Archaeopteryx is compelling evidence that birds could not have evolved from a
pseudosuchian ancestor.
As for the second criticism, it is quite correct that I have assembled many
Archaeopteryx-like anatomical features from a number of different theropods
and that no theropod possesses all these features. The last is due in part to the
fact that many of the taxa involved are known only from incomplete remains
(in most instances, far less complete than most of the skeletal specimens of
Archaeopteryx). For example, the carpus is not known in Ornitholestes (nor is
the pubis or coracoid). The carpus is known, however, in Deinonychus and
Velociraptor which have hands very much like those of Ornitholestes-and
Archaeopteryx. The carpals of these genera also resemble those of
Archaeopteryx. The fact that each of these many Archaeopteryx-like
anatomical features occurs in more than one coelurosaur is extremely
 ARCHAEOPTERYX AND T H E ORIGIN OF BIRDS 165

important, indicating that none is restricted in its occurrence and that all may
have been widespread among coelurosaurs. We may regard them as general
“coelurosaurian” traits.
The observant reader will have noticed that I have made few comparisons
with Triassic coelurosaurs (e.g., Coelophysis, Halticosaunw, Procompsognathus)
that might have been directly ancestral to Archaeopteryx. That much of the
coelurosaurian evidence cited here is drawn from taxa post-dating
Archaeopteryx is a valid criticism, except that I am not proposing any of these
taxa as ancestral to Archaeopteryx. As with the first criticism, there is no
evidence that these same features, or their precursors, did not exist prior to
Late Jurassic times. These characters are considered as representing the
“coelurosaurian state’’ and there is ample evidence (Coelophysis, Segisaums,
Procompsognathus. Compsognathus, Ornitholestes, Coelurtts) that the
“coelurosaurian state” existed long before Archaeopteryx. Triassic genera were
not utilized in this analysis for the simple reason that they are more primitive
in most features and less like Archaeopteryx than are many later coelurosaurs.
With the possible exception of the fused clavicles and the unique form of the
ischium, each character of the skeletal anatomy of Archaeopteryx can be found
in more than one coelurosaur. Obviously, Ornitholestes cannot have been
ancestral to Archaeopteryx, but Archaeopteryx and Ornitholestes could have
had a common coelurosaurian ancestor. Any other explanation for the
“coelurosaurian state” of Archaeopteryx seems contrived, to say the very least.
Tucker (1938a) and several other authors have observed that a major trend
among (theropod) dinosaurs was the pronounced reduction of the forelimb,
“tending to become non-functional”. Tucker (1938b) even went to the
extreme of suggesting that terrestrial bipeds (presumably all kinds) almost
invariably undergo forelimb reduction. Both of those observations are only
partly true. Forelimb reduction was typical of some, but not all, of the large
carnosaurian theropods, and it was not true of most small or medium-sized
coelurosaurian theropods*. In fact, forelimb elongation is clearly evident in
Ornitholestes, Velociraptor, Deinonychus and all struthiomimids. Ample
evidence now exists, much of it discovered since Tucker’s time, to show that
evolutionary trends in the forelimbs of theropods were not all contrary to that
required in the ancestry of the very long-armed Archaeopteryx.
With regard to the fifth criticism predicted above, concerning the
non-theropod-like orientation of the pubis preserved in the Berlin specimen of
Archaeopteryx, it was demonstrated above that the pubis was preserved in an
unnatural position. In my opinion, the evidence for this is beyond dispute, but
for those who are unable to accept that evidence, it must be conceded that the
pubis rotated back from a reptilian position sometime during the course of
Forelimb shortening in theropods has traditionally been correlated with large size (as in
Corgosaurus = Albertosaurus, Tyrannosaurus, Tarbosaurus, Daspletosaurus). but Deinocheirus (Osm6lska
& Roniewicz, 1969), with its nearly 9 foot-long forelimbs was certainly large, even if not clearly
“carnosaurian”. Furthermore, Compsognathus, the classic specimen of a “coelurosaur”. which is almost
exactly the same size as Archaeopteryx, possesses distinctly shortened forelimbs and hands very much like
those of the large “carnosaurian” genera listed above. Forelimb shortening among theropods seems to
have been universally correlated with two anatomical conditions other than that of absolute size: (1) a
very large relative skull size combined with a relatively short neck, and (2) reduction o f the manus to
digits I and 11. Except for Deinocheirus, in every one of the theropods mentioned above, (including
Compsognathus), skull length is comparable to or greater than the total length of the forelimb, and the
manus consists of only two fingers; in Deinocheirus, in any case, the skull is unknown.
166 J. H. OSTROM

avian evolution. Whether this rotation occurred before or after Archaeopteryx

is not of crucial importance. Existing evidence suggests that pubic rotation had
already begun before the Archaeopteryx stage, because in Archaeopteryx its
position in life seems to have been intermediate between that of theropods and
that of later birds.
The sixth point, discussed in some detail earlier, is the only specific evidence
that has ever been raised for rejection of a coelurosaurian ancestry.of birds: the
supposed absence of clavicles in theropods. The avian furcula is currently
believed to represent the co-ossified clavicles, and nearly all developmental
biologists consider the clavicles to be dermal rather than endochondral
ossifications. They are known to be somewhat variable in their degree of
ossification, presumably varying in accordance with the kinds and intensities of
stresses imposed on the shoulder joint and the degree of stability or mobility
required there, Even among carinates, for example, the clavicles may be
unfused or even greatly reduced, as was shown by Glenny & Friedmann (1954)
and reported further by Van Tyne & Berger (1959). As dermal elements, their
apparent absence in any particular theropod specimen might well be the result
of their having existed in a membranous state, which would not be preserved as
fossil evidence.
Another intriguing aspect of the clavicle-furcula problem is that raised by
Lansdown (1968)*. He presented evidence that the furcula of the Japanese
quail (Coturnix c. juponicu) is endochondral, at least in part, rather than dermal
in origin. This raises two important questions: first, is the avian furcula really
derived from fusion of the paired clavicles? and second, are endochondral and
dermal osteogenesis mutually exclusive? If the answer to the last question is
yes, then the answer to the first must be no-and the putative absence of
theropod clavicles has no bearing on the question of bird origins. The furcula
would then be a de ~ O Y Ostructure, inasmuch as neither the clavicles nor the
interclavicle alone could have given rise to an endochondral element.
As with the supposed absence of clavicles, the lack of a certifiable sternum in
theropodst is negative evidence and of no significance, especially since it
apparently is lacking in Archaeopteryx also. The “proof” of this statement is
the complete absence of a sternum in all known specimens of pseudosuchians-
the almost unanimously accepted ancestral stock of all later archosaurs, all of
which (crocodilians, pterosaurs, ornithischians, sauropods, birds, but perhaps
not theropods) possess cartilaginous or well ossified sterna. If the absence of
(dermal?) clavicles among the theropods is sufficient cause for discarding that
group as ancestral to birds, then surely the absence of a (endochrondral)
sternum in all known thecodontians is an equally valid reason for dismissing
pseudosuchians from the ancestry of all later archosaurs-birds included. As for
the ancestry of Archaeopteryx, the importance of a sternum is questionable,
because no sternum has been identified positively in any of those specimens.
The final criticism listed at the beginning of this section, the parallel or
convergent explanation of the “bird-like” features of some coelurosaurian
dinosaurs, is likely to be the most frequently invoked argument against a
I am indebted to Mr John Attridge of Birkbeck College, University of London, for bringing
Lansdown’s work to my attention.
t Lambe (1917) described a poorly preserved bone in Corgo~urus(=Alberrosoums) which he
considered to be part of the sternum.
 ARCHAEOPTERYX AND THE ORIGIN OF BIRDS 167

theropod origin of birds. Fiirbringer (1888) was the first to raise this
explanation and more recently it has been clearly and emphatically restated by
Simpson (1946):
“Almost all the special resemblances of some saurischians to birds so
long noted and so much stressed in the literature, are demonstrably
parallelisms and convergences. These cursorial forms developed
strikingly bird-like characters here and there in the skeleton in one
genus or another. They never showed a general approach to avian
structure (as do Arcliaeopteryx and Archaeornis), some avian
characters were not achieved or even hinted at in any of them, and
they all retain the most conclusive marks of the reptilian nature.”
(Simpson, 1946: 94-5)
De Beer (1954b) followed suit; concluding that the dinosaurs could not be
ancestral to the birds:
“. . . f o r many of the points of resemblance which they appear to
share with birds can easily be proved to be spurious,. . .” (de Beer,
1954b: 45)
These statements, and variations on the same theme, have been repeated so
often that they are now generally accepted as fact. But the important and
surprising fact that has been overlooked by everyone is that these and similar
statements are not directed at the critical issue. They are not addressed to the
question of the origin of Archaeopteryx and they do not explain the
dinosaurian nature of Archaeopteryx.
Although there are differences of opinion as to whether Archaeopteryx is on
the main line of descent to modern birds, there seems to be no difficulty for
most in visualizing the evolutionary transition from an Archaeopteryx-like stage
to the modern bird. The bird-like features of Archaeopteryx are accepted by
all, and only once (Lowe, 1935) has any avian feature of Archaeopteryx
(feathers) been attributed to multiple (parallel or convergent) origins. But
inexplicably, instead of addressing the question to the source of the avian and
non-avian characters of Archaeopteryx, queries have been directed at the
irrelevant “bird-like” characters of some dinosaurs. Consider for a moment: If
Archaeopteryx was derived from a coelurosaurian ancestor, as I believe, then it
is understandable why some coelurosaurs also have a few bird-like features.
This also accounts for the many coelurosaurian features of Archaeopteryx. In
seeking the origin of Archaeopteryx, the crucial question is: Which Mesozoic
reptiles are most similar to Archaeopteryx?-not which ones are most similar t o
modern birds. Rephrasing the above “explanation”: Are we now to believe that
those coelurosaurian-like characters of Archaeopteryx are just parallel or
convergent features? Simpson (1961) observed that:
‘I. . . intricately
co-ordinated structures are less liable to close
convergence and an aspect of that fact is that they tend to be less
labile and to retain ancestral conditions longer. ” (Simpson, 1961:
100) (My italics)
Certainly this must apply to such functionally co-ordinated structures as the
manus, carpus and forelimb, or pes, tarsus and hindlimb of Archaeopteryx and
168 J. H. OSTROM

coelurosaurs. Given the incomplete nature of the fossil record, the almost
simultaneous acquisition of so many coelurosaurian characters in Archaeop-
teryx and in coelurosaurs by means of parallel or convergent evolution is, in my
judgement, infinitely less probable than by means of simple evolutionary
descent from a common coelurosaurian stock.
Some other objections that might be raised to a theropod origin of
Archaeopteryx are:
(a) The presence of a vestigial fourth metacarpal in some theropods
(Ornitholestes) and of a complete fourth digit in the manus of some others
(Coelophysis, Ceratosaurus, Procompsognathus).
(b) The presence of a vestigial fifth metatarsal in some theropods
(Ornithomimus, Struthiomimus, Deinonychus).
(c) The supposed absence of sclerotic plates in theropods.
The existence of vestigial digits over and above the number present in
Archaeopteryx hardly seems to be valid evidence against the affinities suggested
here when it is obvious that the prevalent condition among theropods is the
complete loss of the fourth and fifth fingers and the fifth toe-exactly as in
Archaeopteryx. The retention of these structures in a few taxa is merely the
retention of a more primitive state and neither precludes nor verifies phyletic
relationship with Archaeopteryx. ( I t may be recalled from a preceding section
that there is evidence of a vestige of the fifth metatarsal in the Eichstatt
specimen; see Fig. 16A.) As for the absence of sclerotic plates in theropods, that
too is invalid because they have been reported in the theropods Struthiomimus
(Parks, 1928) and Dromaeosaurus (Colbert & Russell, 1969).
In summary, all the major criticisms that have been, and may still be, voiced
against a direct evolutionary relationship between theropod dinosaurs and
Archaeopteryx have been found to be inconclusive, incorrect or irrelevant. In
my opinion, no conclusive evidence exists for rejecting coelurosaurs as the
immediate ancestral stock of Archaeopteryx. On the contrary, this relationship
is supported by a large body of positive anatomical evidence.

AFTER HEILMANN

Although it was first suggested by Broom (1913), Heilmann is properly

credited with laying the foundation of the pseudosuchian or “common
ancestor” theory of bird origins. However, it has been the opinions and
evaluations of subsequent scholars that have established that theory as the
prevailing view so widely held now by most ornithologists and paleontologists.
Because the thesis presented here is contrary to that theory, it is appropriate to
examine some of the more influential views that have been expressed since
Heilmann-the remarks that have contributed to the increasingly favourable
climate within which Heilmann’s ideas have been examined.
One of the most important participants, and the first to digress from
Broom’s and Heilmann’s theory, was Percy Lowe (1935, 1944), who
maintained that Archaeopteryx and Archaeornis were not birds at all, but were
feathered dinosaurs; that they also were not ancestral to birds; and that ratites
(his Struthiones) were not descended from volant ancestors (a view contrary to
that held then and now by most ornithologists), but had arisen instead from
 ARCHAEOPTERYX AND THE ORIGIN OF BIRDS 169

bipedal, coelurosaurian dinosaurs. His most important contribution was in

recognizing that the skeletal anatomy of Archaeopterjjx was not avian but
rather was almost entirely coelurosaurian, the details of which he clearly
enumerated. Those conclusions were largely ignored by the scientific
community, probably because his other ideas were-in Lowe’s own words-“a
zoological transgression”. Most ornithologists rejected Lowe’s ideas on ratite
origins; this rejection was later validated by de Beer (1956),who showed that
ratite anatomy was inexplicable unless it had been derived from a flying
ancestor. This presumably increased general scepticism about some of Lowe’s
other ideas, especially because his dinosaurian label for Archucopteryx implied
an unlikely diphyletic origin of feathers. Lowe’s paper prompted a strong
response from Tucker (1938a) in which it was noted that Archaeopteryx and
Archaeornis were indeed extraordinarily reptilian, but that there was nothing in
their organization that would not be expected in primitive birds just emerged
from reptilian stock. I am in complete agreement with that statement. Tucker
(19 38b) further wrote:
“The reptilian ancestry of birds is so self evident and so universally
recognized by zoologists that it can be taken as axiomatic in any
discussion. We shall further accept the view that the immediate
reptilian ancestors of birds, if known, would have to be placed in or
extremely close to the group Pseudosuchia. The direct derivation of
birds from dinosaurs, favoured by some earlier writers, would
probably not be advocated by any competent zoologist at the
present day, but the many similarities between the two groups
suggest a common origin.” (Tucker, 1938b: 3 22)
Since Tucker’s remarks, a wealth of new dinosaeian evidence has come t o light
which, I hope, removes all advocates of a dinosaurian origin of birds from the
ranks of the incompetent. Lowe’s second paper (1944) prompted an even
stronger response by Simpson (1946):
“Archaeopteryx and Archaeornis are intermediate between reptiles
and birds in structure and their bearing on the origin of birds is
unchanged by the purely verbal question of whether t o call them
reptiles or birds. . . . I t is, indeed, an interesting point that these
Jurassic birds (as I shall continue t o call them) are more reptilian
than might have been expected in an animal that had already
developed a feathered wing-a point as strongly emphasized by
Heilmann as by Lowe although Heilmann did not question the
position of these animals near or, at least structurally, in the avian
ancestry. The only logical conclusion is, I think, that the primary
avian structure was the feathered wing which developed as a flying
apparatus, a conclusion in no way negatived by its being as yet
unperfected in the Jurassic, even if, as Lowe believes, the wing was
then fit only for gliding rather than flapping flight.” (Simpson, 1946:
94)
Simpson further found Lowe’s designation of Archaeopteryx and Archaeornis
as dinosaurs to be “nothing short of fantastic’’ (footnote p. 94).
After such strong reactions as these, it is not surprising that there have been
170 J . H. OSTROM

so few critics of the pseudosuchian theory. Rut, in response to Simpson’s

footnote, the question must still be asked: How would those fossil remains
have been identified-indeed, how would they now be classified, if no feather
imprints had been preserved in any of those specimens? The skeletal anatomy
of Archaeopteryx, as 1 have demonstrated, is almost entirely coelurosaurian
(just as Lowe reported, and as Heilmann before him had concluded) and
includes only one exclusively avian character-the furcula. In fact, it is only
because of the distinct feather impressions preserved in two of the specimens of
Archaeopteryx that we now have any knowledge at all about Jurassic birds or
about the origin of birds. In the absence of those feather impressions, I d o not
believe that any of the specimens of Archaeopteryx would ever have been
recognized as avian, or even as remotely related to birds. Regardless of how one

Figure 36. Skeletal reconstruction of two Late Jurassic bipedal predators; Ornitholestes
h e m a n n i (above), a coelurosaurian dinosaur from the Morrison Formation (Kimmeridgian age)
of North America, and Archaeopteryx lithographica (below) from the Solnhofen Limestone
(Kimmeridgian age) of Europe. Reconstruction of Ornitholestes is from Osborn (1903); that of
Archaeopteryx is by the author based o n the Berlin specimen. Scales = 5 cm.
 ARCHAEOPTERYX AND THE ORIGIN OF BIRDS 171

assesses the importance of the systematic placement of Archaeopteryx, Lowe’s

dinosaurian label is not as fantastic as Simpson thought. Unfortunately,
however, that designation by Lowe completely obscured the real significance of
his observations.
Holmgren (1955) was another who found the resemblances between
Archaeopteryx and coelurosaurian dinosaurs of special interest. He presented
extensive embryological evidence (from Struthio, Anser and Gallus) which he
compared with the anatomy of Archaeopteryx and various coelurosaurs, and
concluded:
“That the Coelurosaurs agree more closely with the Saururae
[Archaeopteryx and Archaeomis] than the Pseudosuchians d o and
that there is thus better evidence that the birds have a coelurosaurian
than a pseudosuchian ancestry. As the coelurosaurs are probably
derived from Pseudosuchians, we arrive at the phylogenetic series
Pseudosuchians-Coelurosaurs-birds. I f we were to assume that the
birds are direct descendants of Pseudosuchians, we would have t o
seek Jurassic connecting links between the Triassic Pseudosuchians
and the Upper Jurassic Saururae. But no such links have hitherto
been recorded.” (Holmgren, 1955: 307)

I t is unfortunate that so little attention has been given to Holmgren’s paper,

but it seems to have been largely ignored, perhaps because it was preceded the
year before by de Beer’s (1954b) monographic study of the London
Archaeopteryx, which reemphasized most of Simpson’s (1946) conclusions. Or
perhaps it was dismissed because Holmgren, like Lowe (1935, 1944) before him
and Glutz von Blotzheim (1958) later, also believed in an independent origin of
ratites from the larger Cretaceous coelurosaurs. But it must be noted here that
Holmgren’s work was published posthumously, and the introductory editorial
comment clearly states that Holmgren considered the phylogenetic section as
incomplete and only a tentative outline indicating the course that future
research should pursue. In retrospect, though, it appears that all those who
once argued in favour of a non-carinate origin (and especially a dinosaurian
origin) of ratites have paid the price by having their other ideas rejected as well.
The great debate over the separate origins of ratites and carinates was settled by
de Beer (1956) who demonstrated that the structure of the ratite wing,
pygostyle and cerebellum were inexplicable unless they had been retained from
a flying ancestor.
In his discussion of the ancestry of Archaeopteryx, de Beer (1954b) also
considered the supposedly “spurious” resemblances between dinosaurs and
birds, noting the “lost” clavicles of dinosaurs, spurious similarities in the
forelimb, and that the backwardly directed pubis of birds was unrelated t o the
post pubis of ornithischian dinosaurs. On these grounds, he re-affirmed the
conclusions of Tucker (1938b) and Simpson (1946) and accepted the
pseudosuchian theory of Broom and Heilmann, a position he continued t o
maintain in later years (1964). This same phylogeny has been adhered t o by
nearly everyone since, apparently without question (see Swinton, 1960, 1964;
Welty, 1962; Romer, 1966, 1968; and Brodkorb, 1971, to mention only a
few). Only Bock (1969a) qualified his acceptance of a pseudosuchian origin:
172 J . H.OSTROM

“The pseudosuchians are generally accepted as the most probable

reptilian ancestor of birds; however, acceptance of this group is more
by default than by direct demonstration. Pseudosuchians may have
been chosen because they also gave rise to the crocodiles, the living
reptiles with which birds share the greatest number of characters.
Although pseudosuchians have not been proven to be avian
ancestors, they possess no features to discount this possibility.”
(Bock, 1969a: 1480)
Bock’s final sentence above is absolutely correct, but the same can be said of
coelurosaurs-only more so.
Most recently, two papers have appeared which, although accepting my
coelurosaurian ancestry theory, in the long run may generate strong reactions
unfavourable to my thesis on bird origins. Because of that possibility, I feel that
some comment is appropriate here, despite the fact that neither paper is
primarily concerned with the subject of bird origins. The papers in question are
one by Bakker & Galton (1974) on dinosaur monophyly and a reply by
Thulborn (1975) on dinosaur polyphyly. Bakker & Galton present very
interesting evidence which they believe supports a monophyletic origin of the
two dinosaurian orders (Saurischia and Ornithischia), which they elevate to
subclass rank for inclusion in a proposed new vertebrate class, Dinosauria. Their
new class is established primarily on purely speculative grounds that all
dinosaurs were probably endothermic and possessed high levels of exercise
metabolism. They then propose that the class Aves be reduced in rank to a
subclass of their new class Dinosauria on the grounds that:
“the avian radiation is an aerial exploitation of basic dinosaurian
physiology and structure, much as the bat radiation is an aerial
exploitation of basic primitive mammal physiology. Bats are not
separated into an independent class merely because they fly. We
believe that neither flight nor the species diversity of birds merits
separation from dinosaurs on a class level.” (Bakker & Galton, 1974:
171)
While I appreciate their acceptance of my conclusions about the ancestral
affinities of Archaeopteryx and later birds, I reject the assertion by Bakker &
Galton that the avian radiation is merely an aerial exploitation of basic
dinosaurian physiology and structure, as well as their reasoning that birds
should therefore be classified as dinosaurs. Bakker & Galton do not know what
dinosaurian physiology was; no one does. And for them to suggest that the
avian radiation was no more successful (by any criterion) than that of bats is
patently false. The possibility (I would like t o believe, probability) that
Archaeopteryx. and presumably all subsequent birds, evolved from one
particular group of dinosaurs does not justify such a radical departure from
conventional classification schemes.
Thulborn (1975), in a well-phrased reply to Bakker & Galton, claims that
there is no convincing evidence that dinosaurs were endothermic and challenges
their evidence of dinosaurian monophyly with evidence of his own for
polyphyly. But of interest here is Thulborn’s proposed classification which
transfers the ancestors of birds (the entire dinosaurian suborder Theropoda) to
the class Aves.
 ARClfAEOPTER YX AND THE ORIGIN OF BIRDS 173

I confess that I am unable t o accept such theropods as Tj,ru/ztzosuuriis and

All~~saunisas “birds”, and therefore have little sympathy with this
re-classification scheme either. Thulborn’s approach is comparable to that of
Reed (1960), who proposed the expansion of the class Mammalia t o include all
therapsids and sphenacodont pelycosaurs. As Simpson (1960, 1961) and others
have repeatedly urged, classifications should be as stable and utilitarian as is
consistent with the acquisition of new evidence and methodology. ‘The new
view that birds may be descendant from some group of theropod dinosaurs,
rather than from pseudosuchian thecodontians, does not require (or justify) the
radical systematics proposed by Bakker & Galton and by Thulborn. Nor are
their classifications justifiable on utilitarian grounds.

SUMMARY

The question of the origin of birds can be equated with the question of the
origin of Arcliueopterj>x. This last question evokes two possible answers,
depending upon how one views the importance of “primitive versus derived
characters” in assessing phylogenetic relationships. One possible answer is:
Arclzaeopteryx is a direct descendant of some unknown, but presumably
Eirpurh-eria-like pseudosuchian. This answer is predicated on the belief that
Archaeopteryx only parallels or converges with various coelurosaurs in certain
skeletal similarities. This is the view now held by the majority of biologists- a
view that I find unacceptable. The second possible answer is: Arcliueopteryx is
directly descendant from a small unknown Ornitholestes-li ke coelurosaurian
dinosaur. This answer assumes that skeletal similarities between coelurosaurs
and Archaeopter.r*s are derived from a common ancestor, itself a coelurosaur.
This is the view advocated here.
There is no evidence to support an ornithischian ancestry of birds. The pubis
of Archaeopteryx apparently was not reflected backward as in ornithischians
and modern birds, and in any case, the ornithischian pubis is only superficially
like that of living birds. Nor is the so-called ornithopod foot like that of birds.
Evidence of close theropod-Archaeoptervx relationships, however, is
abundant: the presence of the same, multiple, specialized adaptations in both
Archaeopteryx and various coelurosaurs (tridactyl manus, metacarpus and
carpus morphology, forelimb and pectoral girdle structure, four-toed pes,
reversed hallux, metatarsal morphology, mesotarsal joint, hindlimb construc-
tion, pelvic form, plus elongated forelimbs, bipedal posture, vertebral structure
and formula, and basic cranial morphology).
The presence in Archueopteryx, coelurosaurs and pseudosuchians of several
primitive characters in common (thecodont dentition, sclerotic ring, possibly
amphicoelous vertebrae, long caudal series, gastralia, pubic symphysis, short
coracoids) indicates only a probable common ancestry. It does not establish
that the Coelurosauria could not have given rise t o Ardzueopter.vs-and higher
birds. There is n o evidence (outside of Lagostichiis and Lagerpeton) of shared
derived characters to suggest a close evolutionary relationship between classic
pseudosuchians and Archaeopteryx. Similarly, there is no clear-cut evidence in
the form of shared derived characters to link Archaeopteryx with
Splienosuchus.
The absence of clavicles in theropods (now known to be false), once
174 J. H. OSTROM

considered as conclusive evidence against a coelurosaurian ancestry of birds, is

no more significant than is the absence of a sternum in all known
pseudosuchians as evidence against a pseudosuchian ancestry of all other
archosaurs. The absence of any known “ideal” coelurosaurian pre-
Archaeopteryx is only negative and inconclusive evidence, especially in view of
our meagre and exceedingly deficient knowledge about Early and Middle
Jurassic terrestrial vertebrates.
All available evidence indicates that the immediate ancestor of
Archaeopferyx was a small coelurosaurian dinosaur and that the phylogeny of
avian ancestry was: Pseudosuchia-Coelurosauria-Archaeopteryx-higher birds.

ACKNOWLEDGEMENTS

I am very much indebted to Alan Charig and Cyril Walker of the British
Museum (Natural History), London; Herman Jaeger of the Humboldt Museum
fur Naturkunde, East Berlin; The0 Kress of the Solenhofer Aktien Verein,
Solnhofen; C. 0. van Regteren Altena of Teyler’s Stichting, Haarlem; and Peter
Wellnhofer of the Bayerische Staatssammlung, Munich for the many courtesies
extended t o me during my visits to their respective institutions, and especially
for the privilege of studying the specimens of Archaeopteryx in their care.
Without their generosity and assistance, this study would have been impossible.
The manuscript was reviewed at various stages by A. J. Charig, A. W.
Crompton, F. A. Jenkins, Jr., Storrs Olson, Colin Patterson and Dale A.
Russell. I gratefully acknowledge their very thoughtful and valuable suggestions
and criticisms. Discussions with students and colleagues played a major part in
shaping this study: my sincere thanks to all. This research was supported by
grants from the National Science Foundation (Grant No. G.B. 14033), the
Frank M. Chapman Memorial Fund of the American Museum of Natural
History, and the John T. Doneghy Fund of the Yale Peabody Museum.

REFERENCES
ABEL, 0.. 1912. Grundzuge der Palaeobiologie der Wirbeltiere: xv + 7 0 8 pp. Stuttgart:
Schweizerbartsche.
ARMSTRONG. G. T., 1966. The fable of the first fatar flight: 1 2 pp. Pasadena, California: Ambassador
College.
ARMSTRONG, G . T. & KROLL, P. W.. 1967. A theory f o r the birds: 32 pp. Pasadena, California:
Ambassador College.
BAKKER, R. T. & GALTON, P. M., 1974. Dinosaur monophyly and a new class of vertebrates. Nature,
Lond.. 248: 168-72.
BAUR, J . G., 1883. Der Tarsus der Vogel und Dinosaurier. Morph. J b . , 8: 417-56.
BAIJR, J. G.. 1884a. Note on the pelvis in birds and dinosaurs. A m . Nat., 18: 1273-5.
BAUR, J. G., 1884b. Dinosaurier und Vogel. Eine Erwiederung an Herrn Prof. W. Dames in Berlin.
Morph. Jb., 10: 446-54.
BAUR, J. G., 1885a. Bemerkungen uber das Becken der Vogel und Dinosaurier. Morph. Jb., 10: 61 3-6.
BAUR, J. G., 1885b. Zur VBgel-Dinosaurier-Frage. Zool. Anr., 8: 441-3.
BEECHER, W. J.. 1962. The biomechanics of the bird skull. Bull. Chicago Acad. Sci.. 11: 10-33.
BEER, G. R. de, 1954a. Archaeopteryx and evolution. Advmt Sci.,London., 11: 160-70.
BEER, G. R. de, 1954b. Archaeopteryx lithographica; a srudy based on the British Museum specimen:
68 pp. London: Br. Mus. (Nat. Hist.)
BEER, G. R. de, 1956. The evolution of ratites. Bull, Br. Mus. nar. Hist., (Zool.). 4: 57-70.
 ARCHAEOPTERYX AND THE ORIGIN O F BIRDS 175

BEER, G. R. de, 1964. Archaeopteryx. In A. L. Thomson (Ed.), A new dictionary of birds: 58-62.
London: Nelson.
BOAS, J . 1;. V.. 1930. Uber das Verhiltnis der Dinosaurier zu den VOgeln. Morph. J b , 64: 223-47.
BOCK, W. J., 1964. Kinetics of the avian skull. J. Morph., 114: 1-42.
BOCK, W. J.. 1969a. The origin and radiation of birds. Ann. N . Y . Acad. Sci.. 167: 147-55.
BOCK. W. J.. 1969h. Discussion: The concept of homology. Ann. N . Y . Acad. Sci., 167: 71-3.
BOCK, W. J , , 1969c. Comparative morphology in systematics. In C. G . Sihley (Ed.). Systematic biology:
4 1 1 4 8 . Washington D.C.: Nat. Acad. Sci.
BOCK, W. J., 1969d. Nonvalidity of the “Phylogenetic Fallacy”. Syst. Zool., 18: 1 1 1-5.
BONAPARTE, J. P . , 1971a. Cerrirosaurus binsfeldi Price, Tipo de una nueva familia d e tecodonres
(Pseudosuchia-Proterochampsia). Anals Acad. bras. Cienc., 43: 417-22.
BONAPARTE, J . F., 1971b. Los tetripodos del sector de la Formaci6n Los Colorados, La Rioja,
Argentina. Opera lilloana. XXII: 1-183.
BRODKORB, P.. 1971. Origin and evolution of birds. In D. S. Farner & J . R. King (Eds). Aviuti biology:
19-55. New York & I.ondon: Academic Press.
BROOM, K., 1906. On the early development of the appendicular skeleton of the ostrich. with remarks
on the origin of birds. Trans. S. Afr. phil. Soc., 16: 355-68.
BROOM, K., 191 3. On the South African pseudosuchian Euparkeria and allied genera. Proc. zool. Soc.
Lon., 1913: 619-33.
BROOM. K., 1927. On Sphenosuchus, and the origin of the crocodiles. Proc. zool. Soc. London., 1927:
359-70.
BRUNDIN. L.. 1968. Application of phylogenetic principles and evolutionary theory. In T. 0rvig (Ed.).
Currenr problems of lower vertebrate phylogeny: 471-95. Proc. 4th Nohel Symposium. Stockholm:
Almqvist & Wiksell.
CAMP, C. L., 1936. A new type of small hipedal dinosaur from the Navajo sandstone of Arizona. Univ.
Calif. Publsgeol. Sci.. 24: 39-56.
CHARIG, A. J., 1972. The evolution of the archosaur pelvis and hindlimb: an explanation in functional
terms. In K. A. Joysey & T. S. Kemp (Eds), Studies in vertebrare evolution: 121-55. Edinburgh: Oliver
& Boyd.
CHARIG, A. J., KREBS. B., SUS. H.-D. & WESTPHAL, F.. In press. Thecodontia. In Handbuch der
Paliioherpetologie. Teil 13. 1-?. Stuttgart: Gustav Rischer Verlag.
Ct1ARIG. A. J . & KlilG, 0. A., 1970. The classification of the Proterosuchia. Biol. J . Linn. Soc.. 2:
125-71.
COLBITKT. E. H., 1952. A pseudosuchian reptile from Arizona. Bull. A m . Mus. Nut. Hist., 99: 565-92.
COLBEKT, E. H.. 1969. Evolution of the vertebrates, 2nd ed.: mi + 535 pp. New York: Wiley.
COLBERT. E. 11. & RUSSELL. D. A.. 1969. The small Cretaceou’s dinosaur Dromaeosaurus. A m . Mus.
Novit., 2380: 1-49.
COLLESS, D. H., 1967. The phylogenetic fallacy. Syst. Zool., 16: 289-95.
COLLESS. D. H., 1969a. The phylogenetic fallacy revisited. Syst. Zoo/.. 18: 115-26.
COLLESS. D. H.. 1969b. The interpretation of Hennig’s “Phylogenetic Systematics”-A reply to Dr
Schlee. Svsr. Zool., 18: 134-44.
COPE, E. D.. 1867. An account of the extinct reptiles which approached the birds. Roc. Acad. not. Sci.
Philad., I86 7: 234-5.
CRACKAFT. J., 1967. Comments on homology and analogy. Syst. Zool.. 16: 356-9.
CRACRAFT, J., 1970. Mandible of Archaeopteryx provides an example of mosaic evolution. Nature.
Lond., 226: 1268.
DAMES, W.. 1884. Ueher Archaeopteryx. Palaont. Abh.. 2:3: 119-96.
DAMES. W.. 1885. Entgegnung an Herrn Dr Baur.Morph. Jb.. 10: 603-12.
DAMES, W., 1897. Ueber Brustbein. Schulter-und Beckengiirtel der Archaeopteryx. Sber. preuss. Akad.
Wiss.. 2: 818-34.
DARWIN, C.. 1859. On the origin of species b y means o f natural selection. or the preservation of
favoured races in the struggle f o r tife, 2nd ed. London.
DOLLO, L.. 1882. PremiPre note sur les dinosauriens de Bernissart. Bull. Mus. r. Hist. nut. Belg.. I:
161-80.
DOLLO. L.. 1883. TroisiPme note sur les dinosauriens d e Bernissart. Bull. Mus. r. Hist. not. Belg., II:
85-126.
EWER. R. F., 1965. The anatomy of the thecodont reptile Euparkeria capensis Broom. Phil. Duns. R .
Soc. (B). 248: 379-435.
FURBRINGER. M., 1888. Unrersuchungen zur Morphologie und Systematik der Vogel: xlix + 1751 pp.
Amsterdam: Holkema.
GALTON, P. M.. 1970. Ornithischian dinosaurs and the origin of birds. Evolution. 24: 448-62.
GEGENBAUR, C . , 1878. Grundriss der vergleichenden Anatomie: viii + 6 5 5 pp. Leipzig: Engelmann.
GEORGE, J . C. & BERGER, A. J., 1966. Avian myology: xii + 500 pp. New York & London: Academic
Press.
GHISELIN, M. T., 1969. The distinction between similarity and homology. Syst. Zool.. 18: 148-9.
12
176 J . H. OSTROM
GILMORE. C. W., 1920. Osteology of the carnivorous dinosauria in the United States National Museum,
with special reference to the genera Antrodemus (Allosairrus) and Ceratosaurus. Bull. U . S . natn. Miis.
110: 1-154.
GILMORE. C. W.. 1924a. A new coelurid dinosaur from the Belly River Cretaceous of Alberta. Eitll. Can.
Geol. Surv. Dept. Mines, 38: 1-12.
GILMORE, C . W.. 1924b. On Troodon validus: An ornithopodous dinosaur from the Belly River
Cretaceous of Alberta, Canada. Bull. Univ. Alberta. I : 7-43.
GINGERICH, P. D., 1973. Skull of Hesperornis and early evolution of birds. Nature. Lond.. 243: 70-3.
GLENNY. F. H. & FRIEDMANN, H.. 1954. Reduction of the clavicles in the Mesoenatidae. with some
remarks concerning the relationship of the clavicles to flight-function in birds. Ohio J . Sci.. 54: 111-3.
GLUT2 VON BLOTZHKIM, U., 1958. Zur Morphologie und Ontogenese von Schultergiirtel. Sternum
und Becken von Struthio, Rhea und Dromiceius. Revue. suisse 2001..65: 609-772.
GREGORY. J . T.. 1951. Convergent evolution: The jaws of Hesperornis and the Mosasaurs. Evolution, 5:
345-54.
HAUGHTON, S. H.. 1915. A new thecodont from the Stormberg heds. Ann. S . Afr. Mus., X I l : 98-105.
HKILMANN, G., 1926. The origin of birds: 208 pp. London: Witherby.
HELLER. F., 1959. Ein dritter Archaeopteryx-Fund aus den Solnhofener Plattenkalken von
Langenaltheim/Mfr. Erlanger Geol. Abhandl., 31: 3 - 2 5 .
HENNIG, W., 1966. Phylogenetic systematics: 263 pp. (Transl. by D. D. Davis & R . Zangerl) Urbana.
Illinois: Uhiv. Illinois Press.
HOLMGREN, N.,1955. Studies o n the phylogeny of birds. Acta zool., Stockh., 3 6 : 243-328.
HUENE, F. R. VON. 1914. Beitrage zur Geschichte der Archosaurier. Geol. paliiont. Abh. IN. S . ) , 13: 1-53.
HUENE. F. R. VON, 1925. Die Bedeutung der Ursprung der Krokodile. 2. indukt. Abstamm.-u.
Vererblehre, 38: 307-20.
HULL, D. L.. 1967. Certainty and circularity in evolutionary theory. Evolution. 2 I : 174-89.
HUXLEY. T. H., 1867. O n the classification of birds and on the taxonomic value of the modifications of
certain of the cranial bones observable in that class. Proc. zool. Soc. Lond., 1867: 415-72.
HUXLEY, T. H., 1868a. Remarks upon Archaeopteryx lithographica. Proc. R . SOC.Lond., 1 6 : 243-8.
HUXLEY, T. H., 1868h. On the animals which are most nearly intermediate between the birds and
reptiles. Ann. Mag. nut. Hist., (412: 66-75.
HUXLEY, T. ti., 1870. Further evidence of the affinity between the dinosaurian reptiles and birds. Q..I1
geol. Soc. 1,ond.. 26: 12-31.
KIELAN-JAWOROWSKA, 2. & BARSBOLD, R., 1972. Results of the Polish-Mongolian palaeontological
expeditions. Part IV. Narrative of the Polish-Mongolian palaeontological expeditions 1967-1971.
Palaeont. pol., 27: 5-13.
KREBS. B., 1963. Bau and Funktion des Tarsus eines Pseudosuchiers aus der Trias des Monte San Giorgio
(Kanton Tessin Schweiz). Paliont. Z., 3 7 : 88-95.
KREBS, B.. 1965. Ticiflosuchus ferox nov. gen. nov. sp. Ein neuer Pseudosuchier aus der Trias des Monte
San Giorgio. Schweiz. palaeont. A bh., 8 1 : 1-140.
KREBS, B., 1973. Der Tarsus von Rauisuchus (Pseudosuchier, Mittel Trias). Mitt. buyer. Sraarssamml.
Palaont. Hist. Geol., 13: 95-101.
KREBS, B., 1974. Die Archosaurier. Naturwiss., 6 1 : 17-24.
LAMBE, L. M., 1917. The Cretaceous Theropodous dinosaur Gorgosaurus. Mem. geol. Surv. Erch. Can.,
100: 1-84.
LANSDOWN, A. B. G., 1968. The origin and early development of the clavicle in the quail Corurnix c.
japonica. J. 2001..Lond., 156: 307-12.
LOWE, P. R.. 1935. On the relationships of the Struthiones to the dinosaurs and to the rest of the avian
class, with special reference to the position of Archaeopteryx. Ibis, (1315: 398-432.
LOWE, P. R., 1944. An analysis of the characters of Archaeopteryx and Archaeornis. Were they reptiles
o r birds? Ibis. 8 6 : 51743.
MARSH, 0. C., 1877. Introduction and succession of vertebrate life in America. Proc. A m . Ass. A d v m t
Sci.. 1887: 211-58.
MARSH, 0 .C., 1881a. Discovery of a fossil bird in the Jurassic of Wyoming. A m . J . Sci.. 13/21: 341-2.
MARSH,, 0. C.. 1881h. Jurassic birds and their allies. A m . J . Sci., (3122: 337-40.
MARYANSKA, T . & OSMdLSKA, H., 1974. Results of the Polish-Mongolian palaeontological
expeditions. Part V. Pachycephalosauria, a new suborder of ornithischian dinosaurs. Palaeont. pol., 30:
45-102.
MASLIN. T. P.. 1952. Morphological criteria of phyletic relationship. Syst. Zool.. I: 49-70.
MAYR, F. X..1973. Ein Neuer Archaeopteryx-Fund. Palaonr. 2.. 4 7 : 17-24.
MEYER, H. VON. 1857. Beitrage zur naheren Kenntniss fossiler Reptilien. Neues Jb. Miner. Geol.
Palaont., 1857: 53243.
MEYER, H. VON. 1860. Zur Fauna der Vorwelt. Reptilien aus d e m lithographischen Schiefer des Jura in
DeutSchland und Frankreich: 64-6. Frankfurt am Main: Heinrich Keller.
MEYER, H. VON, 1861a. Vogel-Federn und Palpipes priscus von Solnhofen. Neues Jb. Miner. Geol.
Palaont.. 1861: 561.
 ARCHAEOPTERYX AND THE ORIGIN O F BIRDS 177

MEYEK, H. VON. 1861b. Archaeopteryx lithographica (Vogel-Feder) und Pterodactylus von Solnhofen.
Nrues Jb. Miner. Geol. Palaont.. 1861: 678-9.
MKYEK. H. VON, 1862. Archaeopteryx lithographica aus dem lithographischem Schiefer von Solnhofen.
Paleontographica. 10: 5 3-6.
MONTACNA, W., 1945. A re-investigation of the development of the wing of the fowl. J . Morph.. 76:
87-113.
MUDCE. B. F.. 1879. Are birds derived from dinosaurs? Kansas City Rev. Sci.. 3: 224-6.
NELSON. C . J.. 1970. Outline of a theory of comparative biology. Syst. Zool., IY: 373-84.
NEWTON, E. T., 1894. Reptiles from the Elgin Sandstone-Description of two n e w genera. Phil. Trans. R .
SOC. ( B ) , 185: 573-607.
OSBORN, 14, F., 1900. Reconsideration of the evidence for a common dinosaur-avian stem in the
Permian. A m . Nat.. 34: 777-99.
OSBOKN. H. F., 1903. Ornitholesres hermanni, a new compsognathoid dinosaur from the Upper Jurassic.
Bull. A m . Mus. nat. Hist., 19: 459-64.
OSBORN, H. F.. 1917. Skeletal adaptations of Ornitholestes, Srrirthiomimus, Tyrannosaurus. Bull. A m .
Mus. not. Hist., 35: 733-71.
OSBORN. H. F., 1924. Three new Theropoda, Protoceratops zone, central Mongolia. A m . Mirs. Novit.
l4!: 1-12.'
OSMOLSKA. H. & RONIEWICZ, E., 1969. Results of the Polish-Mongolian paleontological expeditions.
Part 11. Deinocheiridae. a n e w family of theropod dinosaurs. Palaeont. pol., 21: 5-19.
OSMdLSKA, H., RONIEWICZ, E. & BARSBOLD, R., 1972. Results of the Polish-Mongolian
paleontological expeditions. Part IV. A new dinosaur, Gallimimus birllarus n. gen.. n . sp.
(Ornithomimidae) from the Upper Cretaceous of Mongolia. Palaeont. pol., 27: 103-43.
OSTKOM, J. H.,1969. Osteology of Deinonychus antirrhopus, an unusual theropod from the Lower
Cretaceous of Montana. Bull. Yale Peabody Mus. nat. Hist.. 30: 1-165.
OSTROM, J. H., 1970a. Stratigraphy and paleontology of the Cloverly Formation (Lower Cretaceous) of
the Bighorn Basin area, Wyoming and Montana. Bull. Yale Peabody Mus. nat. Hist., 35: 1-234.
OSTROM, J . H., 1970b. Archaeopteryx: Notice of a "new" specimen. Science, 170: 537-8.
OSTROM, J . H.. 1972. Description of the Archueopteryx specimen in the Teyler Museum, Haarlem. Proc.
Sect. Sci. ned. Akad. Wet. ( B ) # 75: 289-305.
OSTKOM. J . H., 1973. The ancestry of birds. Nature. Lond.. 242: 136.
OSTROM, J . H., 1974a. Archaeopteryx and the origin of flight. Q. Rev. Biol.. 49: 27-47.
OSTROM. J. H., 1974b. The pectoral girdle and forelimb function of Deinonychrrs (Keptilia: Saurischia):
A correction. Yale Peabody Mus. nat. Hist.Postilla, 165: 1-11.
OSTROM, J. H., 1975a. The origin of birds. In F. A. Donath (Ed.), Ann. Rev. Earth Planet. Sci., 3:
55-57.
OSTROM, J. H., 1975b. On the origin of Archaeopteryx and the ancestry of birds. Proc. Centre Not.
Rech. Sci., Colloq. Internat. 218 P r o b l h e s Actuels de Palkontologie-Evolution des Vertkbrks.
51 9- 5 3 2.
OSTROM, J. H., 1976. Some hypothetical anatomical stages in the evolution of avian flight. Smithson.
Conm'. Paleob.
OSTROM, J . H.. In press. On a new specimen of the Lower Cretaceous theropod dinosaur Deinonychus
antirrhopus. Breviora
OWEN, R.. 1862. On the fossil remains of a long-tailed bird (Archaeopteryx macrurus Ow.) from the
lithographic slate of Solenhofen. Proc. R . Soc., 12: 272-3.
OWEN, R.. 1863. On the Archaeopteryx of von Meyer, with a description of the fossil remains of a
long-tailed species from the lithographic stone of Solenhofen. Phil. Trans. R. SOC., 153: 33-47.
PARKER, W. K., 1887. On the morphology of birds. Proc. R . SOC.,42: 52-8.
PARKS, W. A.. 1928. Stnrthiornimus samueli. a new species of Ornithomimidae from the Belly River
Formation of Alberta. Univ. Toronto Stud. geol. Ser.. 26: 1-24.
PETKONIEVICS, B.. 1923. Uber das Ulnare im Carpus der Berliner Archaeornis. Zenrbl. Miner. Geol.
Palaont.. 1923: 94-5.
PETRONIEVICS. B., 1925. Ueber die Berliner Archaeornis. Beitrag zur Osteologie der Archaeornithese.
Ann. Geol. Pen. balkan. 8: 37-87.
PETTINGILL, 0. S.. 1970. Ornithology in laboratory and field: 524 pp- Minneapolis: Burgess.
PIVETEAU. J., 1950. Origine et Cvolution des Oiseaux. In P. P. Grass6 (Ed.), Daitk de Zoologie, X V ,
Oiseaux: 792-81 1. Paris: Masson et Cie.
PIVETEAU, J., 1955. Archaeornithes. In J. Piveteau (Ed), Traitk d e Palkonrologie. V: 1010-36. Paris:
Masson et Cie.
RAATH. M. A., 1969. A new coelurosaurian dinosaur from the Forest Sandstone of Rhodesia. Mat. Mus.
Rhodesia A rnoldia, 28: 1-2 5.
REED, C. A., 1960. Polyphyletic o r monophyletic ancestry of mammals, or: What is a class? Evolution,
14: 314-22.
ROMER, A. S.. 1956. Osreology of the reptiles: 772 pp. Chicago: Univ. Chicago Press.
ROMER, A. S.. 1966. Vertebrate paleontology. 3rd ed.; 468 pp. Chicago: Univ. Chicago Press.
178 J. 11. OSTROM

ROMIIR, A. S. 1968. The procession of life: 3 2 3 pp. Cleveland & New York: World.
ROMEK, A. S . , 1971. The Chafiares (Argentina) Triassic reptile fauna X. Two new hut incompletely
known long-limbed pseudosuchians. Breviora. 3 7 8 : 1-10.
KOMER, A. S., 1972a. The Chaiiares (Argentina) Triassic reptile fauna XIII. An early ornithosuchitl
pseudosuchian, Gracilisuchus stipanicicontm, gen. e t sp. nov. Breviora. 389: 1-24.
ROMER, A. S., 1972b. The Chaiiares (Argentina) Triassic reptile fauna XV. Further remains of the
thecodonts Lagerpeton and Lagosuchus. Ereviora, 3 9 4 : 1-7.
ROMIIR. A. S.. 1972c. The Chaiiares (Argentina) Triassic reptile fauna XVI. Thecodont classification.
Breviora, 395: 1-24.
RUSSELL, D. A.. 1969. A new specimen of Stenonychosaurus from the Olilman Formation (Cretaceous)
of Alberta. Can. J . Earth Sci.. 6: 595-612.
KUSSKLL, D. A., 1972. Ostrich dinosaurs from the Late Cretaceous of Western Canada. Can. J. Earth
Sci., 9: 3 7 5 -402.
SCIILOTliEIM. E. F. VON. 1820. Die Petrefactenkrrnde: 437 pp. Gotha: Becker’shen Buchandlung.
SEELEY, H. C..1881. Prof. Carl Vogt on the Archaeopteryx. Ceol. Mag. ( 2 ) 8 : 300-9.
SHAHOV, A. C . . 1970. An unusual reptile from the Lower Triassic of Fergana. Puleont. J., 1: 112-6.
SIMONETTA. A. M.. 1960. On the mechanical, implications of the avian skull and their bearing o n the
evolution and classification of birds. Q. Rev. Eiol.. 35: 206-20.
SIMPSON, G . G., 1926. The fauna of quarry 9. A m . J. Sci.. ( 5 ) 1 2 : 1-1 1.
SIMI’SON. G. G . , 1946. Fossil penguins. Bull. A m Mus. nar. Hist.. 87: 1-95.
SIMPSON. G . C.,1960. Diagnosis of the classes Keptilia and Mammalia. Evolution. 14: 388-92.
SIMPSON, G . G . , 1961. Principles of animal taxonomy: 247 pp. New York: Columhia Univ. Press.
SWINTON, W. E., 1958. Fossil birds: 63 pp. London: Br. Mus. (Nat. Hist.).
SWINTON. W. E.. 1960. The origin of birds. In A. J . Marshall (Kd.), Biology and comparative physiology
of birds: I: 1-14. London & New York: Academic Press.
SWINTON, W. E., 1964. Origin of birds. In A. I.. l’homson (Ed.), A new dictionary of birds: 559-62.
h n d o n : Nelson.
THULBOKN. R. A,. 1975. Dinosaur polyphyly and the classification of archosaurs and birds. Aust. J.
ZOO/., 2 3 : 249-70.
TUCKER, B. W., 1938a. Some observations on Dr Lowe’s theory of the relationship of the struthiones to
the dinosaurs and to other birds. Proc. 8 t h In?. orn. Congr.. Oxford, 1 9 3 4 : 222-4.
TUCKER, B. W., 1938b. Functional evolutionary morphology: The origin of birds. In G. R. de Beer
(Ed.), Evolution: 321-36. Oxford: Clarendon Press.
VAN TYNE. J. & BERGER. A. J., 1959. Fundumentals of ornithology: 624 pp. New York: Wiley.
VERSLUYS, J., 1910. Streptostylie bei Dinosauriern. nebst Bemerkungen iiber die Verwandtschaft der
Vogel und Dinosaurier. Anat. Anz.. 30: 175-260.
VERSI.UYS, J.. 1912. Das Streptostylie-Problem und die Bewegungen im Schadel bei Sauropsiden. Zool.
Jb. Sttppl. 1.5, 2: 545-716.
VOGT. C.. 1879. Archaeopteryx, ein Zwischenglied zwischen den Vogeln und Reptilien. Naturforscher,
Berlin. 1 8 7 9 : 401-4.
VOGT. C., 1880. Archaeopteryx macroura. an intermediate form between birds and reptiles. Ibis. ( 4 ) 4 :
434-56.
WAGNER, J . A,, 1861. Ueber ein neuea, augenhlich mit Vogelfedern versehenes Reptil aus dem
Solenhofener lithographischen Schiefer. Sits. bayer. A kad. Wiss.. 2: 146-54.
WALKER, A. D., 1961. Triassic reptiles from the Elgin area: Stagonolepis. Dasygnathus and their allies.
Phil. Trans. R . SOC. (6)-244: 10204.
WALKER. A. D.. 1964. Triassic reptiles from the Elgin area: Ornithosuchus and the origin of carnosaurs.
Phil. Trans. R . Soc. ( E l 9248: 53-134.
WALKER, A. D.. 1970. A revision of the Jurassic reptile Hallopus victor (Marsh), with remarks on the
classification of crocodiles. Phil. Trans. R . SOC. (6 , 257: 323-372.
WALKER, A. D.. 1972. New light on the origin of birds and crocodiles. Nature, 2 3 7 : 257-63.
WALKER, A. D., 1974. Evolution, organic. In D. N. Lapedes (Ed.), Yearbook of Science and Technology
1 9 7 4 : 177-9. New York: McCraw-Hill.
WELLNHOFER, P., 1974. Das funfte Skelettexamplar von Archaeopteryx. Palaeontographica (Abt. A ) ,
147: 169-216.
WELTY, J . C., 1962. T h e life of birds: 546 pp. Philadelphia: Saunders.
WHITE, T. E., 1973. Catalogue of the genera of dinosaurs. Ann. Carneg. Mus.. 44: 117-55.
WILLISTON, S. W.. 1879. “Are birds derived from dinosaurs?”. Kansas C y Rev. Sci.. 3: 457-60.
WOODWARD, A. S.. 1907. On a new dinosaurian reptile (Scleromochlus taylori gen. et sp. nov.) from the
Trias of Lossiemouth, Elgin. Q. Jl. geol. Soc. Lond., 63: 140-4.

ADDENDUM

After this study had been submitted to the Linnean Society, a copy of
Ellenberger & de Villalta’s (1974) preliminary note on Cosesaurus aviceps
 A R C H A E O P f E R Y X A N D THE ORIGIN OF BIRDS 179

reached my desk. This note reports the discovery in Middle Triassic rocks of
Spain of a tiny ( 1 5 cm length) tetrapod with what appears to be a very
bird-like, tooth-bearing skull. Ellenberger & de Villalta interpret this speci-
men as a “protobird” and suggest that it is closer to true birds than are the
specimens of Archacoptcrj~s.Not having seen the specimen, I am in no position
t o comment on that, but their published photographs and dimensions indicate
an animal about the size and proportions of Sclcromoclzlus. No feather
impressions are reported and the post-cranial skeleton appears to lack all of the
advanced characters shared by Arcllueopteryx and coelurosaurs, and shows no
derived characters of birds.

A D D E N D U M REFERENCE
ELLENBERGER, P. & de VILLALTA, J . F . , 1974. Sur la prhsence d’un ancctre probable des Oiseaux
dans le Muschelkalk sup6rieur d e Catalogne (Espagne). N o t e preliminaire. Arra Geol. Hisp.. IX, 5:
I 62-8.

APPENDIX

S.tstcrriatic list o f tuxa

For the convenience of those readers who may not be familiar with the
systematic assignment of the various taxa referred to throughout this paper, the
following listing is provided, together with the age and provenance of each
taxon.

CLASS AVES

Subclass Archaeornithes (Saururae)

Order Archaeopterygiformes
Family Archaeopterygidae
Archaeopteryx litlrogruptiica (=Arcfiaeoriiis siernensi); Late
Jurassic, Europe.
Subclass Neornithes
Order Hesperornithiformes
Family Hesperornithidae
Hesperornis; Late Cretaceous, North America.
Order Ichthyornithiformes
Family I ch th y or ni thid ae
Ictithyornis; Late Cretaceous, North America.
AVES?: INCERTAE S E D l S
Luopteryx priscus; Late Jurassic, North America.
CLASS REPTILIA

Subclass Archosauria
Order Thecodontia
Suborder Proterosuchia
Family Proterochampsidae
Cerritosuurus; Middle Triassic, South America.
Family Erythrosuchidae
Erythrosuchus; Early Triassic, South Africa.
I80 J. H. OSTROM

Suborder Pseudosuchia
Family Euparkeriidae
Euparkeria; Early Triassic, South Africa.
Family Ornithosuchidae
Gracilisuchus; Middle Triassic, South America.
Ornithosuchus; Late Triassic, Europe.
Riojastichus; Late Triassic, South America.
Venaticosuchus; Late Triassic, South America.
Family Pres tosuchid ae
Mandasuchus; Middle Triassic, East Africa.
Prestosuchus; Middle Triassic, South America.
Saurosttchrrs; Middle-Late Triassic, South America.
Ticinasuchus; Middle Triassic, Europe.
Family S cler omochlidae
Lagerpeton; Middle Triassic, South America.
Scleromochlus; Late Triassic, Europe.
Pseudosuchia Incertae Sedis
Hesperosuchus; Late Triassic, North America.
Lagosuchus; Middle Triassic, South America.
Lewisttchus; Middle Triassic, South America.
Rauisuchus; Middle Triassic, South America.
Saltoposuchus; Late Triassic, Europe.
Triassolestes; Middle-Late Triassic, South America.
Suborder Aetosauria
Family Aetosauridae
Stagonolepis; Late Triassic, Europe.
Thecodontia Incertae Sedis
Longisyuama; Early Triassic, Eurasia.
Order Crocodilia
Suborder Protosuchia
Family Notochampsidae (Protosuchidae)
Erythrochampsa; Late Triassic, South Africa.
Notochampsa; Late Triassic, South Africa.
Family Pedeticosauridae (Sphenosuchidae)
Hemiprotosuchus; Late Triassic, South America.
Pedeticosaurus; Late Triassic, South Africa.
Sphenosuchus; Late Triassic, South Africa.
Protosuchia Incertae Sedis
Pseudhesperosuchus; Late Triassic, South America.
Order Ornithischia
Suborder Ornithopoda
Family Hypsilophodontidae
Hypsilophodon; Early Cretaceous, Europe.
Laosaurus; Late Jurassic, North America.
Thescelosaums; Late Cretaceous, North America.
Family Iguanodontidae
Camptosaurus; Late Jurassic, North America.
Tenontosaunts; Early Cretaceous, North America.
 ARCHAEOPTER Y X AND T H E ORIGIN O F BIRDS 1 ni

Suborder Pachycephalosauria
Family Pachycephalosauridae
Hornalocephale; Late Cretaceous, Asia.
Stegoceras (=Troodon);Late Cretaceous, North America.
Suborder Stegosauria
Family Stegosauridae
Scelidosaurus; Early Jurassic, Europe.
Stegosaurus; Late Jurassic, North America.
Order Saurischia
Suborder Theropoda
lnfraorder Coelurosauria
Family Procompsognathidae
Coelophysis; Late Triassic, North America.
Cornpsognathus; Late Jurassic, Europe.
Procornpsognathrrs; Late Triassic, Europe.
Syntarsus; Late Triassic, Africa.
Family Segisauridae
Segisaurus; Late Triassic, North America.
Family Coeluridae
Coelitrrts; Late Jurassic, North America.
Microvenutor; Early Cretaceous, North America.
Ornitholestes; Late Jurassic, North America.
Family Dromaeosauridae
Chirostenotes; Late Cretaceous, North America.
Deinonychus; Early Cretaceous, North America.
Dromueosuurus; Late Cretaceous, North America.
Saurornithoides; Late Cretaceous, Asid.
Stenonychosaurus; Late Cretaceous, North America.
Velociraptor; Late Cretaceous, Asia.
Family Ornithomimidae
Archaeornithornimus; Late Cretaceous, Asia.
Deinocheirus; Late Cretaceous, Asia.
Drorniceiornirnus; Late Cretaceous, North America.
Gallimirnus; Late Cretaceous, Asia.
Ornithornirnus; Early-Late Cretaceous, North American and Asia.
Oviraptor; Late Cretaceous, Asia. (Oviraptor may not be an
omithomimid, but it is a coelurosaur. See Russell, 1972.)
Struthiornirnus; Late Cretaceous, North America and Asia.
Infraorder Carnosauria
Family Megalosauridae
Allosaurus; Late Jurassic, North America.
Cerutosaurus; Late Jurassic, North America.
Family Tyrannosauridae (Deinodontidae)
Albertosaurus (=Gorgosaurus) Late Cretaceous, North America.
Daspietosuurus; Late Cretaceous, North America.
Tarbosaurus; Late Cretaceous, Asia.
Tyrannosaurus; Late Cretaceous, North America.
Carnosauria Incertae Sedis
Zunclodon; Late Triassic, Europe.
182 J . It. OSTROM

Suborder Sauropodomorph a
Infraorder Prosauropoda
Family Anchisauridae (Thecodontosauridae)
Ainphisatrrus (=Anchisaurus); Late Triassic, North America.