Sigma Xi, The Scientific Research Society

Bird Flight: How Did It Begin? Did birds begin to fly "from the trees down" or "from the
ground up"? Reexamination of Archaeopteryx adds plausibility to an "up from the
ground" origin of avian flight
Author(s): John H. Ostrom
Source: American Scientist, Vol. 67, No. 1 (January-February 1979), pp. 46-56
Published by: Sigma Xi, The Scientific Research Society
Stable URL: http://www.jstor.org/stable/27849060
Accessed: 31-07-2016 14:55 UTC

REFERENCES
Linked references are available on JSTOR for this article:
http://www.jstor.org/stable/27849060?seq=1&cid=pdf-reference#references_tab_contents
You may need to log in to JSTOR to access the linked references.

Your use of the JSTOR archive indicates your acceptance of the Terms & Conditions of Use, available at
http://about.jstor.org/terms

JSTOR is a not-for-profit service that helps scholars, researchers, and students discover, use, and build upon a wide range of content in a trusted
digital archive. We use information technology and tools to increase productivity and facilitate new forms of scholarship. For more information about
JSTOR, please contact support@jstor.org.

Sigma Xi, The Scientific Research Society is collaborating with JSTOR to digitize, preserve and extend
access to American Scientist

This content downloaded from 202.3.77.249 on Sun, 31 Jul 2016 14:55:59 UTC
All use subject to http://about.jstor.org/terms
 John h. Ostrom Bird Flight: How Did It Begin?
Did birds begin to fly "from the trees down" or "from
the ground up"? Reexamination of Archaeopteryx
adds plausibility to an "up from the ground" origin
of avian flight

Paleontologie data have provided birds and the intriguing question of Previous speculations on this ques
unique clues about various events in how bird flight might have evolved. tion have produced two quite differ
ent scenarios. Stated very simply,
the history of life that we could not Flight is one of the most remarkable
otherwise have known about?for of all animal adaptations, yet it has these are that birds began to fly "from
example, the question of human ori evolved numerous times quite inde the trees down"?or "from the
gins, the ancestry of mammals, and pendently?more than half a dozen
ground up." The first is the widely
the origins of land vertebrates. It may times in vertebrate animals alone.
favored and very logical "arboreal
be true that the exact lines of descent Flight in vertebrates ranges from
theory," first suggested by O. C.
in these particular examples (and passive (non-self-powered) para Marsh in 1880 and skillfully elabo
other lineages) are not known pre chuting (flying squirrels, flying frogs) rated recently by Walter Bock (1965,
cisely, but few would disagree that the and gliding (the colugo and flying 1969). The second is the often ridi
following general evolutionary lizards) to fully self-powered flight of culed and seemingly less probable
histories are widely accepted as true: birds, bats, and probably the extinct "cursorial theory," commonly at
mankind evolved from a late Tertiary flying reptiles known as pterosaurs. tributed to Baron Francis Nopcsa
apelike creature, probably by way of The most improbable flight is that of (1907, 1923), but originally outlined
the australopithecines; mammals the flying snake, Chrysopelea, but it in 1879 by Samuel Wendell Willis
originated from advanced cynodonts is the varied forms of avian flight that ton.
of the long-extinct mammal-like have fascinated mankind since an
reptilian order, Therapsida; and land cient times, as is evidenced by Egyp
vertebrates arose somewhat more tian and Greek mythology. The di The arboreal theory
than 350 million years ago from a verse ways in which birds fly have In brief, the arboreal theory postu
now-extinct group of fishes, the rhi been revealed in the studies by Brown lates that the primitive ancestors of
(1948, 1951, 1953, 1961, 1963), birds must have been tree-dwelling
pidistian crossopterygians. These are
but a few of many evolutionary Greenewalt (1960a, 1960b, 1962, animals which developed increasing
histories that have been inferred from 1975), Hartman (1961), Pennycuick need and skill to leap from branch to
the fossil record. (1960, 1968, 1971, 1975), Tucker branch, and then from tree to tree.
(1969,1973), and others, to the extent The selective advantage of any ana
One such historical narrative deduced that the aerodynamics of bird flight tomical change that would increase
from the fossil record is the origin of is fairly well understood. But how it body surface area relative to body
all began is not so clear. mass is obvious?first, to slow the
rate of fall from a mis-leap; later, to
A century ago, it was supposed by provide lift and directional descent in
some (e.g. Mivart 1871) that bird longer and longer glides; and finally,
John H. Ostrom is Professor of Geology at Yale flight came into being by a sudden to permit longer and longer gliding
University and Curator of Vertebrate spontaneous appearance of fully de leaps between trees. As Marsh ex
Paleontology at Yale's Peabody Museum of veloped wings, where previously there pressed it in 1880:
Natural History. After receiving his Ph.D.
from Columbia University, he taught first at
had been only normal forelegs for
Brooklyn College and then at Beloit College walking. Today, most of us are confi The power of flight probably originated
before joining the faculty at Yale in 1961. His dent that such a major and complex among small arboreal forms of reptilian
research has centered on the history and ev innovation as flight developed birds. How this may have commenced, we
olution ofMesozoic reptiles, chiefly dinosaurs through a sequence of many small have an indication in the flight of Galeo
and flying reptiles, and on the origins of birds
and bird flight. The author is indebted to
changes that occurred (however pithecus [colugo], the flying squirrels
(Pteromys), the flying lizard (Draco) and
many students and colleagues for critical and slowly or rapidly) over many genera
the flying tree frog (Rhacophorus). In the
lively discussions on the subject of this article, tions, with every stage being well early arboreal birds, which jumped from
but only the author should be held responsible adapted for its own particular setting branch to branch, even rudimentary
for the speculations set forth. Photographs
are by the author unless indicated otherwise.
and way of life. Assuming this latter feathers on the forelimbs would be an
Address: Department of Geology and Geo mode to be correct, what can we say advantage as they would tend to lengthen
physics, Yale University, New Haven, CT about the origins of bird flight? And a downward leap or break the force of a
06520. what is the evidence? fall.

46 American Scientist, Volume 67

This content downloaded from 202.3.77.249 on Sun, 31 Jul 2016 14:55:59 UTC
All use subject to http://about.jstor.org/terms
The flaw in this picture lies with the
analogs chosen by Marsh. They are all
quadrupedal rather than bipedal
arborealists (see Fig. 1), and with the
exception of Draco (where a fold of
skin, the patagium, is supported by
elongated ribs), the flight membrane
is stretched between the fore and
hind limbs. I will return to this point
later.

In 1965, Walter Bock presented a

more detailed version of the arboreal
theory as a model to show that ma
croevolutionary differences between
members of a phyletic lineage were
not necessarily incompatible with 41
known principles of microevolu
tionary change. Listing only the
"major" hypothetical adaptive stages
in their order of occurrence?(1) an
cestral ground-dwelling quadrupedal
reptile, (2) bipedal ground-dweller,
(3) bipedal and arboreal life, (4)
leaping between trees, (5) parachut
ing, (6) gliding, (7) active, powered
flight?Bock made it clear that he
visualized many intermediate grad
ational, but fully adapted, stages be
tween each of these. This sequence
'9

culminated in the macroevolutionary

transition from a reptilian nonflier to
powered avian flight.

The major significance of Bock's

paper was to show that at each stage
Figure 1. Artist Zdenek Burian has recon tice that Burian visualized this transitional
there would be different, perhaps structed a pre-Archaeopteryx stage (com stage as quadrupedal, rather than bipedal,
unique, selective pressures inducing monly termed "pro-avis") in the evolution of following the prevailing view at the time that
still further changes subject to yet avian flight. No fossil evidence exists of any bipedality in birds arose as an adaptation to
other selective forces. In his model, pro-avis. It is a purely hypothetical pre-bird, arboreal life. (From Augusta and Burian 1961,
but one that must have existed?whether or with the permission of Artia Publishers,
flight would be selected for only in the not it resembled the creature shown here. No Prague.)
final stages. The critical point of this
model, though, is that, in order to fly,
the animal first had to be able to
climb. However, considering the de "thrust" surfaces. Flapping action of by loss of traction. A more critical
sign of modern birds, together with these "proto-wings" supposedly flaw, however, is the miniscule
that of the oldest-known bird, Ar added thrust to that provided by the amount of additional thrust that
chaeopteryx, that skill may not have hindlimbs, resulting in greater ac could have been generated by those
been part of the repertoire of primi celeration and faster running speed. earliest enlarged "scales" on the in
tive birds, or even of bird ancestors. I Ultimately, this is presumed to have cipient "wing" pushing against the
will return to this point also. led to flight velocities and to at least air. Certainly this could not have been
partial conversion of the forelimbs anywhere near enough additional
from "propellers" into wings. thrust to produce a measurable in
The cursorial theory crease in running speed, and thus be
The cursorial theory postulates a se One of the key criticisms that has selected for.
quence of stages from a primitive been leveled at this hypothesis is that,
quadrupedal reptile, to a facultative once the animal is airborne, the main The cursorial theory of bird flight
biped (like the lizards Basiliscus and thrust source (i.e. traction of the hind origins has received virtually no ac
Crotaphytus, which at high speeds feet against the ground) would be lost ceptance, apparently for several verf
run on their hindlegs, but at slower and velocity would diminish. How good reasons: Nopcsa's seemingly
speeds move on all fours), to an ever, at that primitive "flight" stage absurd flailing-arm propellers; the
obligatory cursorial biped, followed there might well have been selective seemingly impossible "bootstrap"
by stages of elongation of the fore advantage in lifting up off the ground, effort required for the animal to lift
limbs and enlargement of "scales" on at least momentarily?either for es itself by means of flapping proto
the arms to increase their surface cape, or in pursuit of prey?that wings; the simple logic of cheap
area, thereby forming ever larger outweighed any disadvantage caused gravity-powered flight from an ele

1979 January-February 47

This content downloaded from 202.3.77.249 on Sun, 31 Jul 2016 14:55:59 UTC
All use subject to http://about.jstor.org/terms
 x. .

*0 -0 --- 0 70 :

20~ ~~~ 30 40dr60 70

Figure 2. The fossil imprint (left) of a solitary
feather that has been referred to Archaeop
teryx, and represents the original discovery of
Jurassic birds, is compared with a modern
contour (vaned) feather (center). This fossil
imprint was found on a slab of the famed
Solnhofen lithographic limestone in a Bavarian
stone quarry in 1861. It clearly shows the uni
formly parallel alignment (except at two places and its counterpart is in the Bayerische
near the tip) of numerous barbs, which indi Staatssammlung f?r Pal?ontologie und histo
cates that these barbs bore minute barbules rische Geologie in Munich. (Electron micro
with microscopic hooklets, just as is shown in graph courtesy of A. Pooley; scale bar equals
the scanning electron micrograph of a modern 10 microns. Modern feather photographed by
contour feather (right). The fossil imprint W. Sacco; scale units are in millimeters.)
shown here is in the collections of the Hum
boldt Museum f?r Naturkunde in East Berlin,

vated perch; and the fact that nearly Birds, on the other hand, are skilled served in the feather impressions
all present-day vertebrate fliers (ex bipeds?some of them extraordinar surrounding two of the skeletons in
cept flying fish, of course) are arbo ily so (take the roadrunner, for ex dicate the existence of microscopic
real. Needless to say, some bats and ample). It seems to me that only one feather construction essentially
many birds are not now arboreal, but conclusion is possible: avian flight identical to that of modern feathers.
these may be presumed to be rela evolved by a very different sequence Add to this the arrangement of large
tively recent adaptations for habits of adaptive stages than did that of all contour-type feathers along the arms
and habitats quite different from other fliers. An arboreal origin of of those two specimens (the London
those of earlier arboreal ancestors. flight appears to be the only possible and Berlin specimens) in a pattern
explanation for quadrupedal fliers, very much like that of modern birds'
I noted above that Marsh's analogs and for Draco and the flying snakes, wings. These suggest at least some
were all quadrupeds, and implied that but it may not have been the way bird degree of flight capability?per
they might not be appropriate models flight came about. haps.
upon which to base speculations
about bipedal avian flight origins. It What is the evidence? Unlike the feathers, however, the
is important to note that all nonavian skeletal anatomy of Archaeopteryx is
vertebrate fliers, except flying snakes not so birdlike as we might expect. In
and fish (and possibly pterosaurs), are
Archaeopteryx fact, there is just one skeletal feature
quadrupedal walkers in which all four The anatomical differences among of Archaeopteryx that is exclusively
limbs are involved in the flight ap modern vertebrate fliers point up the avian?the wishbone, or furcula
paratus. Birds are unique. They are probably very different scenarios that (fused clavicles). All other features of
the only vertebrate fliers in which the led to the different forms of verte the skeleton are also found in various
flight apparatus is constructed en brate flight, but they do not provide small carnivorous dinosaurs (coelu
tirely of the forequarters (and the a clear picture of how bird flight rosaurian theropods). It is this dino
tail) and does not involve the hind might have begun. The answer may saurian nature of Archaeopteryx that
limbs at all. By a unique sequence of be indicated, though, in the fossil re led me to theorize that modern birds,
circumstances, birds evolved inde mains of the oldest-known bird? perhaps by way of Archaeopteryx,
pendent dual locomotory systems?Archaeopteryx?from the 150-mil are the direct lineal descendants of
wings for flight and legs for running, iion-year-old Jurassic Solnhofen the small carnivorous dinosaurs (Os
walking, or swimming?and this may limestones of Bavaria, in West Ger trom 1973,1975,1976a).
be the chief reason for their remark many. Although possibly not directly
able success. ancestral to any modern birds, these This is not the main point of this ar
specimens constitute the only direct ticle, however. These striking ana
It is possible that the extinct flying evidence available pertaining to the tomical similarities perhaps indicate
reptiles (pterosaurs) also had wings earliest stages of bird evolution. At much more than a close evolutionary
constructed only of the forelimbs, but the present time a total of only six relationship between coelurosaurs
their hind legs were not designed for specimens are known: five partial to and Archaeopteryx. They also
bipedal walking. It appears that, as virtually complete skeletons and an suggest very similar adaptations, and
with nearly all bats, excursions by isolated imprint of a solitary feather thus similar modes of life. This may
most pterosaurs across the ground that is presumed to have come from be the most important clue about the
surface must have been awkward and the same kind of animal. That feather early stages of bird evolution and the
required the use of the wings as legs. imprint (Fig. 2) plus the details pre origins of avian flight.

48 American Scientist, Volume 67

This content downloaded from 202.3.77.249 on Sun, 31 Jul 2016 14:55:59 UTC
All use subject to http://about.jstor.org/terms
 A

S-4...v ........ .... ... V.

qw
M Al vit

. . . .......

L 7,

'A' . ... ...

O.M.,

A Fa- :21,P 7.0i - 2,

,: -:. :: . . j
... ...... .. -'i . .. . . ... -; .m-Z ', . . . ... ...
I.N.A . . . ... . 'A" 5 e. g.
On
OR 68U; j .."r.. ,
P - -l'.m, , !, -.1--lIx.e.
-!
mi No

Or
.... ... ..
-xz --J
AMAZE

vs

Jr V.

A m

T. id Dr, L 7
ML M
0'. M.

MW
vi. ex

T l .. . .. .....
i-Y

W.

W Pea -
-TS

11 a .4.udt*ff
W.. M

-A

4
q. 1% Gm. Y. X-,! -
I'D OiR 'I
IVC

m M<6 -1 -71'

Tifl 44'

-k

iv.

Figure 3. The Berlin specimen of Archaeop Upon first examination of the speci authors have concluded. But closer
teryx lithographica (some would assign it to a
separate species, even genus, Archaeornis mens of Archaeopteryx?especially inspection reveals some puzzling?
siemensi), from the Solnhofen limestone, of the famed Berlin specimen (Fig. even paradoxical?conditions which
shows the modernlike arrangement of the 3)?we see what appears to be a throw that conclusion into doubt, at
"flight" feathers in an elliptical-shaped "wing." birdlike creature with long arms and least in terms of powered flight.
This individual was about the size of a common
pigeon. The most remarkable of the six known
clear imprints of feathered "wings"
specimens of Archaeopterxy, it was found in remarkably like those of modern First of all, there is no sternum
1877 and now is in the collections of the Hum birds. Surely these "wings" must have (breast bone) preserved in any of the
boldt Museum. been for flight, as nearly all previous specimens. Modern flying birds have

1979 January-February 49

This content downloaded from 202.3.77.249 on Sun, 31 Jul 2016 14:55:59 UTC
All use subject to http://about.jstor.org/terms
a large well-ossified sternum,-which that are prominently developed in
usually has a prominant deep mid modern flying birds and that are the
line keel that serves as the main site sites of attachment of the special
of attachment of the large flight muscles that fold the wing compactly
muscles. Presumably, a sternum was against the back and flanks. This
present in Archaeopteryx, at least in suggests that Archaeopteryx proba
a cartilaginous state, but there is no bly was not able to fold the wing
evidence of that. What does this in tightly against the body?a presum
dicate about the size and power of the ably important action for protecting
flight muscles? Perhaps nothing. the "wing" (if it actually was an air
After all, in bats the sternum is quite foil) when not in use.
small and often lacks a keel. But the
ventral flight muscle (M. pectoralis) All the skeletal f e?tures just described
of bats is small, comprising less than play critical roles in the action of
10% of total body weight, as compared avian wings, both during flight and
with birds, in which it typically equals when not in use. Their absence in
15% to 20% of body weight and ranges Archaeopteryx is clear evidence that
to an astonishing 33% in the dove despite the modern "winglike" ap
Leptotila (Hartman 1961). Thus, the pearance of the feather arrangement,
absence of a sternum in Archaeop the "flight" apparatus of this ancient
teryx seems important?and perhaps bird was far from fully evolved. In
indicates weak pectoral muscles. fact, all these features as developed in
Archaeopteryx are essentially the
Another surprising feature of the same as in certain small carnivorous
skeletal "flight apparatus" of Ar dinosaurs, such as Ornitholestes,
chaeopteryx is the shape and thin Velociraptor, and Deinonychus.
sheetlike construction of the cora
coids?the robust strutlike bones in Figure 4. The left feet of the London (top) and There is one curious exception,
modern birds that brace the shoulder Berlin (center) specimens of Archaeopteryx though. Two of the specimens of Ar
show the reversed hallux and the birdlike form
against the breast bone. This coracoid chaeopteryx (the London specimen
similar to that of a modern, pigeon (bottom).
brace is critical in flying birds, be The same foot design is also characteristic of and the Maxberg specimen) feature
cause it immobilizes the shoulder some theropod dinosaurs. (Bottom photo by W. a robust boomerang-shaped bone that
socket so that the full contractile Sacco.) almost certainly is a furcula, or
force of the flight muscles is applied wishbone. Believed to be formed by
to the power stroke of the wing, and fusion of the paired clavicles, the
there is no loss of muscular force by furcula is unique to birds and gener
downward displacement of the anomalies. The three fingers are not ally is interpreted as a specialized
shoulder. In Archaeopteryx, the co fused together, as they are in all feature of the avian flight apparatus,
racoids are thin, half-moon-shaped modern birds?a condition that is possibly acting as a springlike spacer
sheets of bone?short in length and generally thought to be an adaptation to counteract the transverse com
not at all robust?fragile "braces" (?) related to the need for a solid skeletal pressive component of the force pro
between the shoulder and the ster platform for the firm attachment of duced by contraction of the flight
num. This too suggests that the pec the primary flight feathers. Also, the muscles, which otherwise would tend
toral region was subject to relatively wrist of Archaeopteryx consists of to squeeze the shoulder sockets
weak stresses resulting from relatively several separate small bones and toward each other. If Archaeopteryx
small "flight" muscles. clearly had much more articular mo possessed a furcula, then one is
bility than is found in modern birds, tempted to conclude that it had
Even more interesting is the absence whose wrists permit flexion and ex powerful pectoral muscles and
of the triosseal canal (Ostrom 1976b), tension only in the plane of the wing, therefore it surely must have flown.
the pulleylike structure of the shoul perpendicular to the wing stroke. The "wing" outline seems to add
der skeleton in modern birds that That would seem to be a major me strong support to that conclusion, as
reverses the action of the supracora chanical deficiency in the "wing" of Saville (1957), de Beer (1975), and
coideus muscle so that it powers the Archaeopteryx. others have noted. But what about all
wing recovery stroke, rather than the above-mentioned deficiencies?
adding to the downward power The upper arm 'bone (hum?rus) of
stroke. Without that canal, this Archaeopteryx has a very large del The general consensus seems to be
muscle could not possibly have func topectoral crest, the site of attach that Archaeopteryx probably was at
tioned to elevate the wing in Ar ment of the pectoralis muscle. That least a glider, but no more than a
chaeopteryx, and the recovery stroke would seem to indicate a fairly large feeble, flapping flier?// it flapped at
must have been powered by the rela muscle mass?possibly for flight. But all (Bock 1965,1969; Yalden 1970). In
tively weak and mechanically less this same crest, of comparable rela any case, it is clear that Archaeop
efficient dorsal muscles, such as the tive size, exists in all theropod dino teryx was a long way from modern
deltoids. saurs, which also lack ossified breast birds as far as its flight equipment
bones and obviously were not fliers. was concerned.
Examination of the "wing" skeleton On the other hand, the hum?rus lacks
of Archaeopteryx reveals further all the other processes and tubercles Another very interesting aspect of
50 American Scientist, Volume 67

This content downloaded from 202.3.77.249 on Sun, 31 Jul 2016 14:55:59 UTC
All use subject to http://about.jstor.org/terms
Archaeopteryx is in the construction
of the hindquarters. Although the
pelvis was not co-ossified into a large
synsacrum as in modern birds (does
that indicate that it lacked the LN

shock-absorbing capacity of the Ur

modern avian landing gear?), the

hind legs and feet are much more
OF
similar to those of living birds than is
the forelimb or shoulder. There can
be no question that this animal was A -1

an obligatory biped with a posture

and gait very much like that of to
day's birds, but the hindlimb was not i V%4,:%."'l
1. QK
yet fully modern.

The foot consisted of three main toes

in a near-symmetrical arrangement,
with the median toe (the third) the
longest and directed forward, almost
exactly as in modern birds. The first
toe, or hallux, was short and placed
behind the second in a fully opposa
ble position, again as in modern birds.
But the metatarsals consisted of three
main bones, rather than a single co -..AS

ossified tarsometatarsus. However,

these three metatarsals were fused
together at their upper ends in one
specimen (the Maxberg specimen)
and appear to have been similarly
fused in all the other specimens since . lkl

these bones are not separated from

each other in any specimen of Ar
chaeopteryx (Fig. 4). That would
seem to be a clear indication of an 1.

AL.

early trend, as far back as Archaeop

teryx, toward the modern avian state
of metatarsal co-ossification.

The ankle itself was mechanically

quite close to that of living birds, with
a simple hingelike joint. Flexion and
extension took place at a single level,
between the distal tarsal bones, which
were fixed against the upper ends of
the metatarsals, and the proximal Figure 5. This reconstruction of Archaeopteryx number of primaries [he shows eight], the use
in an arboreal setting was painted by Rudolph
tarsals (astragalus and calcaneum), Freund of the claws on the wing, the method of feeding,
in 1965 to illustrate an article on the and the method of flying/gliding) so that sub
which fit tightly against the lower origin of feathers by K. C. Parkes (1966). No sequent investigators and illustrators will have
ends of the shin (tibia and fibula) tice the nongrasping hallux?a deliberate a target at which to shoot." (Courtesy of K. C.
bones. rendering by the artist, who stated: "I have Parkes, with permission of Carnegie Museum
tried to show all the controversial aspects of of Natural History and The Living Bird. )
The importance of these conditions is Archaeopteryx as clearly as possible (the
clear: as early as the Archaeopteryx
stage, evidence exists showing that
the functions of the hindquarters and
forequarters had evolved entirely in rial animal, fleet of foot, and probably mens that that was so. I agree that it
dependently. In fact, those specimens a ground-dweller. could have been arboreal, but the
show that at that early stage, the specimens themselves indicate oth
running apparatus was far more ad This last item is one of the major erwise. This point may seem unim
vanced (toward the modern avian points of contention among students portant, but in fact it is critical for
condition) than was the flight appa of Archaeopteryx. Contrary to almost choosing between the two hypotheses
ratus. The respective limb propor everyone else's conclusion that Ar of flight origins.
tions (femur vs. tibia vs. metatarsus) chaeopteryx must have been arboreal
also strongly suggest that Archaeop (Fig. 5), I find no compelling ana It is widely acknowledged that Ar
teryx was a highly developed curso tomical evidence in any of the speci chaeopteryx lacked virtually all of

1979 January-February 51

This content downloaded from 202.3.77.249 on Sun, 31 Jul 2016 14:55:59 UTC
All use subject to http://about.jstor.org/terms
 . .

?.:,:--:-.1N-!:.,k.'-'
-, *0 ..
N ... . :
i..r.."..
...
- -:;?.::,
.,.;. .
.......

mi.: 17.
.;., M .
Figure 6. The bony claws, or unguals, from the
feet of four of the specimens of Archaeopteryx
- ;
. !
:

are compared here with the corresponding el

:

. . m m yn.g."g - .;.!.&.;ysr. .... - N. ! !!. ;. .. .. .. . . . . . . . : .. !!!! ! ;.!;!

0 1391, .'. ,.,; Nl.'v
A-,!o: Y.. -.,F-i".
.
....,., .
'in
- -.X,
;.. , -
. i . ..
. . . ....
...:.X...ti
-:,.-:--,; L-
. . . .:
".....
- .::
..
,.....:
.11
,g-,
.. n..
; :-.,.i
... .....
.-i--1i--:r-;v.--:c1-
.. . . .
I :i...
- ..
; --,,, ";
l... .4
-A
'X.-
. , i - ..
....
,,....
,
,. , .! m
":;z5"
-:-.1!P.T, - -.4;-m
.-; .. ....
q.K..0 4.6.j.
-m i.--
:!!::::::.. :..::.
--:. i i ::::
1i ements of several types of modern birds. Each
. I -A 'r -','iS0"',-:--,
.!.A
.. .. . P .- r.." i,; .ig...X;.
'.IN ..% -.-!,.......
4. .;:;;41j..:::i!::f
ii_:,i,--.&1
3 !.I:..,Ilg;..;..'..'.
Am, ;",. -4..::.:"
R.- ; :"--,
.P.". -.j
d.
."
.:.-,; "::k:::
-
--'
..".,:L, 4;-*-SgA'-
:..16.,-, lff.:..!:L,;:,.;.
- , .,;,;..I-
.- .: .,...-..
mW-.-l:-.:-
, ..,--:lv
-'.
,_Aw.."
: -lc .
-!-, . -
...'.. , , '. ;s . Ma.. j'03;;;'
example includes the ungual of the median
-

::'IRK+.,
-m I1; R .I.Ir .- ... - .-ip" """
- .;K'i'..!:.:%?
;O.k,.:,P.
- :A.,
" .". -% -W.
:','. ..,;,;..:.:.,.:.:
-- ?,.,I
F .":R7---,
.-" l::i:i
ill I .m.
I .. ...
-ill
,.:.. .-.-
i .-mmI; ,,
.1. - ..:4'.1
,"N--&." >
,"Zik, ---,,J;
.:::.Rm
_ :' , ... ,,'Di,it
ip....,,,,,:,:!:!
: P.I. !: , U.-
I,.-,.:!;:::..k:..
...'_'.;:! o-i
.:?: 1'Nir
..-",;..;:,.
1:3i!.... -I .. ,--. ..;,.!S!::
'....
,:..:;:n:i....:.,..-O. .,.:: .. ......
,lrl;:_ ::...- .."
-
.....m , ......:?
......,-:p &.C iid'--'.-1-
..M ..
.. . ."S 1,- .. ,,:
.,
-Imv -::iff:4 Nip 1 -f.. - - 1j,:-:-: -:-yr- M -1,-,WM , x;f;e1%i-jv; W. -,!j: : . :,:., .-_ ... I-"-!F -.4. e - f::.-,;. f: 1:,p, k. ;!; .:! .---';. 5": !. !: , . .A wem_-.!

., "-.X ':'i " '...

.. M.,
-5.'.?,g!, 12M
."AfR -.4
.. . it ,.,.K
il.v. .iii -_....Pc. *.M - _..
::, I.-
. 0MV;- 1! ..: . c .ii ........ ;l."
il 1; --- .iil'r-, -.1-.1.... I ...... - .. :, :

,., K.I -11R.3p

-? .- .l.i;
-, .. -,
,;?-'O,
,',M:; -...
?il i!:
.ii;.'mi:.,.
:- ff iii Pn'ji;
i:,:!,-..
', .:;::':""-..,.i:..
.im
i::i.;.i.: -i,.1i-,.,i "n4;g-:-
:;:-.v',m, -.r.,m!m?.i-.V1
.. -- W
.- ,! R Pi- z5i:.1.
.4 f. - .;
11114 ., .,-. ,...!J,.P.-.'!' '.,,-.6.": A.F%1I ,, ', :! i.,
""!2ii --4M
11 xv!i:..Mf;.
- --_ ...ml
... . ffl .1,.,.
..-
%.q.
..
! .I
I. .1ill
..1:0
.. 11!
-,:,... S.,
.1.
..
.;r ,il
4&
.-,- v .r
. ;:-m.-.,
..
.,.,:.
.
5?R ,m . ,.. -
,.,-.
.m....'o...
. .0.01._W51,1A
. : .,: "'-
- C-
F.
;j2i:;i'-..
--.1 1; ,'..
!..,;. ...-
xt.. - .m - .-
m - .:
.. :.
.. ... w
-::ii
-i;;,,.-f-J .0! --a
!.. C.: -.
(third) toe and of the hallux. All third-toe un
".-'.,..
.,i .!!-!O
,.Nti i'l:, ii. ... Ki - .5,
..",--!i; .l .;:;:
L. ..7"i" ill"'. gm, -.,!;M A -&- : .,i::X .-: !gVi!
". ..-...4- - -mf;
:
I., ,_
-..:
.,
, ,-:-'-
-.m:Z-11 . . -m-%:.Pliii
. :.--..... :..:;.
.fp,.,ll;i-;;9--.-5g .
; ;;rdi.-.I;i:,t-.
-:k-V---::%::,: 2,-:
.,
X.....
.-.,-:,,.-- .,O
.7
.2
s.y-a;..*-.', . .ii:.-.
m
--: ,....".2.M
.m...
----.-i!S
K,:i,--:-.:.m: I,-,!:i1Lj-.,
ij:R -.-'.%-'.-.
..... j: ml:jt.::-m;-', 'I
--x.,,.-
;.' :%' F
:*.g k. -, ...,V ::::
l,O:;K..M..Fx.,i;i---
...Z..
...ia .;. ;:
.,..;M;:; li^ '. . . ..
:8.".-,i:----;
g"e.:.i, 3,; .. -.__
. .j s:f...:... :3:im!
l.-C.,:,.,: . : '-'.-'ilk'r-'::
.. '.
-- ::-il ,. ;z-.1.
--L. iii:-.i:.f. -. -:,,:,,-,,,, "-
guals are drawn to the same unit length for
" " " . - . .- jjk2;.5
.. -. . 3i..-
.W.. 'O.
- 'p, ,
._-.'l!,:i .-
-.:Ip
.. . :::::':--'L-
.. . . :.,, ,:-:; , .
:,m---.,
:-,.,9 .
.1- ..--f'.,i.
M.-
l--.k-;-- ..!A
----
-,:;.-, -P:-,.,:-:-.,--:r
--------,--'-
--.,J'.' ...
i--!Ki,
;!:V' -
rl;.1,
'.- - -_-
...." ..,. .4
. :.-_m'A
_,Zl :.C:.i. ...
, j:i,':
.1--w -, . -,
-,. q;:-:!lo?-
- -:....
iijR
.,::.. -,: ,
:i.7
'%:
-:.-.. -. 1'.......;:..:
-,:.:;--,-,** ,;.
. _W1114
1, .,:........
--.-6- -:.. -11510
.. .!.- I .... 0M
,-.-.-.- ,m-.j.j-.--
."...1%
:j
.q
- !! .1.
'r,:Ax 1w
:.. p '--L
:.,.,:
-,4:5":...-:.H...,;V..3
m". .j.1m._.- ;;..!,.$:
:.:.:i;. __m., .F:!....
P. :v:...1
. , _.
.,. ,...:.-
.!.:%.:. I.,
::..... -'r.:;.-i
. .. -.--.-,...:
-%
:.j F..:,.;..
.W1.4-4.11
. .L?
_ --- A '..... ..? -:.. ..%, 4...1%i%3.::.:-. mt;;m-m-1m-.-;F.
. .

;....L p V- I : j%
.!!.m:l-:C:
..:.....:...,:.: X::j.:::
-6x j. , : .m...
.0.41,111.
; -,.--X
,5lf!l:-.
. "'.11,
.: ... !,-,qi
.f.,,.:.:.:
.-ml ......
%%f,
..:..
.2 .f'..t ,.,--..-..;.;..-.:
.4!: F4;
-e ;1li.4i
,---e.e-m-+mm-m-----i1
,li7.:-,il,.j,,,.::f- .. :..:..
i!!, *R
-,.:-.-. N.,
!iim2-
%.4R.-.-.:-ii---.i-.,,
Lz.'.
m.
!
; i -Y
.-
r;F:4
A.,.!:--i g
...:.:
....- i: i .......
q.k
:,.Z?-",...:f?.
, fi:
IN
-,
iii,.:
5..",.-
. :i .. !:lf
;i;
-
: : ,!,;r
- ..::.4.:
.:
iii.
, .i ii-::;m,:,]';:n--l-m.-.-.
iZ..,
,i.
:1..0z." 1 .5
. .
,:-!
.;n,;i,.,.",.::::!:
j. . ..
.- -,,"
., ..;
::.....
"
..-.,:.-..,: ...
.,X,.
.. .....
...
-::-.--.j:
f,::"..
?',. .. :..
.,.,. .0i
!l :'
.tgj-.". convenient comparison, and each companion
. ,.:!n.::.
ini
:.,..
. .., Uf-.-,.,P, .:::
.4-:: ::::%%!
.01:! .fl:ii-:,.
!:.:9!-:
.. . : : . ...... . ,
. -..
.1 -x,;:!
--N-.iR
:;.-..:, K
.,:,i.:.:.:.
% . ......
..:..;.;.,:.:%
:2m!mi::%%-m
-
:.-.,.p'.-.. . - - Pim gw-zm4,.,.in:1r ::m: ...
V.-.9m:- -
.:..
,,!::-..:.
.. -
' :, A ..::i;,
:.;.;'..:;.:;..::::. %:9,11ill
.
_:X--
!:%".1:45-
%!,:
,.-..f...::..,:,:
".,. --'.f.:._.:i
.,. ;:.,:
--."
.:.:,.%'.%..
".: ..i.: .-...:..:2.1:
,. ; ..:..:.;:::.i.--.:;U., .,.-,,.
:!::f.,.,..- ;. :!I ....
'. -
.'Illillltia:
"
-,:f
;,.I.:.-. R..
.2 ;,;pAi...
..i.ii..
-:-,.v%-,-
,::.:..,.-,::b:i . ::J!-.,-----
...'... - ........
-..:.-Z;m--...-.:
.. : -! i;l._ . ... .C. . '. 91:4-x'..., ,
. - - . ---- .. - .-
,i! :,.
&--&
i::%:. -:&% . .
e :,4r
N..
sketch of the hallux ungual is drawn to the
..-A
.:!,. :, " ". :--:%,;:k ....,,.l'.-.-:.:
S. " 4. ....,%;:!
....
.1,::;: :.i
.? . . .
:. .::i
.:::X;,Rji:! ,;
1.... ..- ;,
,-'-
;,!,;!'1,!
...'..'.-".it?..,
.,.,.,::;:::..,!,;!",!
.::;il,:;h m. ::;fM
v:,i:i !:!.::.;:::r
- - . ,-'.
:,s,'--!ii:?i ... ::%. :! '. . rii: .....ii !,.,P '.f,
. I zm.m:.:-';P--:-:i:'1.
, .'-e..."
.... .. ,.r I~ ;:-,-'4W1, 4i_:-,
~ .i,-:,-:k..?1.' - :-
Vi--.,5:::.-,-,.4?. 4,--L;-,:..,-, f. -
":i-.. ...ImVni
.',%ll,;ri?..!,::!s i-m;
::.. .pt"
. .-. ....
. j . i..:...
.. il ::-x ..". . . I..
e -!1A- "; -,.. ,1
..... .. :: ,:1!'.z
i.!%-.:.1-.:-
.-.., . .0'.. ..
j.,...%;. . ,. .:::.:.::.
.:":.,....!.:.:,-.":
.,..",
.1 :,.::.,:::
:....
:."::i:i:i;
.-..,;
-
.
:. ..!;
. !-:.m..
,,,
..
.. . .::
*m:-'...
...
.--f;.,.
..
....
.. :...'..,.,i:.
.:,- -r
1. -
.;:.jj:
. ..
-.:
.....
:1,
......:. ., i
,-:.4-'
i's :.._
E....
-'..%.:.%:,!,::.;.',
:,.!.,..,:!:; , -..-:--.-F-,
:::!'.,
......,
,..
..
-f
i;%:w-xm---,
;i;l !-.:
.R. - ,%:>:,:;
'!T4,T.i,.:A
:.:,
;:....il,,:'%.-.,
-
.. . ..,i:.,.
%L.1.
p! I"fW.
'.'-'-.-;i3:
;,
%? .1/xitm.-',
..-;%i'.sj::f
... ;!,f!
r, ::5 , - -O,:,*,7,%.i
, k
,
F..I ii. ..,-;M;j-,f
"i x;ieii, 2i
-.ii . :,:-!...,-, .,..-
-A,
..:::,. -: _.-:i...1.:....;..
-;
-.-,-
:,::-
:::::
?.-.li:ii-.,i:;:;---..
:.
-_ . .;:,
11
.- I i"::-;:Y.:!::.;..:.
m .......
?-.'
, , -. -:!..
'v
-' ...
....... .%i&..
w,
.... .O.r,. ..
.. same scale to show relative size differences
" -i ..:-,
: : : . -L gnqi -.-it-.:,lii1 -it,;. -:-.-- ..,:,M -::::,.. .;:.,....;:,:,.:,.::!i::-' :-el , ,!.:.A...:.:..:L.;!
1. ii':i
. .
.., ., ii'. ... ...... .. .
:6N-..-- , ;:. --:--m,-1:E . " . .1q::e1:-aivf
::. - 4 :::!! _m ...
V--.-V -,
...- : . ,-
-i, .
.. .. ,
;,:-,;*. ::,..,
.. ..,:.
... %..9%. , '.,.,
...:.,4,. - :
--:"! :?".
-,..-..:--.:.--.,
.- --:j-:-4;-.:,%.l-.j--:
.. ;
: ! ..U. , .:., . -:
...::.::.
: !:.,-.,::
*,ii::1:! 'y
-. .!i,-
-...-.I-.--
.f:im; .,
........lw.
- :;,i'.'
.......
,.'
,,:..-i .,::-
ia-.--giiii
::; " "'i
-....:!% , .,.ii.: ! i,
:ff4.,:.:::.
* : -il... "., :R '.!! N:'.'
: plli:.' ,l
-,I.'-,::;
f,-,z.:.,.
:,-,-;:-i'.,ilK .,;5, , ....
::q;:.:,...
. ?..;:.,.:
. W : .l. m.. : .", X.
l!:,-;:-:i::i!
zol
n'-':--..
---
%.".
....:.-..::ww.:Il...%
- ..""..2
- ,
ic.:,::,
.r.mi-i;ii3l
.,..."A..F..:.
.
;,;...,.::r,
_.
, '.I.:.,.;g,
. . -,. .....
F% :
. .. ...4 *":.;,
...
-
:.;,."...-.?.' "'.
.: :. ....
.'..kA.".:"P:..
... .
.: "-.'
_ .
- ::.
.
.. ...
. :-.h.:
..'.! i
: .m "!i;:.
,.:.:
"..:::...,:.:,:,:,."::
.
' :' . ZXM
. .
.
.-;
..:
. ,
..", , -.,A,.
.
,
...,.P.!:
..: .,::::
;..k;p:.%.
-
,if,-*.,::
.;.:,:.P.
..;
% :
p: - ,'
-.:::qc:,%-,:,l-:c
Z?:i"
.:.:
.:::
-'.-
'..,
... .
;,xyi:,
i -i%
.
:i :i:.i
.-
,
.
...,
.... .:
.:
-
%
':;--....,-?- ..
-- .---:
%...-
: A:i..i.:!:.p
,,.F.,c.:.-
-.-,%:N
.,
;-- ..:, :Pjp
.....:"j;m
% .
.- - ..:.,.V..,
_...
....i
A.'' .!,,ii-5.",
. . . .
W
SimX
....
.;.
:mi:-.,.!::i
::K;M
"
-,i1:.i,1f'.-":-;
:;.::. :m%:j.,m,..:n.::,!
-::,%:: .. ..:f.:::::,
_-:...:;*FO.mM - -
i,:.";?..,::..:,. .:,.Z:..:::,;,.:.:.,.
'% .-A-Y,,....
. . .. . ..
i.,.-,;:f:,
1. --'m
. ..
fiii:l;! -.1.;.g'M
.,.,.,."!.;.:...w.:.."
,?mRX:m:,0
".
:::.:;:::.!..::f,.:: !.i-.
el, M
-..:i.
:.::....!,:i
::.%,..,!,
:-i:,-
;:.4:r.:.......'
.
-.%'-.'
-
.
WN.. ".-i :;"!
.
:!1
..
f.....
Im 9; , r between these two claws; the degree of en
Log?
.... V._
, "..-Na
-,
... V.
...-
... ....
.. 'A. " .-
il'A
,:-. :4. ..,-
- -:;!S'--1iz:i
!
..k....
A. ..-.
. . .
.,.::..-::.Z;.:::!
, .... %::",.
..':-.'-:f-.:.-...-i.'.'-....
i -.-Y.
- . .- 'i
.. ...:..::.::.:;
-: ... .s
.... :::,
.. .::
..!%:!.-.,..,:.,::
:-: ..... -;:"::.,::1-1-:i
.... ...... .......,:..-. ..,:::;,..%;:
.'-. ::;.....
- -..-.-:.,
... 1 :.ji:
....
:j.t :...
..:
.---.:..!...i:x:: I ..:::i:
.. .-..ek,.,
;:i..::%f.!-!
, .... -. .-:,
...., -:e
... .....
......
":.." ,Q;?!-.
,...r
- ,. .
v.
ioi:.,-:f-.;%x,1
.f:
.j... ,_:l:.. p;;
N:.m.-
.:::,:%
.1
.--.j-.'- : , ;.-,::::
- ....... . ...
,'..'.-.-.',-
..- - ,!,
- ...... - . ..'' .'..
I :i , .. - .., ..
..... :. , . -s!: .
.,.:P,;A
,t1K
.,
F-VA
.. -(
-_,T'-- . jrK .....
- ". "
-,.,
iiAip0
. I..
.....
- .
1'-,.;.-
---'
...i
...!;:,
,.
....
. ....,.-A
.:
...::
.-:F .%
-
. , %...
.a!i:,;-:---:
r.
. .;1
.-1.
. .1.i;:,:-.,.
-..:
.,!,;;;
1_1:f:: .,.;
....
,.:..:.:..
...:.i :%.
.
,
...
f.....
.kt-;1-
.,;..:.;..
-!-.-: *.
..
:'%
.
i"''.'.',S...i.::.-:-%:
..........
-': :
. -
,
-
-::,.,.-::,;
.1::",.i:: '':
.--.-;:.
...
, ..:
-:-:-,
,:,;., .:i-
. %..,P.-
..,:.:
-.
.:A::,
.::.
.....'%
X%.:- . ..
.:::
. .....::.
.::..,.:
.!:: ;, ..
. -...
-.
.r!-,:-1';;
.
-'.
.. .
-
-!..
...
... .,
.
.,.':;-."L,'::V-
..
..-!:;:-.-:!!---.
.
..
---.!:...i,%-,--.:
.:.:iZ:'-
. -- ..
&.Iii'.
'.::
.. ,
.
....
..-..
::-.s
1i:;i::0--!,:*1..y
- . :
.-.
..
..
,
'.;gr.
:-.!--
."
-
:i: 't
.;V-!:.!-;;
,..,..
..
p
:,
,.
4-00!p
:...
:'-.-.'.-i',
....
.....
.- !"i:
.!::.-.,:..-.,.":.
, ."4
4
' '.
...
... .,::, .- .
.......
"
-;Ae..:..---
.%i,
O.W
...l.--
....-.
..,:..:::."...,.i%:
,.P;.::!:.
,-:!:.;-.:?:..
-- - - -
.I
",:"
"
f!.
O .
0 .U.: U."..U.
U-m-.
ZMii::: ..- -
7-go!
....,.::;:
.,;:.;:.
i,,.'. LN
.l, . ;:..
ii
... M
,-
-- ....4.....
';'-t%:;"
.." ._
- .-A-.
......
.,:
;..Im ...
,;.
, " . ..j.
..F
!! .1-.-
.;
.I.I.Q
-w .. 5 k,5,
.'..-+;,Vd::fg,
..
..
R.
:.,:r. .: -, largement or reduction of each claw is indicated
-.; ,. . . ... .. . . .. .- . " ...%-- j!i! 'mi:-Al. .. .;..-...I. ."
...,.-y:;..i-:. mm -.._-_:.1..-..
. . ,.: . - ... N, -: . .

,,-, .......:Iiii,
'(j. .Ii -.. .,..
.............
."'! .. !... ...
: -.. - :..,;,i:.. % ..- -... ......:1 . .'.!.:f:f::4,:,.
:;.,:.",;:.:
.... .... . ... .; .-.. :.... ...... .,::.. ..-, .... .... ." -,iii ..12--l'i...'S!F,..'.?,
. I-, , _ i. I b: "
. '.. "'; ...". ... ..:: ..::
.:,
.n., .-')
'-.-
.-
i.-.11.9 -
...
:,..:
...,
...
.... ..I_
l,:?"r:,i ,
..:...'.....
.. ..I. .... -
... ... !i
:.....;..
,..:
.:..::.: ... ....: .; ........
.... ,.. ::.. i .:--C--;---....-.:-..,-.
!...W
L ?., .L ':...
.'-Eg!6 .. :.Y.
%-i. '_:. ... !- ,i
R:,%-::i-::%;N
.... ...
oi:j 2.i'- im
. ,.,:iz-:
ei!: ,r
-
,- -;. .m,
M. ." ,;-,.
-A--:-,--.-;,;i:-!;:,::-. .. -..
. ,.,:r
Uli..., .- -". 24
?!,'i '. ,
-. ""p
by the horizontal lines, all of which equal 5 mm.
, ,
.", 51:1. .1 .... ..:. .':.. :.,......,.,.;:".,;,. :...:
1,5 ----'!: i: ...;li :: .!-.,
.-. :a. .::,::, :: ,:4:,.;: 'X . -m! .%:,.."! .. ... . ... :,. '. .... :.
'....... i /. ft -.0 ..: : .- - :.. - -, - --$ : ..'. ...- ......:.
... . .
.i...:.X
-1
,.; .-..... .. :
:... i:-
..'..%.--:
. .......-?.:,... ,:;,.:U.::.'.X;.-.i.f,-.
..:. : .. ,-;i..i.% ,iF
.::,:.-.,.
:.... ...
.. ,...I!
kyd
.......
,i::-
iffi.,
..: II.I.-If%M.
--lq-T.W
":.::.:i-
. 'QP
. .1.
-'A;-h
-.-:1
!.-
:.;,:.;::;,:,;.,I1!-M I,z- . .iiiiii....
..w.,.kr.
- .-
%.
. ;;:..-
..-.--'..'...".-,; . ..:.
i.:. N ,IO.'ei-M"E"
. -;:j,-, ..,.-,i::...,.,
.....,161;,112,Z
., .,,,,r..M
- .....'.
'..,:
1, '..,
.. 'ge: .-
_,. .;
_R
q. -.: .:... .., .. .: -..:, .. ., :' ..:... . A"Xi-.. -.. , ."' -:;.i.i. !.4.:.-;:!l
...,:% . ,':'.I ..x, . . _;;m'l 'li' .A-le% MR
. :.:..4:.. : :.. ...':;m:-,
:l'..
. :M;
.. .;..,e
:.:i..-: -.,i:
... ..:-i,
-'-.,',!:i..;::
...i.: i::!ili
:-:---:, ...--.-.--.----.:.
,'. ', .F
,; -i:K.
-6 . .....:m lpg
.

.":!fm: .: -.f
,. r.., . *...%:,:::::,.. :.!'.:..- .:.::
.? -...: .: % ...... e.,. .., ,::;:i.. .;,... :i:: .,:-'pm ::;:!:;;:i.",'.' .,,;. 'im,
..J..x:.i.?-', f.
...'; !. .:.V..z :.. .,i. . ..."!N:
. .:..:. - ..:::,..
.....
.. ...:
:... :..:.i.
...
..,.. .. :.:' .,:::;,
..--
..%....:.: .
....f :,!:....:.........
..........
- ..r-Q
.: :.. .. -
......- ,w
_
.!,.;.
..
gn.,i
-..m.::m.:m:-.,. I.
it...,:.-..'l'.-
::. m-.,::-
"m.
.: ....M
:.........!; -
. . m:
'...;_k-:
.
i.; ..-;;:Z!;.:-.. ...,. .- V.:,. ,.....
:...:
":. .'..
..-f:.,.:.:.
'..,:C'. ... .: -. :"
..
.11*1111411; -;: Xj
....
:, Zi;,
-,-;j%,:-.V.-:-Fi:
-
.,
.. :,.:,-!.-,;.;Ci.!...-:!.g.-"!:A4p:-::.,..
11, z-ri:my.
IVI&I'M ::3 .,.
--,:i ..;--.:Rr: ...%':x
.. -,
..
.-.':' . . ".
:.,:,.::" ...
'I.i
".:. I . ,. ....
i.. '. :U;.:;
:.::.::..,;:,.r2...:.:
-.,:mi-%::--'1.;:!'m!.
. .:;:-m
.,..,:...-.i-..:.
:::
.4;
..-...-.,.::.
M ..WA,
. -.,.. '.'f:
-'.-;:. " ..: . -
II-.:-a-:4....F..!.1.
- .';.
. . R .11
.. ... .l. . c-i:-'!N--1T.1.rT .
- .. -. _ -__ Orientations are standardized by showing the
-- 2 1,
. -. i;e
-!,::
--wAm.-.-.-:-Z.V
:,.: ;---,..,,%--.
, .;.": -,
,-:1.:- .::.....
-%
';;+
.. ... %
. ...:" ':.:.,..:, ... . .: - ... 11 , -F-i--.iq.--;,-,:- -fi.-_--n---.
. . .-W!
. , ..i!:"
'-!:-'i-:,!i,
-: . - : L ...:
...:,i11 ..
..
?.
.
......
" . -.-
aw
. ,l:._,.r-,!;.1_1.
. .:
-.:..
-
. _ .:...'
.:;.
-
.
.
.:
:
.:.::
.::.
_ __ _ .
-
..
.A'.
.
.i...
: _
..
..
...
..
,m%
..ii.-m.m
.
..
.
......
_
.
.
.....:
..
. _-4
m .
;.-1:;i-:
:::i:......
-
: ".
.Y
-
;!:.;-.-
.
.:,;.,.:...::.
-.m..-mi..i.
.
.:
.
-
-
.,.:,
...
-:--:-.%-::-:::.::j;%f:,-
.. ...;-...
". M.
..
.
....
:
,.
,
ve. .
..
'. , .:.
:
.
.
y
-
;:::::
..
.
.
- ...,
-,!
..
..
_.
....
m.--
:X.:.....,
::jj.:;i:
.
-.igf.
.-
%.
.:
!e
- :-
. .
...
,
....
.
-.
;.:.;.
-
,
.:-,:
.
I--
- :
-:-
.1m
..:
... -nV-?
-:;..,.,-*'N
. f:g-..j;,m.ji;m;l
.::::
.i;p:iii0--.i.im;in;
... .
.:..::.::.;.., N
:;,
;.i
.
.....%!:
-'
...".....
.:
.:;
'. F1,
i'.j
:1f.
'.
P, - _,i
4..."..X..
'.
I ...
-i:.M
"f::,%
;.4..:.17
:%;j::!-,m-.-L..
fijl
;
. ?.!:
.:
ff
::.:
. F.-
- :.--
:::
e.
.,.:.
M
. , -?..j;:
-. W
.:
, -M'.-
-..
-M
.,
-% 0
........-ti
!,
i
.,
I . ... 0,
I
..
-.
N,
-
.,.q;;. ..
.,:,!1-
P
'.-.
.."
__
.
.
-1,
.Cmi
.
-W..
.-.
' -
..
::,:;;.
.....
r.:
:rl.:%!:!%F
. -I - .
.1-1
-.---::,-
,!10.
0:
..
.m
....
....
. -O' .., r
....
i. ,
g
PP'.,
.- .";',
.f...' .,
,:'i..
. F.7&':;V_.
.
-.i,.Lff1.
, ,....v--. ,
-M - , .
.
.1. 4'.::... ,.:.:'..::
-'.,i:,emll:;1
. i::: -.'j.;.
.. ...':..:--.
::;.:i::..
14 .
I .,;;.
,j.. . .... ::_
. ......,
.r.,....::... ..:.
- ;- ..
. - .. -,
.... .: :..,:..
-i? :,
- ,... .:i.....
.---.--.----v;ii . .... .-,,!!:.-,.:.i
.:.,.., -::ijj
.-"i
.,
!.4:.."'144
.M --,1;-;-j;gkF:-x
.:.: :"-
.: ;;"..I,
.,.,!.!";:.,f.;:.. .:,;! ..Ti r- ;.. -.,i..%lp,
:..
::. ,..,:,-.,iL:!?'..it.'K.
A-.w.,-:,.- '....
1 1--
....,; %-.;.'-;i
- ...:1.:-..,pi.-
.........
:;!-..-:; e.,t
;i
-i;;
4 -,
.., .'! ....
.-
-:;:
M .IM
.: "':
P,. .S-0
- I-..-rd!%
.-,0 15'....
- .4p;
j ...,
m articular facets in vertical position. In nearly
J .,... " :51 .:... -m.--- .: ..-_
, -,.....
. 4 :. .y
. .,. ....':..-'ii,'iF-M-
- -,-
.,. '':..' .:::::::
-'.. I.:,..- !....::: .: -: .-,%: .r -"Y.-e" L. -f ,... - X....lil,!;.f.,
-!-5.ii::;; 1;
""""" f-i
-_ :gg !M
M k ,,,, -.- ,; :-- ,.,? S,.;,?r, g4 L;
; 1. .1: . .I. .. ... 'N i.' - , * Ir .,: 'i . -g...
!7,'.'-.-i:m!i;; f, . 4. i!:-.::i:j.. 6 :ii, :;,.-..-.; .l[:. ..m:
:;:, ..:.m::;.-. . ..;: ..":.... :!,.
..f%. % .......
...:: . ...
- .: ". -:;::.;
- '-: ..:;45...
.%;::,Z.,.":
.. .. ... .....
:!..,!,:",-.,!..,.,....: . I..q- 1.4,;.,.,.:i
..:.:.:,..r ,.:..i:;s.,;I, . 'i.y-,:;,, .',.,%;...
! t-e,. -...,.. .:..i--,
.,!.....,i.,...,: -.g.
'.lf')i-g
,gg'n .-. mo all respects, the unguals of Archaeopteryx re
.i'.::::.,
.:. - - . . . . .1 " 4, - - .-, 7: :". 7 . i.
. :
. !.N.
I..
"!: : . .::.,
1 ,.:
r;:: .A--
'..i,-"i,. . . :;.
;i?.:.. :i
..:;-:iK
. l'.
E!; ,-.., .
"'? -A1:k;.?';'-;z.6
.,-"i?"",. -- 1-5
.:
:..:. . 1 . c;7ji.c
4:,Oi:.:..;.;i4P..-.N;lsAll - - - .. Y. . _M .
.- .-.
!,
m ...f:...
1.:.%
.. ...:
.:..
...:..:.. .
- - ... . . . .,
.. :-:---,
...
. ....
. .......1m%.
.
. ..
.
-
I :..:.;.... ..
'...,:'.-.
..... .: ... .'- :--
. .... . :--.,
: .
.' .-:../.:-::
. .
, ._-l-.V.,4
........ .m.--rm-
: ,.::.;,:;P.'
... .. i,;
-::,, :-- ..
. ,%-';'e
-':,:-
.4 ,K-
. .-;:i: -f'.
;.-
.:. l1':-.:..-
-, , -:;.W
:;Y5
rj.,
. ::,:.....
!,lrr
.,:% - i::;;'Ij
......
-:,: .jii.,;& :.. g
Q::;:..'......."....
.. ..
l'.. ..
.- ..... ..:i
. ..:.:.:%.
4 . ii. .m .__
pg ,---mm;-- w:, iZ ;v, ..
% ..
.. , ...
IR.,
7.;;lP!'S:' M -- .. .M
....... -
, . .P.M1
,?itn vm.
. il.l. . z" . :.. 4'.
_...,
Yp... _.
- :Api ..'g, l .
semble those of ground-dwelling birds more
:::
Fi -.-
:::::: -,,... . :-
-, .-.:
.. .:-. ..
... m , -.- .-
,:i
.. .., . i...,
.. _
'k
. . -.-,
..:.. ..i. .%..
.,: '- :,. -, ..
.;-:q-xii..p-
?,::...
;:
:::.,,...
.Z" - . -
Ob '.,"
-":'..
. N - i:
..W ,%.l%.-M1
:. ., ;11141%
,..-..:.,..,?..!
-iii'M-P:,: 2 . iiitv.m.
. .", ...'-
ilii:; 511-M ig.,t::,
- .:
i,.-,-.-,, ;.-.
, , N:
O, .V,
:i ..%;,-....
..yi ..ijvtii'e.i
:'ti: mm
4,:!i:i:-.:!,,e.
i- ...!." . I1x
.1 ,:,
11
W;..
1
-.0 1
fm
.-
..-:-:---.- ?M-
.AVWgl-m
M ..! :ir ;. il:
:g
- -
- .-.y"
I-.,. . .: . . w
%. : . :- ..m,
... ,:;.,:...
.i.. ..
.: . . . -:--.,-:'j.-.: ...:.:.:,: ....... .1. %- '...; u -- - ..-. ,., , .... . I M.I.- . Ri ,.- .1. ...., ..... I..,,,. .... %,R y
..-.-:.-"..
-... . .w .Z ... i:::ii i--,.:- ,m - --. q, f%-; i_.."-,ga.
A. . .1. . ii I.. 1- --:.- ... - .:", A .....r4c.
, ---
l;::...: .I..... .... . ..,:. . :. . ... .S.M.; ., ., . . tv .,;ll
:..,:,.: .:. , ,-::.. .i:P!i!i;'
...i ... ..--..,.: ,,,,, m . -, .::%-j:
.-. .,.:.. ,
....
., ..-M
'..".,:.,Rms?
., .,-1l;M
......
.l-.F.;l..%
. --Pn ., .-:! It-IM.
i.,lk-:.1'K-'. . w". 1.,:p.,,
;:.V.
P -,51Zj.
J, .cm. .!&.,
.,'i!-'-i;:mle-.'.;
, ..i
.m: .,.:;,. .---- -.M .:.. I.Y.
-,-: -,.,., iiw closely than any of the others: they feature
..
.:...,
.. -
. ,,-
m .",
m ,:,.
:-...
!"---- ! M m. , t: i6. . .., . .; -, _.
-.. -','. .m..;
. I:::. ;.,...: . .m..., .:::, -
. m..%?.
:.-, :-.-..-i.'.'.
C.
A"ll, X..
l.-,;,
. . -MUVOWIN,----
:i
" ,.--' .m,
. i: S-u-.
...'..' -.F...:%-4
9 ,r.K-_-,-.-'-Z
-
.....-,.-
:-i . :!;
.-WIMMM .:
jvZ!! -,:_Wrl .. -.-Xim.2;!
... -
.X;,.i--';;ipxr
IR , ,-.;!;
"A. ':-.,!-,-XfW,.
-,.1-.-6.-U 1-t -,iii,
. .gx..
r11A
m -m-
:.
.-- , mr.l.-M
:.:.-..i4:y.,
;P: ;N'
...m._r.:i,
-1, ...-
... I-.%.&:
.:.%
-'. .'. .
P...;j ,msw --
.--:-.;i;m-o!5:m$
-:.,l""Y111A.
. E-e ...1 mw .4
p .q.? .mz ai.-ii
...., ...Im ig..
.. & .
.....''."
.......".;::..' ..:,.,..::
-.-. i ....
61PIONOP0111-110"
.. ..
...
. .. :
:..
-!-
-!lfiiii7i
. I":
%i
., :...... 1, . . . .
-..f;;fm.:,...
. ..i: :.. . m
ilf , it!
- ,
, . .
,.::n. .3 .. . , f5.::::.
.., : . .--.,.
!N-'i!g
---1 ..;
ili... . IN: ; Wg -i
-.,
.-
:.;"' .,
.. ..%.;
,. ii
--.
.,.
:._ ...
..::
..
.-_..
.:.l..-.;. .., :-.-
..1&1-
...
;i--i.,
.1 ....
.:..
...
!'.'.".,;,M .a.:!"........ i.i;:- i:ii, :v
.I-,...Agm ..,.
!L-:%', .., -*'.1" ,O
.:,::.,:,:
:M
, ,i:,n'
l121. I... ,.1 ..7.
2%..
... ., .I
". - .. . , .. .. . .: .... . .. 7-S KS &SM.m . - iii ,:;i..,: e: .,..:e.i-'..'.'..': ii" , .,, .:U%.,ff:;r,.!, '-'.,!",:" ;.,:,.,.,.:t -, . ..,:. .. ', .i:..'-.I:., !:n ,...!:..-, ,,.:; .1 W W : ....... . . sfil
.-...%....'t: . .. . ...- --..i4.-- n.... "i.- :..... :.:,::"!.;k .l..r:;" ,l%-i-'IE-.Z!P ......... 1, ,-Y*..--. -. - .. ........ - ...'.
.,::.: ..*.--!:.,i:-...:f!;-.-j-.-.-- :14. . m!.m.; -.:::. .". .::..' -%,k m , .."'. . .... ..m.
-!;.:;
*.
.---: .....'::
.;:
:..
.:.%.
I :.
.F..
. .,% ,
. :...
... '::. .:g
.----.
....
,-... .m. ...
---.',- , ..---: !...-_. ......
.-.-1.-- .15
....:
--x:.As; .'Rz?
..'
-- .. ..
::
---,1ft ..
.... ..
, l.1;:
:i- ."..
-.z.-..i,::.,..
,"o, -
;-,ii:12:: .
... ' ..jp,
',' .,k
'.1"im".1110
f. -:
::. ,w.
",?i!
.:
. M-.p.'F,
.
!,"..-ig',.!:.lj.
-'a IN r. M ..
..
ON
.4
M.;:O
"M :-
.;'o
"i - . ,-
-KE
N.V..':Xe.,
-, Y. ,Pi
. :-.,i-lZ,4---U--
:-
---f;i-:-
. . m, ,-FA...4..
-..Gap - ... ,
...
:,:;A.:, .-.....a!
- ,i ..-iffi
-,,P,,ifo g ..
S.,3
.N.E slight curvature, robust form, shallow, poorly
-,-.,:,-. .:-: , , -., ,..,.- .:,,-::i. ! 6k'..."......
1..-:, Oii, -0 M . ;
. .. .%...-,, ::'':: . .:.,,-.::.. ., M117 ""
:.,. . -i:;i : in I . .N -, iZ NZ., ii'. .%. .?..
.:.;
.
.:.. . i:: . i,. ...::
...
..i
..:.
5'r-1.10..:t4,
. -.:.:!.% . !.: ,j '...,.
;c 11, " ESU
.-,..,/,:i.
-r:iii-7&f:;.-;g-%
,.. ,ii '-ZIN M
i I- ..1:.",.:..-,;::.
-::,::-'
7,.i;.. i%::..:.:
k,"'..'.. ?,C.,." ;i: ic;...
'....' -:%Z; .:,; .-.:-.- -.; 0 .:.1I. Z-mi.-Z
IQ%;;
_1iOM . --
IM ..rmi,OAn.: ,,,: ....;-.%:i20 V--S!i-.wmrW
-R:rg-.vy.,-:
..,,.: m- -.-.-.jj, ...- R- ".., '.: ,R . ,i.. , I _!; ...... ;;.i R ... defined flexor tubercles, and a hallux claw
.. X.% iA .-C.! k1? PU :; lli:,:ili. 3 i :,. ---"
:.,.lm!:-........:......-1:..'%.,.-
i n'-?*,:.-'-:: -:-.- -. ...".: .. ., :.

'-'i'-' ,!:.:.:j-!-. ,.,.:-

-,
; :- m
M. .:.
: -, :..,::
.. . . ,;i.
.. ---.,
i.
.b . .:.::...-
."....: ....
.
:.,
::;
.:i.: i: ..
.:
% .1
- ,
,.....:'-i:i'.:,m
...
-.., ..:,:
::::.:,:.::
..
,.,r
k : -:....:
,--- -
n-,.:;,,------
11 .:.,
...... . .
-:-*-,.4-.,,iO,.__-t'
.

R - -,,,"
.-.,.1..
.-....:
,
:,;;t...
.::.&:
. -
-
VUX, i
f....
',i-i,.-,:i.-,-:
-I...u
, ...;
."..i:
-g-:------
..!:
:-xi!i:;,;-:.f.:.:,-
qw
...... . pm......
-,
-
;...
- ..,
__
;
. -.-
?!,i.-?tl:; ,_
---,----------
- -
s..,- -'-
i.I.. , --
......:.- 2'r.:.
.,;-
"'
W_
- .
-i.-ii, "-
;i I
':
-: , .
::j A.A. .., . -1.
.:--,-.i-.---
: ....:.. :. - .....
.... ..': .,:..
:i;:-,:-:i'.,i...:
ggiivae.-nw. -:-,
:::: -
.. - ..
:..
,:,:Vi;, .- . ..,..
.i:.i.:::
..
.-;:.::.:-
- M .1 Am__..
,_ -
.. , !.-f,,!!*
.
.. IN
:,:
.., .wi-:
.. ..M.M.,mmm.
,
9
:.
-:; ..., .1.. 0. ;!I!
.. .: $':U:;-',::
.fio-ii;
. .;
. .
....:.n -,
,:,.
..CNj-Zmp.:-
:. " ',
49 -i
?.11!'<1".,Yw
:%W W ....--..".,..",",Ll
...."'..".1.-
-+::,.
... S.-;.
'C ,;!m
f,,!!*'C, 1:
... .m.n.-i..
.'4 --
'P -:8;r
i.'.---'-'%'
.. . ...-$:' .J-P:,
...2
r.,::.,...,:::.,.I.rk:
, .,..,::
,i.M
:...
.:__ ,:
.2 ,
.f.,"f-.
'.....AI .....
.i"..
-,:.4..,.....
'.....A
1, -- . ,,"..."
.;i. .
"..."'..."I
"..." '...I - .I .. .I
.-..,
e. 'k-:0
.:-.--:-4,...
P.M.. ,-;,-_..
% .....
...
:.. /.-
....:..
i..,
-
. ,.:
." ..;-;:ii. ..
.
?:fm...
,.,,,-
...,
-..
- lf
::..I.
"...
.4..-.:f-:;:
f.. .:.
:, .1.1
:i.. .
.S.
,:.:,.:.,
-..l;Q
1.
-:.
m....:..
. ..
. "....%:%
-.1.-i4j.
...
a.,j*,
,
,;:..."::..-,;,.;...;!:.:..:,
.,:,:
.1-mm"..W.
f-.%, .i..
-..::
,!,i.mX-
,:..
::i,:%,;;.?i
-g_ i
,-
.) .:x".
I--,'.
-'SftlmuiLi
.
%
.
..........
-:,...:.
"
: ..:.
...
IRN .. !::
'.',:1
i. .?
.--- .....
-.K_;.m-m1-.1.,;,!i5?:
::,.,.!. -.I
't
...._..rf;:L.
::: MiK....iFiK!
F!Uxv;
4.. ;
I..,
'::',
j?..:
L
io_-
.
if;"
-
.-,
:'. .,n:..
il.:....
.. :
:.1mil!-m;
":;R.A-&,l,.:jP..-.f
,ar,
,
- --.7 -
"f:
.....p r.-
s
..
_-
-,., .y
.
.:-,.1 11
--:-',,:.
m:i.-.::
:.-A.;.m.
:..-'. ....
...2
......
-
.:
. iiiii
11
-/ ;..v. .....2
.--- "' .
.:-tfZ..;;.:!
'.....A
..--T.ym:-
-J.A.:
F
.
k3;,,"..:,:;
. .
.--Z:j3g%-;
,
-......
.1i ..11
'-.,--
'...-R
-.!: :t
'._1-11
, -.:,Ei:;
.
....
.W.
-......
1 IP,
.1 ....
.:;:.;:;:.e
:i:
--l.::;.
UK .- "' .11:
. 3i
jR.,
?1H *,
h3iF
..
shorter than the other claw. (From Ostrom
I-f,:!! ii. ,, :: ;-:i . ,.. ,!aLr-...........,.,. ."..-,',

.%---::%.' .r...1- _!li

::-.i.4,i:.,
:.... '-:.-:i:i%;,.-i1
_-,
. !..:: .. -'--i
"....
. -,if
....:: .i.:
-1.. '.;:: .::
......
...:: *4:,ii ...
,:m:;.-;Zn-,.,
....- .-, .1.-
1-sl p
.-:-...
-.-. p
-%%
I......
,.!:5. ......:.. ...
--?!iiin,
...... !. -
- --.:;.,-
-AA% .. - .,
... ..
-,;.,
i, .-m
RPE-Ziv,;% :."!
-.-.-.,i-,l'-,.if . ....:;j.
%.
.. , .;.,,.k', ,::.....:.:;f.-. ,:kle...... 11. ... .... ... :. I-S.-
- . ..-
..
'.!,.:.:.-- 'r
:W
;j...&
1 k.,--,K-:-1;-:j-.,5-e.
Qp.
9:,.,; .T. iv..
1.-.--:-::...-.-..
:.-X,;!.lq,":-,_4i?:
.- _..
. i,:!i
,DAll'.
:X'51?
,.,;.;:...",..,..;:.::.f
-, ..,V
.,.:r, .". iil
".. .R_
x.,-.
.-::i-i:;iiF%!M
:ilJ--m-cZ0,,!:K.Ar; ';;e:;
. m,
--miq ,,,
..w : .U-
- _."
, tt'g'.
-,%- . . ...
-..........
-Xl .iVot-m-
% isy. .',!Rii,'.-
.'i....--
.tb m ........
.;n :5O-F.'.'r
.';.-;ky-,
... .- .".,-.'
.....
,ii::ii--'l'.
!F -I.-:'
--
.,,M! I i; ,'i-i.--
-:: _Ulgm ,.E
L.......,11.1
..I
Iii ."i
.,; . 1974.)
?. % ?/.
__
! A X. ,!...
/. . . 5i .1 ... '...
i.:i &.-r. :..I..:..!.
:-;i4ii-,.-,-. %!. - ---','f..;;-. .,.. .... %.. .F::::--i % ,,% ..... g::. -M: im ,%A:& .. .- . .g 1? .kM "-Wli t.:fk..;;
?:!..,4 -..' _._
.-,:;: .,-,, :4 ...... -
.i. U"::::
., i -.i.,,. -.
.;:. .:f:.:.. - :'?.
.....
i:i
....
vs; .. ,..-...i!r,.,,
'm
-"f ;O5,,, :.Ne
-i:::--.,..._t-
M.... -Ilx... -..--.--i-!rl-,,
.'.-ii".,:A.;-:..!,-.i.'
..M.:.:
,I N.m 1 ....
A!"..
- ...f.
.1 .......:.. ,,,
,illml-_li:
Ur .,"'-.:..
,: -:%-:----...,-
-;x.,:C...A
:.w7,.u::..:I . ;, . .-
%.;.'.-,,: if .-,!.,
'..'..'-.,-':9
.. ..
fil.-,F;,
..:-,.:::4lwl'
::.-..';,.'i'
14-
.. '.-!"'
, !1 ;,.. l_:. .91- I
.. .-i
-. ,-'%'i::..-:.'-.
. ,.'f
..-to. -ij
...."-" .,-I !.
,-, 1:,- ..,-,- !:";im.
:,:
.. .-.,:ii,--!
...
.-'.-..!i:-m .. M.,
.,A ,'_ -.1.I.-.1--m
.. ,
...- .- 050.0
.-.,,%, --r,,,;-Z ;.. .-,-:.;- %,* --.. ;,g, ;..-u;,-.:s4Fn---g;i:5;.__
_r.k.i;g.
".::::
...1. !;-- ....
:-?: .1 -.., -
...!-: -!m
. . .---
a
w .....
. . .k::.:,.
e.. :.
-..::: ? ....-
I
..-K!
:...-- ... ..
'-.-,::?.g
:h
':.1'.
!--.z
.. ...:g...
....
......... I.1
-:R-m!.::vg;if
I ;m;,
- l.1
.., .- ..
-;
I.M.
.1: ..O.. ,MV-,,ry;
,Cl
-1-41;.. .1 . ..
- - ;-q'-.giig-'1 -:.- E-,. -.%,
e.,1-x-- :-.,;- . rs- 2 . . . . . . . .
t:.1 --:,-::N-: ;:::%
j.....
. ..11 ..
.._
m P , -:,
.-_.:
....m
.: A
. . ..;:.,:
11w
'.:.,..'. l*
..... N;-
X
-ir. .. .....
. i:,p
...
-: -m ,_-.5T1 F. %a
....
, :..i:,.__..
_:
i::i%. -, ....
-.:.,!i.
.U
.";X R:
V:i:ii---
xt
9;.-,i:-:N',
--W
.....1m...
.,..-M,...
-gif
--,--N5-.._m,..m
-. ..:.il: 6:....
lil;
.1,
."... ...-....
......
m -Zr!--1;Yde::.;:i:.U..:::
-,.-
....",."N
!. ;:.r.il
.... -. :-
..
Z.,: -.
.. ','-
. .: . - - I -
.,.w
;U. - :K...
. M46.
..".
,% ,;:;!..,
. -;?;
0 i f.; ,_,.; .,..
,.,%-;
;:_ ._ 5lif-.:7.
-m
:.l..-"-:m:m.
" .4
.;.' . ".::
, .'." ,_: -: '.-. :.,. .!
-:;-i.%i-.,e;,4,. ': --,... ',:-. -
M.
'. : -::-.,.:.-,:,:.p-. Ile! .:, ":, ,., . , 'ilk '. . _ .
:.......iI:..::_ :.. . : ... . ::.:. . :.. .. - .

':
4..
.
- . ..4 .:.

:-
; -.---
...: . . v. M.M.
. .T
.. ...;.....K
:.:.:f:.
..
.,;%---;Ax.-
.: , , :.: : .:... .I.-
.--.P
. . : .. a:.!
. :. .... . V..y.
. ..
..:.
. j .r . :,"

-.m.; :.i
.
.. .I,!::.
1,
.%:,mm
,T!.i::,-.Si,-.;.,
, :.-.-frA!i
gx.-z...:iliR. .:f
I "ON; .. ... , - - ,.":" :. : : s . %, ,
.: i .""..-.M
.:.!...:.:.,
1011:1't
.
!'11
. A.p
%::..'.-
1' '1..R:,.
i.'M
.
: .m..
!.. I . . ,
r.1jry
'. ' a2ex-miiim-gm
m.i--:-.-:-
10.--.-i:.-.-:1i:
.,. ... .-/1.

.. ,..:
.
_ iUr
-, -,: i" '. -, ."; .-L.",
;;U!;,?.,,-.
- . !8 , .:e11r11. .: . C,
ly-
!i'.':
P..
.':.i!f .. . - . ,

....
:....
,; : ilk,
w.-.... . .... , .

.. . .,
....: ..:.:.
..,..
. . .
.. .. . ..
.......
1. ._. :
.
... ..
m
..- m -;...%
....:...
... .M ."' , ....
w
;....%%1:,..... ...
.... W
..;.....
.. %, ?!,
..._.._.
. ..?.:... ... . n
. :.- ;.
K.il.,f. :,
;-14 --.,
..
- .,. ,
....
_ ., :,
.........
:, : r I . :1
.. '!
, . .. Lv
:pri-,I~.:
.,.! . 4.14 -P; ;-;j .,_._ ..Z14.
.1 .1. . . . u . ....l...."..,
-1.1 - .. - ., X k, '- i i- '. 0-11
. -. iil,
..le:,
,:---:-:,, ,.;.:,: .!; .. . -:s
.:. k-. ..::
:.'.%:::: ;A --.- .----:-e -'-.'.'-F.; -.,; ,.. . :,;:::.i:! "A.0 .... . ." - ,
.. .. .. ,:-.i:,.-,r .. ;!M Kii! gap -:Lr.

" X% inm*, ;."i: kV-' '4' ":""'-;F,..

. .:.:.
. . - ..
i....."
.. -
,;... !'..'
"i.,i%
..f'...'.. - - ..
- F.
::,.,:4 ., il
...
.:
:,:"., M V vl
_. .1
".,.. . %,
.,.'i- M;F0,M
r,.-X.-.,.,::,;X-- -, Ii-S.
R- . . ..
M-
, ..iw .. 9,1 i .nm:rM..-l[:*;,%
.--
.-,. - iii .- in

pected in a perching or grasping foot.

Second, the hallux is relatively short,
as compared with the three main toes
and with the hallux in passerines or
predatory birds, where grasping is
critical. Third, the bony claw, or un
gual, of the hallux is distinctly shorter
than that of the median, or third, toe,
whereas in modern perching birds
and birds of prey it usually is consid
erably longer than the ungual of the
third toe. Finally, and perhaps most
important, the shapes of the bony
claws of all four toes in Archaeop
teryx are only moderately curved and
have very weakly developed flexor
tubercles. In all living perching birds
and birds of prey all claws are
strongly curved and have very prom
inent flexor tubercles. The relatively
low tubercles in Archaeopteryx
(compared with those of passerines)
clearly indicate that the claws could
not have been tightly flexed with
comparable force. It is significant, I
think, that the claws of Archaeop
teryx most closely resemble the rel
atively straight, low-tubercled claws
of modern ground birds, such as the
pheasant, partridge, and quail (Fig.
6). These anatomical facts point to a
ground-dwelling existence for Ar
chaeopteryx, rather than to life in the
trees.

the skeletal specializations that are vocates point to the reversed hallux of Of course, one can argue that Ar
essential parts of the flight apparatus Archaeopteryx, which, according to chaeopteryx might have reverted to
of modern powered avian fliers. Yet, them, must have been for grasping, as a ground habit from an arboreal an
the general consensus is that Ar in all living perching birds. That evi cestry in which the earliest phases of
chaeopteryx was at least a glider, dence, however, is not as conclusive as flight were achieved. But if that were
which leads to the seemingly logical it might seem. true, then how, or for what, were the
conclusion that Archaeopteryx must "wings" of Archaeopteryx used, since
have been arboreal, since it obviously First of all, the hallux of Archaeop the skeletal features seemingly es
could not "glide" from the ground up. teryx is situated higher on the meta sential to powered flight are lack
In support of this arboreal habit, ad tarsus (see Fig. 4) than is to be ex ing?
52 American Scientist, Volume 67

This content downloaded from 202.3.77.249 on Sun, 31 Jul 2016 14:55:59 UTC
All use subject to http://about.jstor.org/terms
A hypothesis of preflight
evolution
As I mentioned above, the overall
nature of the skeleton of Archaeop
teryx is remarkably similar to that of
certain small theropod dinosaurs
(coelurosaurs). The similarities in
clude almost identical three-fingered
hands, metacarpale and carp?is,
shoulder region, foot and ankle, parts
of the pelvis, and the vertebral col
umn, including a long unbirdlike tail.
Some critics claim that these ana
tomical similarities do not necessarily
mean a close phyletic relationship
between Archaeopteryx and the
theropods, arguing that they are the
result of convergent or parallel evo
lution. That is a possibility, but a very
remote one, as I have tried to show
elsewhere (Ostrom 1976a). There
simply are too many detailed simi
larities. However, regardless of which
explanation is accepted, these strik
ing anatomical similarities also mean
that these two kinds of animals were
similar in many aspects of their life
styles (Fig. 7). They must have been
doing the same kinds of things.
Small theropods such as Compsog
nathus, Ornitholestes, and Veloci
raptor clearly were fleet-footed bi
pedal predators. The foot in some
(perhaps all) bore a fully or partly
reversed hallux, but that certainly
was not for arboreal activity. The long
arms and three-fingered, strongly
clawed hands obviously were for
grasping and, in view of their sharp
piercing teeth, probably for seizing
prey, not branches. For theropods as
a whole, prey size probably ranged
from cat-size up. As for Archaeop
teryx, the tiny sharp teeth in its jaws
make it clear that it too was a preda
tor. But its small size, about that of a
common pigeon, indicates that it
lived on much smaller prey, most
probably insect-size organisms. The
image that comes to mind is that of a
fast-running, bipedal predator
flushing small animals or insects and
running them down. In short, Ar
chaeopteryx appears to have been a
miniature theropod?except that it
had feathers.
Figure 7. Comparison of the left hand of the
Berlin specimen of Archaeopteryx (top), that
of the small theropod dinosaur Ornitholestes
(center; scale bar equals 5 cm), and that of a
modern pigeon (bottom) shows the striking
similarity between Archaeopteryx and the
dinosaur. (Bottom photo by W. Sacco.)
tremely efficient insulation, mini
mizing heat loss when ambient tem
peratures are lower than body tem
peratures. But on the wings and tail,
they form airfoils producing lift and
propulsion.
We don't know what the original se
lective value of primordial feathers
might have been. It has been sug
gested that they originated in con
junction with flight (Parkes 1966), as
heat shields (Regal 1975), for colora
tion (Mayr 1960), and as insulation to
retain body heat (Ostrom 1974).
Whatever the original function, it
seems most likely that feathers ap
peared and evolved as a general body
covering, rather than in isolated
patches, and most probably were re
lated to epidermal functions or
properties. If this presumption is
correct, then our quest for the original
factors behind the origin of bird flight
can be rephrased, in part, to: What
were the reasons for modification of
small contour feathers on the arms
and tail into greatly enlarged contour
"flight" feathers? Was it to increase
the surface area to slow the rate of
fall? Or could there be some other
explanation?
The arboreal theory, as described by
Bock, depicts a bipedal ancestor
moving from the ground into the
trees, becoming increasingly arboreal
and perfecting the ability to leap from
branch to branch and tree to tree. The
specimens of Archaeopteryx clearly
indicate that bipedal carriage pre
ceded evolution of the flight ma
chinery. Indeed, it is that prior de
velopment of bipedality that released
the forelimbs from the normal busi
ness of support and locomotion,
making them available for other ac
tivities. In theropods, especially the
small coelurosaurs, the ultimate (and
perhaps the original) activity was for
grasping prey. The specimens of Ar
chaeopteryx seem to indicate that
that probably was true of the ances
tors of Archaeopteryx?and perhaps
of Archaeopteryx as well, since its
forelimbs are virtually identical to
those of some coelurosaurian thero
pods (Fig. 8).

The most critical components of the Figure 8. Sketches of the right arm and hand of The arboreal theory is a very logical
avian flight apparatus are the flight Archaeopteryx (top) and of the theropods model; otherwise it would not have
feathers, which are greatly enlarged Ornitholestes (center) and Deinonychus been so widely accepted. But it does
(bottom) show the striking anatomical simi seem to have one major weakness: it
versions of the contour-type feathers larities. The humeri are drawn to the same
that cover the rest of the body in length to minimize size-related differences; is predicated on the invasion of an
modern birds. In today's birds, these true sizes are indicated by the scale bars, which arboreal niche by an obligatory bi
body contour feathers provide ex equal 5 cm. (From Ostrom 1976a.) pedal ancestor (Tucker 1938). That

1979 January-February 53

This content downloaded from 202.3.77.249 on Sun, 31 Jul 2016 14:55:59 UTC
All use subject to http://about.jstor.org/terms
 - -- l task of grasping the next landing spot,
and not to have evolved into a para
a- -
chute, or a wing. Or, if the forelimbs
of this obligatory bipedal pro-avis had
been used to assist in climbing, why
did the lineage not revert to quadru
pedal clinging?
84

The distinctly independent actions

- - .

and functions of the hind and fore

-

-
A.
-
-
-
A
4 -!:S
- quarters of modern birds (and of Ar
chaeopteryx as well) indicate com
JP
plete functional separation of the two
ever since bipedal locomotion was
*0
A first achieved in bird ancestry. I
suggest that, unlike other vertebrate
-A*.
* T

fliers (which remained quadrupedal),

-> *r birds became arboreal after they had
*(
P.
developed the power of flight?not
before. I further suggest that the
unique and highly diverse nature of
151 avian flight was possible only because
- - 4- -A
it was preceded by uncompromised,
obligatory bipedal posture and loco
motion that allowed the forequarters
- - 9 - -
* 1.. - - jK
in a proto-Archaeopteryx stage to be
adapted for other entirely separate
and unrelated activities. The question
* * - * - - -r
remains, though: How did this lead to
flight? What was the original selective
advantage of enlarging the initially
small contour feathers that covered
the arms and hands?

It seems to me that two important

clues point to how, or why, this
feather enlargement came about.
First, the small size of Archaeopteryx
and the presence of tiny sharp teeth
in its jaws are surely solid evidence
Figure 9. This pigeon-sized specimen of specimens of Archaeopteryx. The head has
that Archaeopteryx was insectivo
Compsognathus, the smallest known carniv been twisted back over the body. Except for the rous. Second, the coelurosaurian di
orous (theropod) dinosaur, came from the same shorter arms and hands, it is very similar to nosaurlike design of the forelimb and
Solnhofen limestone that produced all the Archaeopteryx (Fig. 3). shoulder, and especially the hand and
wrist, suggests a coelurosaurlike
use?at least in its ancestors (al
may not seem odd at first, because so equal in size, and all four feet are though it is so nearly identical in Ar
many modern birds are tree-dwellers modified for clinging (Rothschild and chaeopteryx that I find it difficult not
and they are bipedal. In fact, birds are Dollman 1936). Apparently all four to believe a coelurosaurlike action
the most common of all arboreal ver limbs are used in moving about in the there too, despite the presence of
tebrates. But in modern birds, the trees. feathers). These lead to the thought
wing is fully perfected for flight, and that there may have been a behavioral
the feet are all that remain for In short, it seems highly unlikely to connection between this coeluro
grasping any perch. When we look me that a bipedal (and presumably saurlike predatory design of the
around for other arboreal creatures, cursorial) animal would shift to an forelimb and the development of en
almost without exception they are arboreal habitat and still retain larged "flight" feathers.
quadrupedal?with all four limbs obligatory bipedal stance?unless the
adapted to provide maximum cling forelimbs were already highly If the "proto-Archaeopteryx" lineage
ing capacity. Furthermore, several adapted for some critical, nonsup evolved from ? somewhat larger
kinds have even modified the tail into port, nonclinging action. Even given predaceous coelurosaurian stock into
an additional clinging appendage. the unlikely invasion of the tree world a smaller animal insectivore niche
The tree kangaroos (Dendrolagus) by an obligate biped, if this lineage (see Figs. 9 and 10), and if these ani
would seem to be an exception as evolved into more and more profi mals used the three-fingered hands to
"bipedal" tree inhabitants, but, un cient leapers (as the model specifies), seize ever smaller and smaller prey,
like other kangaroos, they have fore I would expect the forelimbs to have any modifications that would im
limbs and hindlimbs that are nearly been perfected more and more to the prove insect-catching ability would be

54 American Scientist, Volume 67

This content downloaded from 202.3.77.249 on Sun, 31 Jul 2016 14:55:59 UTC
All use subject to http://about.jstor.org/terms
highly advantageous. Is it possible to be a logical sequence within the large flying insects so plentiful during
that the initial (pre-Archaeopteryx) insectivorous precursors of Ar Solnhofen time (Late Jurassic). The
enlargement of feathers on those chaeopteryx. structural changes necessary to make
narrow hands might have been to in it a skillful powered flier, even com
crease the hand surface area, thereby From this beginning, it would seem to parable to the contemporaneous
making it more effective in catching require relatively small evolutionary pterosaurs, were still great.
insects? Continued selection for steps leading to brief flapping flights
larger feather size could have con after fleeing insects, with selection When I first proposed the "cursorial
continuing to improve the adductive predator" theory several years ago
verted the entire forelimb into a large,
light-weight "insect net." It is not powers of the pectoral muscles, to (1974), it generated quite a bit of in
difficult to visualize how advanta firm up the shoulder skeleton, and to terest but apparently very little ac
geous these paired "insect nets" align and firmly implant the "wing" ceptance. As one friendly critic put it,
would be in snaring leaping insects, or feathers, with the final achievement "The enlarged insect-nets' would
even in batting down escaping flying of an Archaeopteryx grade and the produce so much increased drag that
insects. The "proto-wing" at first may beginnings of powered flight. There they would work against the cursorial
have been used in sweeping move is no question in my mind that Ar running skills." I don't find this a se
ments to flush insects concealed in chaeopteryx was well beyond the rious problem because, more than
ground cover or in low vegetation, but simple leaping, insect-catching stage likely, the "nets" would have been
subsequent enlargement of the and perhaps was at the threshold of held in streamlined position during
forelimb surface area seems best ex powered flight. Its wing form, so the chase and not employed as snares
plained as increasing the snaring similar to the elliptical wing form until within range of the prey (see Fig.
range?reaching those insects that (Saville 1957) widespread among 10). It is also possible that the ex
were flushed beyond the range of the modern birds, is strong evidence that panded snaring range would have
mouth. From simple sweeping arm Archaeopteryx probably was at least more than compensated for any loss
movements, to flapping swipes at a weak, flapping flier. But it seems of running speed resulting from in
leaping or flying insects, to flapping unlikely that its flight proficiency was creased drag. Possible too is increased
leaps up after escaping insects seems enough to catch in flight the various leg muscle power t? overcome what

Figure 10. The author's attempt at recon
struction of a hypothetical stage in the early
evolution of birds shows a pre-Archaeopteryx
stage (top) and Archaeopteryx (bottom).
Proto-Archaeopteryx illustrates an early stage
in the enlargement of feathers on the hands
and arms as aids in catching insects. The en in a similar predaceous pose. Body, limb, and
larged tail feathers are hypothesized as aero feather proportions on the wings and tail of
dynamic stabilizers enhancing agility and quick Archaeopteryx are based on the Berlin speci
manuevering during the chase after prey. Ar men. Proportions of proto-Archaeopteryx are
chaeopteryx presumably was at or just past the based on Archaeopteryx specimens and the
threshold of powered flight, but is drawn here theropod Ornitholestes. Scale bars = 5 cm.

1979 January-February 55

This content downloaded from 202.3.77.249 on Sun, 31 Jul 2016 14:55:59 UTC
All use subject to http://about.jstor.org/terms
ever added resistances were encoun Bock, W. J. 1965. The role of adaptive mecha backed vulture, Gyps africanus. J. Exp.
tered. nisms in the origin of higher levels of orga Riol. 55:13-38.
nization. Syst. Zool. 14:272-87. ?-. 1975. Mechanics of flight. In Avian
More serious, perhaps, is the widely -. 1969. The origin and radiation of birds. Biology, ed. D. S. Farner and J. R. King, vol.
Ann. New York Acad. Sci. 167:147-55. 5, pp. 1-75. Academic Press.
held view that the excessive energetic
Brown, R. H. J. 1948. The flight of birds. I. The Regal, P. J. 1975. The evolutionary origin of
cost for a prOto-Archaeopteryx to flapping cycle of the pigeon. J. Exp. Riol. feathers. Quart. Rev. Biol. 50:35-66.
flap itself up off the ground makes 25:322-33. Rothschild, L. W., and G. Dollman. 1936. The
any cursorial theory most unlikely. -. 1951. Flapping flight. Ibis 93:333-59. genus Dendrolagus. Trans. Zool. Soc. Lon
That view may not be correct, though.
-. 1953. The flight of birds. II. Wing don 21(6):477-551.
Clark (1977) has commented: function in relation to flight speed. J. Exp. Saville, D. B. 0.1957. Adaptive evolution in the
Riol. 30:90-103. avian wing. Ev?l. 11:212-24.
If birds are derived from bipedal archo
-. 1961. The power requirements of birds Tucker, B. W. 1938. Functional evolutionary
saurs, and the forelimbs developed strong in flight: Vertebrate locomotion. Zool. Soc. morphology: The origin of birds. In Evolu
powers of adduction in their capacity as London Symp. 5:9,5-99. tion, ed. Sir G. de Beer, pp. 321-36. Oxford:
insect nets, then it is conceivable that the Clarendon Press.
-. 1963. The flight of birds. Riol. Rev.
first fliers may have had muscles capable 38:460-89. Tucker, V. A. 1969. The energetics of bird
of the power outputs required for hov Clark, B. D. 1977. Energetics of hovering flight flight. Sci. Amer. 220:70-78.
ering or low speed flight. -.
and the origin of bats. In Major Patterns in 1973. Bird metabolism during flight:
Vertebrate Evolution, ed. . . Hecht, P. Evaluation of a theory. J. Exp. Biol. 58:
We can add to this that the earliest C. Goody, and . M. Hecht, pp. 423-25. 689-709.
ascents were not powered by the NATO Advanced Study Institute Series,
Series A: Life Sciences, Vol. 14. Plenum
Williston, S. W. 1879. Are birds derived from
forelimbs alone, but almost certainly dinosaurs? Kansas City Rev. Sci. 3:457
Press. 60.
were initiated by powerful leaps de Beer, G. 1975. The evolution of flying and Yalden, D. W. 1970. The flying ability of Ar
generated by the strong hindquarters. flightless birds. Oxford Biology Readers chaeopteryx. Ibis 113:349-56.
Clark goes on to say: 68:1-16. Oxford Univ. Press.
Greenewalt, C. H. 1960a. The wings of insects
However, if Archaeopteryx was capable and birds as mechanical oscillators. Proc.
of brief periods of hovering or of ascend Amer. Philos. Soc. 104:605-11.
ing flight (for which power requirements ?-. 1960b. Hummingbirds. Doubleday.
are also quite high) then it should imme
diately have been capable of sustained-. 1962. Dimensional relationships for
flying animals. Smithson. Misc. Coll. 144
level flight at moderate speeds, as the (2):l-46.
power requirements for this are mini -. 1975. The flight of birds. Trans. Amer.
mal. Philos. Soc. 65(4):l-67.
Hartman, F. A. 1961. Locomotor mechanisms
My cursorial predator scenario lead of birds. Smithson. Misc. Coll. 143 (1):1
ing up to Archaeopteryx is specula 91.
tive to be sure, but no more so than Marsh, O. C. 1880. Odontornithes: A Mono
the arboreal theory, given the anato graph on the Extinct Toothed Birds of
North America. Vol. 7, Report of the geo
my of Archaeopteryx and modern logical exploration of the fortieth parallel.
birds. Enough points of evidence Prof. Papers of the Engineer, Dept. U.S.
contradict, or raise serious questions Army, No. 18, Washington, DC. 201 pp.
about, a preflight arboreal stage in the Mayr, E. 1960. The emergence of evolutionary
history of birds to give us cause to novelties. In The Evolution of Life I, ed. S.
Tax, pp. 349-80.
reconsider carefully whether the "up
from the ground" origin of bird flight Mivart, St. G. 1871. Genesis of Species. Mac
millan.
is so improbable after all. Make no
Nopcsa, F. 1907. Ideas on the origin of flight.
mistake: I emphatically reject the Proc. Zool. Soc. London, pp. 223-36.
Williston-Nopcsa hypothesis that has-. 1923. On the origin Of flight in birds.
the proto-wings developing as pro Proc. Zool. Soc. London, pp. 463-77.
pellers to increase running speed. But Ostrom, J. H. 1973. The ancestry of birds.
I do believe that the predatory design Nature 242:136.
of the wing skeleton in Archaeopter -. 1974. Archaeopteryx and the origin of
yx is strong evidence of a prior pred flight. Quart. Rev. Biol. 49:27-47.
atory function of the proto-wing in-.a 1975. The origin of birds. Ann. Rev.
Earth Planet. Sci. 3:55-77.
cursorial proto-Archaeopteryx,
-. 1976a. Archaeopteryx and the origin
which turned out to be preadaptive of birds. Biol. J. Linn. Soc. London 8:91
for flight. It is conceivable that a 182.
predatory function of the forelimb -. 1976b. Some hypothetical anatomical
still existed in Archaeopteryx and stages in the evolution of avian flight.
Smithson. Contrib. Paleobio. 27:1-21.
that it was this adaptation that led to
the unique nature of avian flight. Parkes, . C. 1966. Speculations on the origin
of feathers. The Living Bird V:77-86.
Pennycuick, C. J. 1960. Gliding flight of the
References fulmar petrel. J. Exp. Biol. 37:330-38.
-. 1968. Power requirements for hori
Augusta, J., and Z. Burian. 1961. Prehistoric zontal flight in the pigeon, Columba livia. J.
Reptiles and Birds. Prague: Artia Publish Exp. Biol. 49:527-55.
ers. -. 1971. Gliding flight of the white

56 American Scientist, Volume 67

This content downloaded from 202.3.77.249 on Sun, 31 Jul 2016 14:55:59 UTC
All use subject to http://about.jstor.org/terms