Pterosaurs

Pterosaurs
Neither
Neither dinosaurs
dinosaurs nor
nor birds,
birds, these
these creatures
creatures were
were flying
flying reptiles
reptiles
that
that endured for 135
endured for mlllion years.
135 million years. The
The ones
ones with
with wingspans
wingspans of 12
of12
meters are
meters are thought
thought to
to have
have been
been the largest animals
the largest animals ever
ever to
to fly
fly

by
by Wann
Warm Langston,
Langston, Jr.
Jr.

ew prehistoric
prehistoric animals
animals havehave cap­ is
is reptilian,
reptilian, including
including thethe shape
shape of
of the
the gard
gard the
the ptcrosaurs
pterosaurs as
as bats,
bats, birds
birds or
or fly­
fly-

F
Few cap-
tured
tured the
the imagination
imagination so so complete­
complete- teeth.
teeth. For
For another,
another, the
the pelvis
pelvis and
and hind
hind ing
ing marsupials.
marsupials. In
In 1830
1830 Johann
Johann Wagler,
Wagler,
ly
ly as
as have
have the
the flying
flying reptiles
reptiles known
known feet
feet are
are those
those of
of a
a reptile.
reptile. It
It seems
seems clear,
clear, a
a German
German zoologist,
zoologist, linked
linked the
the ptero­
ptero-
as
as the
the pterosaurs.
pterosaurs. Extinct
Extinct for
for the
the 64
64 mil­
mil- however,
however, that
that in
in their
their adaptations
adaptations to
to saurs
saurs to
to extinct
extinct marine
marine reptiles.
reptiles. He
He
lion
lion years
years that
that have
have passed
passed since
since the
the end
end flight
flight the
the pterosaurs
pterosaurs departed
departed so
so far
far considered
considered them
them intermediate
intermediate between
between
of
of the
the Mesozoic
Mesozoic era,
era, these
these dragons
dragons of
of from
from the
the reptiles
reptiles popularly
popularly known
known as
as mammals
mammals and
and birds.
birds. His
His reconstruction
reconstruction
the
the air
air have
have nonetheless
nonetheless figured
figured promi­
promi- the
the dinosaurs
dinosaurs that
that no
no investigator
investigator would
would showed
showed a
a swimming
swimming creature
creature with
with pen­
pen-
nently
nently in
in man's
man’s view
view of
of the
the earth's
earth‘s dis­
dis- now
now confuse
confuse them
them with
with either
either of
of the
the two
two guinlike
guinlike wings
wings and
and rudder-shaped
rudder-shaped feet.
feet.
tant
tant past
past ever
ever since
since Arthur
Arthur Conan
Conan Doyle
Doyle dinosaurian
dinosaurian orders.
orders. The
The pterosaurs
pterosaurs and
and The
The modern
modern consensus
consensus that
that the
the ptero­
ptero-
made
made them
them part
part of
of The
The Lost World. For
Lost World. For the
the dinosaurs
dinosaurs appear
appear to
to have
have evolved
evolved on
on saurs
saurs are
are flying
flying reptiles
reptiles was
was established
established
almost
almost two
two centuries
centuries paleontologists
paleontologists divergent
divergent paths
paths from
from earlier
earlier forms
forms of
of by
by the
the turn
turn of
of the
the century,
century, and
and the
the ptero­
ptero-
have
have been
been puzzling
puzzling over
over the
the fossil
fossil re­
re- reptilian
reptilian life.
life. saurs
saurs are
are now
now regarded
regarded by
by most
most investi-
investi-
mains
mains of
of the
the pterosaurs,
pterosaurs, and
and surely
surely oth­
oth- It
It also
also seems
seems clear
clear that
that the
the pterosa urs
pterosaurs
ers
ers have
have wondered
wondered how
how thethe pterosaurs
pterosaurs did
did not
not evolve
evolve into
into the
the birds.
birds. In
In this
this re­
re-
solved
solved the
the problems
problems ofof powered
powered flight.
flight. gard
gard the
the telltale
telltale anatomy
anatomy isis that
that of
of the
the
Hang-gliding
Hang-gliding enthusiasts
enthusiasts might
might well
well be
be wing.
wing. In
In a
a pterosaur
pterosaur the
the fourth
fourth finger
finger of
of
curious,
curious, because
because the
the larger
larger pterosaurs
pterosaurs each
each forelimb
forelimb was
was greatly
greatly elongated.
elongated. It
It
weighed
weighed about
about asas much
much as
as a
a human
human supported
supported the
the front
front edge
edge ofof aa membrane
membrane
hang-glider
hang-glider pilot.
pilot. that
that stretched
stretched from
from the
the flank
flank of of the
the body
body
Until
Until recently
recently it
it was
was thought
thought that
that to
to the
the farthest
farthest tip
tip of
of that
that finger.
finger. The
The oth­
oth-
pterosaurs
pterosaurs with
with wingspans
wingspans as
as great
great as
as er
er fingers
fingers were
were short
short and
and reptilian,
reptilian, with
with
eight
eight meters
meters represented
represented the
the maximum
maximum a
a sharp
sharp claw
claw at
at the
the end
end of
of each
each one.
one. InIn a
a
size
size for
for flying
flying animals.
animals. Nine
Nine years
years ago,
ago, bird
bird it
it is
is the
the second
second finger
finger that
that is
is the
the
however,
however, Douglas
Douglas A.
A. Lawson,
Lawson, then
then aa principal
principal strut
strut of
of the
the wing,
wing, and
and in
in the
the
student
student at
at the
the University
University of
of Texas
Texas at
at bird
bird much
much ofof the
the extent
extent of
of the
the wing
wing con­
con—
Austin,
Austin, discovered
discovered aa number
number ofof wing
wing sists
sists of
of course
course of
of feathers.
feathers.
bones
bones from
from anan unknown
unknown species
species of
of pter­
pter-
osaur
osaur in
in the
the Big
Big Bend
Bend National
National Park
Park in
in Although
lthough the
the fossilized
fossilized remains
remains of
of
West
West Texas.
Texas. The
The bones
bones were
were surprising­
surprising- .£\.. pterosaurs
pterosaurs have
have been
been found
found on
on ev­
ev-
ly
ly large.
large. The
The radius,
radius, a
a forearm
forearm bone,
bone, ery
ery continent
continent except
except Antarctica,
Antarctica, most
most
was
was almost
almost complete.
complete. It It was
was nearly
nearly pterosaur
pterosaur fossils
fossils come
come from
from the chalk
thechalk
three-quarters
three-quarters of
of aa meter
meter long.
long. No
No addi­
addi- deposits
deposits of
of western
western Kansas,
Kansas, from
from cer­
cer-
tional
tional remains
remains of
of this
this animal,
animal, named
named tain
tain sedimentary
sedimentary strata
strata in
in England
England and
and
Quetzalcoatlus northropi after
Quetzalcoatlus l1orthropi after the
the Aztec
Aztec particularly
particularly from
from thethe Solnhofen
Solnhofen lime­
lime-
god
god who
who took
took the
the form
form of
of a
a feathered
feathered stone
stone of
of Bavaria,
Bavaria, aa fine-grained
fine-grained rock
rock laid
laid
serpent,
serpent, have
have come
come to
to light
light since
since then,
then, down
down inin a
a quiet
quiet lagoon
lagoon during
during the
the Meso­
Meso-
but
but bones
bones of of smaller,
smaller, quite
quite similar
similar pter­
pter- zoic
zoic era
era and
and used
used by
by man
man for
for lithogra­
lithogra-
osaurs
osaurs havehave been been found
found in in another
another phy
phy throughout
throughout the the past
past few
few hundred
hundred
part
part of
of the
the park.
park. These
These further
further discov­
discov- years.
years. The
The first
first pterosaur
pterosaur fossils
fossils were
were
eries
eries allow
allow a a hypothetical
hypothetical reconstruc­
reconstruc- found
found in
in aa limestone
limestone quarry
quarry near
near the
the Ba­
Ba-
tion
tion of
of Quelzalcoatlus
Quetzalcoatlusthat that gives
gives itit a
a wing­
wing- varian
varian village
village of
of EichsUitt.
Eichstéitt. Their
Their discov­
discov-
span
span ofof 1111 to
to 12 12 meters.
meters. Quetzalcoatlus
Quetzalcoatlus ery
ery was
was reported
reported inin 1784
1784 by
by Cosimo
Cosimo
may
may thus
thus have
have been been the
the largest
largest creature
creature Collini,
Collini, a
a former
former secretary
secretary to
to Voltaire.
Voltaire.
ever
ever toto fly.
fly. Although
Although the the Texas
Texas ptero­
ptero- The
The fossils
fossils plainly
plainly included
included a
a winglike
winglike
saurs
saurs are
are still
still imperfectly
imperfectly understood,
understood, II structure.
structure. Nevertheless,
Nevertheless, Collini
Collini con­
con-
shall
shall review
review here here what
what isis known
known about about ceived
ceived them
them to
to represent
represent an
an amphibi­
amphibi-
them
them today.
today. The The subject
subject can
can bestbest bebe ous
ous mammal.
mammal. Similar
Similar misconceptions
misconceptions
approached
approached in in the
the context
context of of what
what is is persisted.
persisted. Indeed,
Indeed, even
even after
after Georges
Georges
known
known (or (or suspected)
suspected) about
about the the ptero­
ptero- Cuvier,
Cuvier, the
the 19th
19th century
century master
master of
of com­
com-
saurs
saurs inin general.
general. parative
parative anatomy,
anatomy, pronounced
pronounced what what
Perhaps
Perhaps the the least
least controversial
controversial asser­asser- he
he called
called the
the "ptero-dactyle"
“ptero-dactyle” (literally
(literally PTERANODON,
PTERANODON, a
a pterosaur
pterosaur of
of the
the Creta­
Creta-
tion
tion about
about the the pterosaurs
pterosaurs is is thac
that they
they wing-finger)
wing-finger) toto be
be a
a flying
flying reptile,
reptile, some
some ceous period, which lasted from
from 135
135 through
were
were reptiles.
reptiles. For
For one
one thing,
thing, their
their skull
skull of
of his
his contemporaries
contemporaries continued
continued toto re-
re- 64 million
million years
years ago,
ago, is
is depicted
depicted in flight. The
in /light. The

122

©1981
© 1981 SCIENTIFIC AMERICAN, INC
gators
gators as
as making
making up
up an
an order
order of
of reptiles:
reptiles: The
The pterosaurs
pterosaurs resembled
resembled both
both birds
birds models
models of of the
the head
head inin aa wind
wind tunnel
tunnel sug­
sug-
the
the Pterosauria.
Pterosauria. Roughly
Roughly 85
85 species
species are
are and
and bats
bats in
in their
their overall
overall structure
structure and
and gest
gest that
that when
when it it was
was turned
turned at at a
a right
right
known.
known. They
They are
are arranged
arranged in
in two
two sub­
sub- proportions.
proportions. In
In a
a way
way this
this is
is not
not surpris­
surpris- angle
angle toto the
the direction
direction of of the
the animal's
animal’s
orders.
orders. The
The older
older suborder,
suborder, the
the Rham­
Rham- ing.
ing. The
The design
design of
of any
any vertebrate
vertebrate animal
animal flight,
flight, itit increased
increased the the drag.
drag. The
The crest
crest
phorhynchoidea,
phorhynchoidea, appears
appears abruptly
abruptly in in that
that flies
flies is
is subject
subject to
to aerodynamic
aerodynamic con­
con- thus
thus may
may havehave been
been an an air
air brake
brake for
for
the
the fossil
fossil record
record in
in 200-million-y. ear-old
ZOO-million-year-old straints.
straints. AA more
more detailed
detailed examination
examination landing.
landing. It It also
also could
could have
have served
served asas aa
Triassic
Triassic limestone
limestone inin northern
northern Italy.
Italy. The
The reveals
reveals both
both similarities
similarities and
and differences.
differences. front-end
front-end rudder
rudder to to offset
oflset the
the lack
lack of
of aa
distinguishing
distinguishing features
features of of the
the Rhampho­
Rhampho- For
For one
one thing
thing the
the pterosaurs
pterosaurs had
had hollow
hollow tail.
tail. Further,
Further, it it may
may havehave helped
helped toto bal­
bal-
rhynchoidea
rhynchoidea are are many.
many. They They hadhad a a short
short bones.
bones. No
No doubt
doubt this
this represented
represented aa sav­
sav- ance
ance the
the long
long beak
beak andand inin that
that way
way al­
al—
face
face andand a a short
short neck.
neck. In In the
the wing
wing thethe ing
ing in
in weight.
weight. Birds
Birds too
too have
have hollow
hollow lowed
lowed a a reduction
reduction in in the
the mass
mass ofof the
the
wrist
wrist was
was short
short and
and the
the fourth
fourth finger
finger was
was bones,
bones, but
but they
they usually
usually are
are reinforced
reinforced neck
neck muscles.
muscles.
long.
long. TheThe animal
animal had had a a long
long tail.
tail. The
The more
more massively
massively byby internal
internal struts.
struts. In
In Other
Other large
large pterosaurs
pterosaurs seem,
seem, how­
how-
Rhamphorhynchoidea
Rhamphorhynchoidea included included some some pterosaurs
pterosaurs as
as in
in birds
birds the
the fusion
fusion of
of vari­
vari- ever,
ever, to
to have
have managed
managed without
without a
a crest.
crest.
of
of the
the smallest
smallest pterosaurs,
pterosaurs, which which were
were ous
ous bones
bones limited
limited thethe flexibility
flexibility ofof the
the Moreover,
Moreover, the the interlocking
interlocking of of the
the neck
neck
roughly
roughly the the size
size of
of a a sparrow.
sparrow. trunk.
trunk. Movable
Movable joints
joints were
were few,
few, particu­
particu- vertebrae
vertebrae in Pteranodon may
in Pteranodon may havehave kept
kept
The
The other
other suborder,
suborder, the the Pterodac­
Pterodac- larly
larly in
in the
the wings
wings andand the
the ankles,
ankles, thethe the
the animal
animal from
from rotating
rotating its
its head
head toto any
any
tyloidea,
tyloidea, appears
appears just
just asas abruptly
abruptly in in the
the places
places where
where the
the stresses
stresses onon the
the body
body great
great degree.
degree. Finally,
Finally, some
some Pteranodon
Pteranodon
late
late Jurassic,
Jurassic, about
about 50 50 million
million years
years lat­
lat- were
were great.
great. The
The long
long tail
tail of
of the
the rham­
rham— skulls
skulls lack
lack a a crest.
crest. If
If this
this isis aa natural
natural
er.
er. These
These animals
animals were
were distinguished
distinguished by by phorhynchoids
phorhynchoids was was stiffened
stiflened by by long,
long, condition
condition andand does
does not
not merely
merely reflect
reflect
a
a long
long face
face and
and a a long,
long, curved
curved neck.
neck. overlapping
overlapping outgrowths
outgrowths of of the
the verte­
verte- the
the imperfect
imperfect preservation
preservation of of the
the fossils,
fossils,
Both
Both thethe wrist
wrist and
and the
the fourth
fourth finger
finger con­
con- brae,
brae, and
and other
other vertebral
vertebral outgrowths
outgrowths the
the crest
crest may
may have
have been
been a a sexual
sexual char­
char-
tributed
tributed impressively
impressively to to the
the length
length of of limited
limited the
the sideward
sideward bending
bending andand twist­
twist- acteristic.
acteristic. Whether
Whether it
it would
would then
then have
have
the
the wing,
wing, and
and some
some of
of the
the species
species were
were ing
ing of
of the
the neck
neck of
of the
the great
great pterodactyl
pterodactyl adorned
adorned thethe male
male or
or the
the female
female cannot
cannot
exceptionally
exceptionally big. Quetzalcoatlus was
big. Quetzalcoatlus was called
called Pteranodon.
Pteranodon. be
be guessed.
guessed.
a
a pterodactyloid.
pterodactyloid. The
The Pterodactyloidea
Pterodactyloidea Many
Many pterosaurs
pterosaurs hadhad a a ridge
ridge oror crest
crest
were
were almost
almost tailless.
pearance
pearance of
tailless. The
of both
The sudden
both suborders
sudden ap­
suborders of
of the
ap-
the ptero­
ptero-
on
on their
their skull.
treme
treme in
skull. The
The crest
crest was
Pteranodon, where
in Pteranodon.
was most
where in
most ex­
in some
ex—
some spe­
spe- T headaptation
he most remarkable evolutionary
most remarkable evolutionary
adaptation in the structure of the
in the structure of the
saurs
saurs without
without any
any obvious
obvious antecedents
antecedents cies
cies it
it doubled
doubled the
the length
length of
of the
the head.
head. pterosaurs
pterosaurs was
was of
of course
course the
the wing.
wing. In
In its
its
is
is fairly
fairly typical
typical ofof the
the fossil
fossil record.
record. It
It The
The function
function ofof the
the crest
crest is
is aa puzzle.
puzzle. fundamental
fundamental design
design it
it resembled
resembled the
the
emphasizes
emphasizes thethe random
random nature
nature of
of dis­
dis- It
It surely
surely had
had an
an effect
effect on
on the
the aerody­
aerody— forelimb
forelimb of
of a
a tetrapod:
tetrapod: an
an animal
animal that
that
coveries
coveries in
in paleontology.
paleontology. namics
namics of of the
the animal;
animal; experiments
experiments with with goes
goes on
on all
all fours.
fours. Here,
Here, however,
however, the
the

animal.
animal has
has employed
employed the
the updraft
updraft along
along a
a cliff
cliff to
to gain
gain altitude,
altitude, then
then of
of the
the head
head could
could have
have been
been an
an air-control
air-control surface;
surface; the
the crest
crest may
may
turned
turned in
in a
a glide
glide toward
toward the
the sea
sea to
to search
search for
for fish.
fish. It
It is
is thought
thought to
to have
have thus
thus have
have offset
offset the
the lack
lack of
of a
a tail.
tail. The
The wingspan
wingspan of
of a
a big
big pteranodon
pteranodon
flapped
flapped its
its wings
wings only
only slowly
slowly and
and infrequently.
infrequently. The
The crest
crest on
on the
the back
back was
was as
as much
much as
as seven
seven meters,
meters, making
making it
it large
large among
among the
the pterosaurs.
pterosaurs.

123

© 1981 SCIENTIFIC AMERICAN, INC

©1981
greatly
greatly elongated
elongated fourth
fourth finger
finger had
had be­
be- ly,
ly, however,
however, the
the scapula
scapula and
and the
the cora­
cora— notarium
notarium waswas formed
formed by by the
the fusion
fusion of of
come
come a a long,
long, slender
slender wing
wing spar.
spar. The
The coid
coid were
were fused,
fused, and
and inin the
the larger
larger spe­
spe— several
several of
of the
the vertebrae.
vertebrae.
wing
wing was
was attached
attached toto the
the trunk
trunk ofof the
the cies
cies the
the scapula
scapula no
no longer
longer lay
lay loosely
loosely Many
Many ofof the
the pterosaur
pterosaur fossils
fossils that
that lie
lie
pterosaur
pterosaur by by aa massive
massive shoulder
shoulder girdle:
girdle: embedded
embedded in in the
the muscles
muscles along
along the
the flank
flank in
in fine-grained
fine—grained sediments
sediments include
include im­im-
a
a bony
bony ring
ring consisting
consisting of
of the
the scapula,
scapula, or
or of
of the
the animal,
animal, a a position
position typical
typical of
of the
the pressions
pressions of
of the
the wing
wing membrane,
membrane, which which
shoulder
shoulder blade,
blade, the
the sternum,
sternum, or or breast­
breast- scapula
scapula in in all
all the
the tetrapods
tetrapods except
except the
the extended
extended from
from thethe tip
tip of
of the
the fourth
fourth fin­
fin-
bone,
bone, and
and a a third
third bone,
bone, the
the coracoid.
coracoid. InIn turtles.
turtles. Instead
Instead thethe scapula
scapula was
was a a mas­
mas— ger
ger to
to the
the side
side of
of the
the trunk,
trunk, atat least
least in
in the
the
the
the smaller
smaller pterosaurs
pterosaurs the
the girdle
girdle rough­
rough- sive
sive rod
rod turned
turned inward.
inward. At
At the
the midplane
midplane rhamphorhynchoids.
rhamphorhynchoids. The
The membrane
membrane
ly
ly resembled
resembled thethe one
one in
in aa bird.
bird. The
The main
main of
of the
the body
body the
the top
top of
of the
the scapula
scapula was
was was
was reinforced
reinforced internally
internally by
by long,
long, thin,
thin,
difference
difference was
was the
the absence
absence ofof a
a fourth
fourth inserted
inserted into
into the
the notarium,
notarium, a
a bony
bony bar
bar closely
closely spaced
spaced collagenous
collagenous fibers
fibers that
that
bone,
bone, the
the wishbone
wishbone or
or furcula.
furcula. Typical-
Typical- unique
unique to
to the
the largest
largest pterodactyls.
pterodactyls. The
The probably
probably prevented
prevented sharp
sharp bending
bending ofof

FOSSIL REMAINS of the sparrow-size pterosaur Pterodactyills

Pleradactylus e/e­
ele- the
the animal
animal dried
dried out
out on
on a
a beach
beach before
before it
it drifted
drifted to
to the
the place
place where
where it
it
gans lie
galls lie in
in limestone
limestone from
from Solnhofen
Solnhofen in
in West
West Germany.
Germany. The
The lime­
lime- finally sank.
finally sank. In
In addition
addition to
to fossilized
fossilized pterosanrs
pterosaurs the
the Soluhofen
Solnhofen lime­
lime-
stone
stone solidified
solidified from
from sediment
sediment that
that lay
lay at
at the
the bottom
bottom of
of a
a lagoon
lagoon stone
stone has
has yielded
yielded fossils
fossils of
of small
small dinosaurs
dinosaurs and
and of
of the
the first
first known
known
some
some 150
150 million
million years
years ago,
ago, in
in the
the late
late Jurassic
Jurassic period.
period. The
The attitude
attitude bird, Arc/Iaeapteryx. The
bird, Archaeopteryx. The stone
stone has
has been
been used
used for
for centuries
centuries for
for litho­
litho-
of
of the
the skeleton
skeleton is
is unnatural.
unnatural. For
For example,
example, both
both wings
wings extend
extend toward
toward graphic
graphic printing.
printing. The
The specimen
specimen of
of Pterodactyills elegaus shown
Premdactylus e/egalls shown here
here
the left
left and
and then
then fold
fold under
under the rib
rib cage.
cage. Apparently
Apparently the
the carcass
carcass of
of is
is from
from the
the collection
collection of
of the
the American
American Museum
Museum of
of Natural
Natural History.
History.

124

©1981
© 1981 SCIENTIFIC AMERICAN, INC
the
the outer
outer part
part of
of the
the wing.
wing. Additional
Additional
fibers
fibers were
were arrayed
arrayed toward
toward the
the trailing
trailing
edge
edge ofof the
the wing,
wing, where
where they
they could
could de­
de-
crease
crease the
the wing
wing membrane's
membrane’s flutter­
flutter-
ing.
ing. The
The fibers
fibers were
were slightly
slightly arched
arched in
in a
a
fore-and-aft
fore-and-aft direction,
direction, thereby
thereby contrib­
contrib-
uting
uting to
to the
the camber,
camber, or
or upward-directed
upward-directed
convexity,
convexity, of
of the
the pterosaur's
pterosaur’s airfoil.
airfoil.
The
The fourth
fourth finger
finger was
was stretched
stretched out
out by
by
a
a long,
long, slender
slender tendon
tendon running
running from
from aa
muscle
muscle in
in the
the inner
inner wing
wing (the
(the homo­
homo-
logue
logue of
of aa tetrapod's
tetrapod’s upper
upper arm)
arm) to
to aa
bony,
bony, elbowlike
elbowlike outgrowth
outgrowth on
on the
the first
first
segment
segment of
of the
the fourth
fourth finger.
finger.
Adding
Adding to
to the
the area
area of
of the
the wing
wing was
was a
a
triangular
triangular membrane
membrane thatthat stretched
stretched for­
for-
ward
ward from
from the
the wrist
wrist to
to the
the base
base of
of the
the
neck.
neck. The
The triangle
triangle was
was supported
supported at
at the
the
wrist
wrist by
by a
a uniquely
uniquely pterosaurian
pterosaurian inno­
inno—
vation,
vation, the
the pteroid
pteroid bone,
bone, which
which evident­
evident-
ly
ly was
was a
a place
place of
of attachment
attachment for
for a
a ten­
ten-
don
don and
and aa muscle
muscle that
that held
held the
the membra­
membra-
nous
nous triangle
triangle taut
taut and
and braced
braced the
the wing
wing in
in
front.
front. It
It is
is known
known from
from well-preserved
well-preserved
specimens
specimens thatthat some
some rhamphorhyn­
rhamphorhyn-
choids
choids had
had aa vertical
vertical flap
flap of
of skin
skin at
at the
the
end
end of
of their
their tail.
tail. This
This was
was clearly
clearly some
some
kind
kind of
of rudder
rudder that
that must
must have
have enhanced
enhanced
the
the animal's
animal’s maneuverability
maneuverability in
in flight.
flight. It
It
could
could also
also have
have served
served in
in water
water if
if the
the
rhamphorhynchoids
rhamphorhynchoids were
were swimmers.
swimmers.
I

T hree
hree other
other aspects
anatomy
aspects of
anatomy deserve
of the
the pterosaur's
deserve separate
pterosaur’s
separate mention:
mention.
The
The first
first is
is the
the pterosaur's
pterosaur’s hand,
hand, or
or rath­
rath~ k4, _A—"r. "—15:11“
er
er the
the three
three digits
digits of
of the
the hand
hand that
that pro­
pro-
jected
jected outward
outward from
from the
the wing.
wing. If
If the
the
pterosaurs
pterosaurs walked
walked on
on all
all four
four limbs,
limbs, the
the r'/- . \ ‘»
hand
hand may
may have
have been
been employed
employed for /fl\
for
grasping.
grasping. Gorillas
their
Gorillas walk,
their fingers
fingers folded
walk, however,
however, with
folded against
against the
with
the palm
palm of
of
I 4
the
the hand;

done
hand; in
their
done much
have
in other

much the
have been
been a
other words,
their knuckles.
knuckles. The
words, they

the same.
same. It
a biped.
biped. In
they walk
The pterosaur
It may,
In any
walk on
pterosaur may
on
may have
have
may, however,
any case
however,
case the
the long
long
I I
fourth
fourth finger
finger was
was folded
folded out
out of
of the
the way
way TWO
TWO SUBORDERS
SUBORDERS of
of the
the order
order Pterosauria
Pterosauria are
are compared.
compared. The
The Rhamphorhynchoidea
Rhamphorhynchoidea (top)
(lop)
by
by means
means of
of a
a joint
joint at
at the
the base
base of
of that
that had
had wingspans
wingspans ranging
ranging from
from .4 to
to 2.2 meters.
meters. The
The face
face and
and neck
neck were
were short;
short; the
the tail
tail was
was long.
long.
finger
finger when
when the
the pterosaur
pterosaur walked.
walked. The
The Each
Each hind
hind limb
limb bad five long
had five long toes.
toes. Tbe
The Pterodactyloidea
Pterodactyloidea (bottom) had
had wingspans
wingspans as
as great
great as
as 12
12
joint
joint allowed
allowed the
the fourth
fourth finger,
finger, and
and with
with meters. Tbe
The face was long; tbe
the neck curved. Tbere
There was almost no tail. Fifth toe was rudimentary.
it
it the
the wing,
wing, to
to turn
turn upward
upward along
along the
the
side
side of
of the
the animal's
animal’s body.
body.
The
The second
second notable
notable aspect
aspect of
of the
the pter­
pter- suggests
suggests maneuverability
maneuverability in
in flight,
flight, take­
take- The
The third
third notable
notable aspect
aspect of
of the
the anato­
anato-
osaur's
osaur’s anatomy
anatomy is is the
the brain
brain of
of the
the ani­
ani- off
off and
and landing.
landing. my
my is
is the
the pterosaur's
pterosaur’s skin.
skin. Although
Although rep­
rep-
mal.
mal. In
In this
this regard
regard an an analysis
analysis of
of endo­
endo- The
The olfactory
olfactory bulbs
bulbs of
of the
the brain
brain were
were tiles
tiles are
are typically
typically covered
covered by
by scales
scales and
and
cranial
cranial casts
casts is
is illuminating.
illuminating. TheThe casts
casts small.
small. This
This suggests
suggests that
that the
the animal's
animal’s mammals
mammals are are covered
covered by
by hair,
hair, it
it has
has
form
form when
when sediment
sediment fills
fills the
the cranial
cranial cav­
cav— sense
sense of
of smell
smell was
was poor;
poor; thus
thus the
the ptero­
ptero- been
been suggested
suggested from
from time
time to
to time
time that
that
ity
ity of
of a
a skeleton
skeleton during
during its
its fossilization.
fossilization. saurs
saurs did
did not
not rely
rely on
on smell
smell for
for hunting.
hunting. the
the pterosaurs
pterosaurs may
may have
have had
had aa hairy
hairy
They
They therefore
therefore suggest
suggest the
the surface
surface fea­
fea- In
In contrast,
contrast, the
the optic
optic lobes
lobes were
were well
well coat.
coat. The
The reasoning
reasoning begins
begins with
with the
the hy­
hy-
tures
tures of
of the
the brain.
brain. Perhaps
Perhaps the
the best
best ex­
ex- developed.
developed. Hence
Hence the
the animal's
animal’s eyesight
eyesight pothesis,
pothesis, offered
oflered by
by T.
T. H.
H. Huxley
Huxley more
more
ample
ample of
of aa pterosaur
pterosaur endocast
endocast was
was dis­
dis- was
was good;
good; doubtless
doubtless the
the pterosaurs
pterosaurs used
used than
than a
a century
century ago,
ago, that
that aa flying
flying verte;
verte;
covered
covered in
in Jurassic
Jurassic rock
rock in
in northern
northern vision
vision for
for hunting
hunting andand navigation.
navigation. brate
brate such
such as
as a
a pterosaur
pterosaur must
must have
have
England
England late
late in
in the
the 19th
19th century.
century. The
The Moreover,
Moreover, they
they were
were probably
probably active
active in
in been
been aa warm-blooded
warm-blooded animal.
animal. After
After all,
all,
specimen
specimen is
is only
only 25
25 millimeters
millimeters long.
long. It
It daylight.
daylight. Overall,
Overall, the
the encephalization
encephalization flying
flying implies
implies a
a high
high rate
rate of
of metabolism,
metabolism,
nonetheless
nonetheless shows
shows that
that the
the cerebellum
cerebellum quotient
quotient ofof the
the pterosaur-the
pterosaur—the ratio
ratio of
of which
which in
in turn
turn implies
implies that
that the
the animal
animal
was
was substantially
substantially larger
larger in
in relation
relation to
to the
the the
the volume
volume oror weight
weight of
of its
its brain
brain to
to that
that must
must maintain
maintain a
a high
high internal
internal tempera­
tempera-
size
size of
of the
the animal
animal than
than it
it is
is in
in reptiles
reptiles of
of the
the brain
brain of
of an
an earthbound
earthbound reptile
reptile ture.
ture. A
A coat
coat of
of hair
hair would
would furnish
furnish insu­
insu-
that
that do
do not
not fly.
fly. Indeed,
Indeed, the
the floccular
floccular of
of the
the same
same size-was
size—was relatively
relatively large.
large. lation
lation against
against the
the loss
loss of
of body
body heat.
heat. It
It
lobes
lobes of
of the
the cerebellum
cerebellum were
were larger
larger than
than It
It indicates
indicates that
that pterosaur
pterosaur brains
brains were
were might
might also
also streamline
streamline anan animal's
animal’s body
body
they
they are
are in
in birds
birds of
of comparable
comparable size
size to­
to- large
large for
for the
the brain
brain of
of a
a reptile
reptile but
but to
to reduce
reduce its
its drag
drag in
in flight.
flight. In
In 1971
1971 A.
A. G.
G.
day.
day. This
This suggests
suggests a
a high
high level
level of
of muscu­
muscu- had
had not
not quite
quite attained
attained the
the relative
relative size
size Sharov,
Sharov, a
a Russian
Russian zoologist,
zoologist, presented
presented
lar
lar coordination.
coordination. More
More specifically,
specifically, it
it achieved
achieved by
by the
the brain
brain of
of a
a bird.
bird. the
the first
first clear
clear evidence
evidence that
that this
this reason-
reason-

125

©1981
© INC
1981 SCIENTIFIC AMERICAN, INC
ing
ing is
is correct.
correct. He
He had
had discovered
discovered inin Ka­
Ka« the
the wing
wing ofof Quetzalcoatlus northropi. The
Quetzalcoatlus Ilorthropi. The apart,
apart, and
and soso it
it cannot
cannot be be established
established
zakhstan
zakhstan thethe fossil
fossil skeleton
skeleton of
of a
a pigeon­
pigeon- remains
remains of of other
other 'parts
parts of
of the
the body
body have
have that
that all
all the
the fossils
fossils represent
represent a a single
single pop­
pop-
size
size rhamphorhynchoid,
rhamphorhynchoid, Sordes Sordes pilosus. not
not been
been found;
found; evidently
evidently the
the wing
wing of
of ulation.
ulation. Nor
Nor can
can itit be
be shown
shown that
that all
all the
the
The
The fossil
fossil was
was clothed
clothed inin "long,
“long, dense
dense the
the animal
animal waswas separated
separated from
from the
the body
body animals
animals lived
lived at
at exactly
exactly the
the same
same time.
time.
and
and relatively
relatively thick"
thick” hairlike
hairlike fossil
fossil ma­
ma- before
before the
the burial.
burial. The
The second
second group
group of
of Nevertheless,
Nevertheless, the the larger
larger and
and smaller
smaller
terial.
terial. It
It appeared
appeared toto Sharov
Sharov that
that the
the en­
en- fossils
fossils consists
consists of
of the
the scattered
scattered bones
bones of
of bones
bones would
would scarcely
scarcely be
be distinguishable
distinguishable
tire
tire body
body (except
(except for
for the
the tail)
tail) was
was indeed
indeed at
at least
least aa dozen
dozen smaller
smaller animals.
animals. All
All of
of if
if it
it were
were not
not for
for the
the disparity
disparity in
in size.
size.
covered
covered by by hair.
hair. them
them were
were found
found inin one
one area,
area, where
where it
it The
The smaller
smaller bones
bones might
might therefore
therefore be
be
appears
appears that
that all
all the
the animals
animals died
died over
over viewed
viewed as
as representing
representing immature
immature speci­
speci-

O ur knowledge of the giant West Tex­

Our knowledge of the giant West Tex-
as
as pterosaur, Quetzalcoatlus, comes
pterosaur, Quetzalcoatlus, comes
a
a short
short span
few
span of
few years.
years.
of time,
time, perhaps
perhaps only
only aa mens
mens of
ing
of Quetzalcoatlus
ing new
new discoveries
northropi. Pend­
Quetzalcoatlus Ilorthropi.
discoveries that
that may
may settle
Pend-
settle the
the
mostly
mostly from
from two
two groups
groups of
of fossils.
fossils. The
The The
The sites
sites where
where the
the two
two groups
groups of
of fos­
fos- issue
issue definitively,
definitively, the
the smaller
smaller bones
bones are
are
first
first group
group consists
consists of
of fragments
fragments from
from sils
sils were
were found
found are
are some
some 50 kilometers
kilometers classified
classified in
in taxonomic
taxonomic shorthand
shorthand as
as rep-
rep-

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it
E EOSUCHIANS
EOSUCHIANS

PTEROSAURS EVOLVED in the early part of the Mesozoic era. chians and the saurischians) were the reptiles called the thecodonts,
Their
Their appearance
appearance preceded
preceded that
that of
of the earliest
earliest birds
birds by about SO
by about 50 mil­
mil- which for their part evolved from the early,
early, lizard-shaped reptiles
lion
lion years
years and
and followed
followed that
that of the earliest
of the earliest dinosaurs
dinosaurs by
by about
about 20
20 known as eosuchians. Among the pterosaurs the rhamphorhynchoids
million years. It is hypothesized that the precursors of the pterosaurs first appeared some
first SO
50 million years before the pterodactyloids. The
(and
(and of
of the
the birds
birds and
and of
of both
both orders
orders of
of the
the dinosaurs,
dinosaurs, the
the ornithis-
ornithis- last
last pterodactyls
pterodactyls died
died out
out with
with the
the dinosaurs 64 million
dinosaurs 64 million years
years ago.
ago.

126

© 1981
1981 SCIENTIFIC AMERICAN, INC
INC
a PTEROID
PTEROID BONE
BONE NOTARIUM
NOTARIUM resenting Quetzalcoatlus sp.,
resenting Quetzalcoatlus sp., an
an undeter­
undeter-
\ mined
mined species.
species.
SCAPULA I

N
A
A pterosaur
pterosaur vertebra
vertebra found
found inin the
the
1940's
1940’s in
in Jordan
Jordan bears
bears on
on this
this problem
problem
because
because itit resembles
resembles the
the neck
neck vertebrae
vertebrae
of Quetzalcoatlus sp.
of Quetzalcoatlus sp. The
The vertebra
vertebra was
was
long
long and
and slender,
slender, and
and so
so it
it was
was mistaken
mistaken
at
at first
first for
for a
a wing
wing bone.
bone. It
It clearly
clearly repre­
repre-
sented
sented anan animal
animal larger
larger than
than any
any ptero­
ptero-
saur
saur known
known at
at the
the time.
time. The
The animal
animal was
was
therefore
therefore named Titanopreryx. The
named Titanopteryx. The point
point
comcolo I is
is that
that the
the vertebra
vertebra found
found in
in Jordan
Jordan is
is
almost
almost large
large enough
enough to
to fit
fit Q. northropi.
STERNUM
Q. northropz‘.
On
On the
the other
other hand,
hand, the
the fossil
fossil from
from Jor­
Jor-
dan
dan is
is older
older than
than the
the fossils
fossils from
from Texas.
Texas.
ALULA For the present TitallOpteryx
Tz’taizopreryx and Quet­
Quer-
,\ ,‘ CLAV|CLE zalcoatliis can
zalcoatlits can be
be regarded
regarded as as distinct.
distinct. If
If
I) g
future
future discoveries
discoveries demonstrate
demonstrate thatthat they
they
are
are the
the same,
same, the
the International
International Rules
Rules ofof
Zoological
Zoological Nomenclature
Nomenclature stipulate
stipulate that
that
Tiranopreryx must
Titanopteryx must replace
replace Quetzalcoatlus
Quetzalcoarlus
as
as the
the name
name for
for the
the Texas
Texas pterosaur.
pterosaur.
The
The preliminary
preliminary estimates
estimates of
of the
the size
size
of
of Quetzalcoatlus northropi were
Quetzalcoarlus norrhropz' were based
based
on
on comparisons
comparisons between
between the
the wing
wing bones
bones
of
of that
that animal
animal and
and the
the wings
wings of
of some
some
considerably
considerably smaller
smaller pterosaur
pterosaur species.
species.
\\ By
By this
this method
method itit was
was inferred
inferred that
that the
the
wingspan
wingspan ofof Q.
Q. northropi ranged from
northropiranged from 1111
to
to 21
21 meters.
meters. Extrapolations
Extrapolations from
from two
two
-=/ .g / ‘ .91. -=. CORACOID
large
large pterosaurs, Querzalcoatlus sp.
pterosaurs, Quetzalcoatlus sp. and
and
SCAPULA
SCAPULA
Pteranodon, refined
Pteranodon, refined the
the estimate
estimate to
to 15.5
meters.
meters.
Aeronautical
Aeronautical engineers
engineers quickly
quickly point­
point-
CLAVICLE ed
ed out,
out, however,
however, that
that a
a pterosaur
pterosaur with
with
the
the shape
shape of Pteranodon and
of Pteranodon and aa wingspan
wingspan
of
of 15.5 meters
meters might
might have
have weighed
weighed as as
much
much asas 136 kilograms.
kilograms. ItIt would
would then
then
have
have lacked
lacked the
the muscle
muscle power
power to to main­
main-
tain
tain level
level flight
flight byby flapping
flapping itsits wings.
wings.
Moreover,
Moreover, the
the strength
strength of
of the
the wing
wing
bones
bones would
would perhaps
perhaps have
have been
been insuf­
insuf-
ficient
ficient toto bear
bear the
the stresses
stresses the
the wings
wings
would
would have
have had
had to
to endure.
endure. Of Of course,
course,
Quetzalcoatlus northrapi did
Quetzalcoatlus northropi did not
not have
have
exactly
exactly the
the proportions
proportions ofof Pteranodon.
Pteranodon.
v: I Even
Even so,
so, an
an animal
animal with
with aa wingspan
wingspan of
of
/
15.5
15.5 meters
meters would
would probably
probably have
have been
been
SCAPULA I
I
I
at
at or
or beyond
beyond the
the engineering
engineering limits
limits for
for aa
STERNUM flying
flying machine
machine made
made ofof muscles,
muscles, ten­
ten-
dons
dons and
and delicate
delicate hollow
hollow bones.
bones.

CLAVICLE nfortunately the

Unfortunately the precise
precise length
length of
of
(I U the
the wing
wing of
of Quetzalcoatlus northropi
Querzalcoarlus norrhropi
,
~ CARP/1L8 .
m— ,
1”t I Y \x
is
is still
still uncertain.
uncertain. It
It is
is known,
known, however,
however,
-
that
that the
the metacarpal
metacarpal bone
bone of
of the
the fourth
fourth
. HUMEHUS _. ~ ‘11.: I 3} <1 finger
finger was
was relatively
relatively longer
longer in
in Quetzal­
Quetzal-

3
3;.
x ._
_ /' .
I g
x I 2} coatlus than
coat/us
that
than it
that some
some of
it was
of the
was in
the other
Pteranodon and
in Pteranodon
other bones
bones of
of this
and
this fin­
fin-
ger
ger were
were substantially
substantially shorter.
shorter. These
These
4 """'\\‘;\.\ ;/" fr", ' , I \ «Jf 3 t)II differences
differences suggest
suggest an
an adj usted calcula­
adjusted calcula-
. l - ,
f tion
tion by
by which
which the
the wingspan
wingspan of
of Quetzal­
Quetzal-
" /

DIGITS CARPALS {Ky

If I coarlus sp.
coatlus sp. emerges
emerges as
as no
no less
less than
than 5.5
5.5
ULNA “.7
SCAPUU‘ STERNUM meters,
meters, and
and that
that of
of Q. northropi as
Q. northropi 11 to
as 11 to
12
12 meters.
meters. Such
Such an
an animal
animal might
might have
have
WINGS
WINGS of
of a
a pterosaur
pterosaur (a), a
a bird
bird (b) and
and a
a bat
bat (c) are
are evolutionary
evolutionary variations
variations on
on a
a forelimb
forelimb weighed
weighed 8686 kilograms.
kilograms. In
In spite
spite of
of the
the
that was suitable for an earthbound
earthbound animal that walked on all fours. The variations are distinc­
distinc-
recent
recent discovery
discovery in
in Argentina
Argentina of
of an
an ex­
ex-
finger that supports the wing; in the bird it is mainly the
tive: in the pterosaur it is the fourth finger
tinct
tinct vulture
vulture whose
whose wingspan
wingspan is
is estimat­
estimat-
second, and
second, and in the bat
in the bat it
it is
is the
the second
second through
through the
the fifth.
fifth. In
In each
each animal
animal the
the wing
wing attaches
attaches to
to the
the
trunk
trunk by
by means
means of the shoulder
of the shoulder girdle,
girdle, a
a ring
ring of
of bones.
bones. The
The girdle
girdle of
of the
the larger
larger pterodactyls
pterodactyls is
is ed
ed to
to have
have been
been more
more than
than seven
seven meters,
meters,
peculiar
peculiar in
in that
that the
the scapula,
scapula, or shoulder blade,
or shoulder blade, turns
turns inward
inward and
and abuts
abuts the
the notarium,
notarium, a
a unique
unique Q. northropz' still
Q. Ilorthropi still would
would rank
rank as
as the
the larg­
larg-
pterosaurian
pterosaurian bone,
bone, at
at the midplane
midplane of
of the body.
body. The
The notarium
notarium is
is several
several vertebrae
vertebrae fused
fused togeth­
togeth- est
est known
known flying
flying creature.
creature.
er.
er. The
The arrangement
arrangement provided
provided a
a base
base for
for the
the action
action of
of the
the wing.
wing. The
The arm
arm of
of man
man is
is shown
shown in
in d. Few
Few assertions
assertions about
about how
how the
the ptero-
ptero-

128

©1981 INC
© 1981 SCIENTIFIC AMERICAN, INC
a

_ . ~ _~ , ..—~ ~ *‘ _. " '----'

L._J
.. ’ 1 CENTIMETER
CENTIMETER

.-"> ~

.r J

k . )4- " , ( {

is?
.7, .
fig)? V.-.»v~:7",7
“-,' V , _. . "777"” "V'~\.,
77—17.

b
b
_

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\‘

I
V
‘

q
.

f
. k )7

’7‘
d
‘

"
, _, "
,1 ,
'7 . I

>------...J
g___l
1 CENTIMETER

. . _'_‘,,
', -5 "
_.
.” 1ICENTIMETER
CENTIMETER d
d . .

, {/ I

7 .7 ' ' ' , ' «V . ‘ '------'

L__:
, , , 77,,e »’ 1o CENTIMETERS
10 CENTIMETERS
7 i , , 7v»~i—'“' , 7 7,, _, \ ,

t ‘ ' —-— ——»_ _. _.__.____._/

SKULLS
SKULLS of
of four
four pterosaurs suggest that their diets varied. Ptero­
Ptera- had
had peglike
peglike teeth.
teeth. It
It may
may havc
have eatcn
eaten insects. Dorygnathus (c) was
insects. DOI)'gllathus
dausrra (a) had
daustro had teeth
teeth that
that resemble
resemble the
the baleen
baleen with
with which
which some
some whales
whales toothy; Pteranadon (d) was
toothy; Pterallodoll was toothless.
toothless. It
It seems
seems that
that both,
both, however,
however,
strain
strain plankton from seawater. Allurogllathus
Anurognarlzus (b), a tiny pterosaur, ate
ate fish,
fish, because
because fossil fish have
fossil fish have been
been found
found within
within their
their rib
rib cage.
cage.

0
Q " /OLFACTORY I) (v ”.
,/ ‘\ BULB r I 4/, ' ‘ ,
/ I \_ / / x
‘ I
\\ / ‘ l I \
. I \,CEREBRUM
REBRUM / \ A/ i
I .
/A i A ”L“ (

-. \OPTIC LOBE ‘ \ 1,,

CEREBELLUM
F CEREBELLUM 9
\ X) FLOCCULUS
FLOCCULUS
\_,\fl/2

BRAIN of0f the ptcrosaur

pterosaur (a) is reconstructed from endocasts,
endocasts, which crn reptile, the alligator (d). The view is from
from above. In the pterosaur
formed
formed from
from sediment
sediment that
that filled
filled the
the skull
skull during
during fossilization.
fossilization. The
The the optic lobes
lobes (indicative
(indicative of
of vision)
vision) and
and the
the cerebellum
cerebellum (indicative
(indicative
other brains are
are of
of Archaeopteryx (b), a
a modern
modern bird
bird (c)
(8) and
and a
a mod-
mod- of muscular coordination) arc
are well dcveloped for a reptile's
reptile’s brain.
brain.

130

©1981
© INC
1981 SCIENTIFIC AMERICAN, INC
saurs lived are weIl supported. The dif­

YOU CAN TAKE IT WITH YOU

ficulty is that vertebrate animals in gen­
eral are seldom buried immediately in
the precise place where they lived. In­
stead the carcass lies exposed to scav­
engers, decay, weathering and transport
by flowing water. The Kansas pterano­
dons, for example, are found at sites that
were at least 160 kilometers out to sea
in the Mesozoic. Almost surely the ani­
mals had died in or over water far from
home. The Solnhofen pterosaurs are in
stone that was laid down as limy sedi­
ment in lagoons. The attitudes of many
of the skeletons suggest that the carcas­
ses had dried out on a beach and then
had floated to where they sank.
Efforts to imagine how the pterosaurs
became airborne have led to suggestions
that they launched themselves by jump­
ing or faIling off a cliff, by dropping
from their roosting place in a tree or
even by rising into a light wind from the
crest of a wave. Each hypothesis has its
difficulties. The first one rests in part on
the assertion that the hind feet of the
pterosaur resembled those of a bat and
therefore could serve as hooks by which
the animal hung from a cliff in prepara­
tion for flight. A recent study by Kevin
Padian of the University of California
at Berkeley suggests, however, that the
hind limbs and the feet of pterosaurs
were much more like those of birds and
dinosaurs than has generaIly been as­
sumed. None of these animals is thought
to have been much of a cliff-hanger.
As for the second hypothesis, it seems
unlikely that large pterosaurs could
have landed in trees without damaging
their wings. It is conceivable, however,
that the pterosaurs .climbed the trees.
The hooves of the biIly goat seem to
make the animal suitable only for walk­
ing, and yet biIly goats can climb into
3Smm photography
the low branches of trees by clinging to is as easy as adding
them with other parts of their limbs. The your camera for exciting
third hypothesis caIls for high waves to close-ups of your favorite subjects.
channel the updrafts the animal would

have employed to soar. The wind that
made such waves, however, might well
have been too strong for the animal to
control its flight once it was airborne.
The setting of the Texas pterosaurs
presents a special problem for imagin­
ing how the animal left the ground. The
fossils were found in siltstones and sand­
stones that were deposited in a broad
alluvial fan some 400 kilometers inland
from the nearest seacoast of the time.
Although highlands extended to the
west, there is no indication in the geolo­
gy of the region that any mountains or
cliffs were nearby. It appears, then, that
Quetzalcoatlus may have lived on fairly
flat, low-lying ground. There, as is the
habit of a vulture, it may weIl have
had to wait each morning until the sun
warmed the ground and strong thermal
updrafts developed. In the larger ptero­
saurs the musculature that animated the
wing was not impressively massive, and
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131

© 1981 SCIENTIFIC AMERICAN, INC

 “Physicist’s rire"
"Pb,sicist's Fire” the hind
the limbs were
hind limbs
considered, it
things considered,
things
Quetzalcoatlus could have
Quetzalcoatluscould
were long long but

have run
weak. All
but weak.
unlikely that
seems unlikely
it seems
run on on its
All
that
its hind
hind
pterosaurs may
pterosaurs
Bramwell and
Bramwell
the University of
the University
may have have lived.
George R.
and George
Reading have
of Reading
Cherrie D.
lived. Cherrie
Whitfield of
R. Whitfield
have mademade a
D.
of
a
legs and
legs flapped its
and flapped wings energetically.
its wings energetically. detailed examination of
detailed examination Pteranodon as
of Pteranodoll as
Still, if
Still, if the animal could
the animal could stand stand up up on on a flying
a machine. They
flying machine. conclude that
They conclude that a a
its hind legs
its hind legs and catch the
and catch the appropriate
appropriate pteranodon with
pteranodon with a wingspan of
a wingspan 69 me­
of 6.9 me-
breeze, a
breeze, single flap
a single flap of of the wings and
the wings and a a ters and
ters and a weight of
a weight of 16.616.6 kilograms
kilograms
kick with the
kick with legs may
the legs may have have beenbeen all all it
it could have
could flown only
have flown only in light to
in light to moder­
moder—
needed for
needed for takeoff.
takeoff. ate winds. A
ate winds. wind is
light wind
A light is one that blows
one that blows
Landing would
Landing would have have been been lessless prob­
prob- eight to
at eight
at to 12 miles per
12 miles hour; it
per hour; will lift
it will lift
Evidently the
lematic. Evidently
lematic. the air speed at
air speed at which
which loose paper
loose paper off off the ground and
the ground make a
and make a
a flying
a pterosaur stalled
flying pterosaur stalled was was less less forfor stand out
flag stand
flag out by by a third of
a third its length
of its length
a pterosaur than
a pterosaur than it is for
it is for a bird of
a bird of the
the from its
from staff. The
its staff. aerodynamic proper­
The aerodynamic proper-
weight. Hence
same weight.
same Hence the pterosaur could
the pterosaur could ties of
ties of the pterosaur in
the pterosaur in a glide would
a glide would
NC d 069 805 touch down
touch down gentlygently on on its its hind limbs. The
hind limbs. The give it
give only a
it only tendency to
small tendency
a small to sink.
sink.
animal‘s body
animal's would pitch
body would slowly onto
pitch slowly onto Thus it
Thus easily rise
could easily
it could rise in an updraft.
in an updraft.
"The Fireframe (grate)
Texas Fireframe
“The Texas channels an
(grate) channels an
hands as
its hands
its as the animal folded
the animal folded its its wings.
wings. All these
All calculations are
these calculations based, how­
are based, how-
fireplace into
amazing amount of heat from a fireplace into
How did
How did it move on
it move on the ground? Some
the ground? Some ever, on
ever, on a model of
a model Pteranoa’on in
of Pterallodoll in which
which
the room, without using any "
moving parts .. .
any moving . investigators suspect
investigators suspect that that thethe hindquar­
hindquar- the wing
the membrane stretches
wing membrane stretches from from the the
BETTER HOMES
BETTER AND GARDENS
HOMES AND GARDENS (2/80)
(2/80)
ters of
ters Pteranodon. at
of Pteranodon, were too
least, were
at least, too tip of the
tip of the fourth finger to
fourth finger to the ankle of
the ankle of the
the
THE SECRET OF SUCCESS weak to
weak support the
to support the body body at all. They
at all. They leg. The
hind leg.
hind The fossil record gives
fossil record gives nono evi­
evi-
Place logs novel Texas Fireframe®
logs on the novel Fireframc® instead that
suggest instead
suggest that the animal rested
the animal rested on on dence for
dence this. In
for this. fact, in
in fact, some ptero­
in some ptero-
grate to form a cavity which opens up to you.
you. its breast, which
its breast, which theythey assumeassume was was well well saurs the
saurs wing is
the wing known to
is known have been
to have been
Ignite paper in fire
cavity.. Eureka! The fire
in the cavity padded. The
padded. animal would
The animal would then then have have narrow at
narrow the base.
at the base.
takes hold quickly, spreads,
spreads. burns evenly, dragged and
dragged pushed itself
and pushed itself in the manner
in the manner In general the
In general relation between
the relation between the the
steadily
steadily in
in cavity.. That means the cavity
the cavity
certain bats.
of certain
of bats. Pad
Padian suggests, to
ian suggests, to the
the area of
area of the wing and
the wing the total
and the weight of
total weight of
throws a beam of radiant energy
energy at you, so the contrary, that
contrary, that the pterosaur had
the pterosaur had a a two­
two- the pterosaur indicates
the pterosaur indicates that that thethe wing
wing
is hot,
fire is
fire efficient,, fuel-saving.
hot, efficient fuel—saving.
legged stance.
legged Finally, certain
stance. Finally, certain fossil fossil bore less
bore weight per
less weight unit area
per unit than it
area than it
Texas Fireframe's
Fireframe‘s arms are height­
height—
tracks from
tracks from a a Jurassic sandstone in
Jurassic sandstone in Ari­
Ari- does in
does in a bird of
a bird of the same size.
the same size. Ordinari­
Ordinari-
adjustable—the key to easy
adjustable-the set-up. unique
easy set-up,
zona are
zona are attributed (perhaps erroneous­
attributed (perhaps erroneous- ly this would
ly this suggest that
would suggest that the flight of
the flight of
control fire.
control of the fire.
ly) to
ly) to a pterosaur. They
a pterosaur. They were were made made by by the pterosaur was
the pterosaur was slowslow but but maneuvera­
maneuvera-
MORE QUOTES FROM THE MEDIA
MORE an animal that
an animal walking on
was walking
that was on allall fours.
fours. Pterosaurs, however,
ble. Pterosaurs,
ble. however, lacked lacked manymany
"Hot, . . . easy to start
“Hot, even, slow burning ... the control
of the
of surfaces that
control surfaces that are are avail­
avail-
L.
by physicist
. . . designed by
...

“Cranberg’s conception
''Cranberg's
L. Cranberg."
Cranberg."
TIME (12121175)
TIME (l2/2l/75)
of the radiation
T
The mechanics of
he mechanics
subject also
subject
ties. Yet
ties_
of pterosaur
fraught with
also fraught
too sheds
it too
Yet it
is a
flight is
pterosaur flight
with uncertain­
light on
sheds light
uncertain—
how the
on how the
a able to
able to the
ample, an
ample,
The bird's
er. The
er.
birds. They
the birds.

bird’s tail
lacked, for
They lacked,
horizontal stabiliz­
effective horizontal
an effective
serves such
tail serves such a
for ex­
ex-
stabiliz-
a func-
func-
pattern from his log-holder is correct."
log-holder is
SCIENTIFIC AMERICAN (8178)
SCIENTIFIC AMERICAN (8/78)
is the steady, slow­
“The big advantage is
"The slow-
fire that results."
burning fire
g,’\
\

THE
THE NEW TIMES (12/31178)
YORK TIMES
NEW YORK (12/3l/78) \.
assembled. heavy­
Available in four, fully assembled, heavy-
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versions as follows:
Model
Modcl 5—25: 25"
S-25: 25" front 2]” back width, 13"
front width, 21" I3"
deep, l5" high. Model V-25:
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U-25: 25" l5" x 15".
2l” x 15" 15",
U-l7: 17"
Model V-17: 14" x 13"
17" x 14" U-33: 33"
13". Model U-33:
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x 15" x 15".
29" x 15" instructions.
15”. Copyrighted instructions.

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Name
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_____________ State

FIREFRAME CO.
TEXAS FIREFRAME
Austin,
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Austin, Texas 78764
Zip

EVIDENCE OF
EVIDENCE HAIR on the body of the pterosaurs was provided in 1971,
OF HAIR 1971, when these fossil
impressions of the pigeon-size pterosaur Sordes pilosus were found in Kazakhstan in the
USSR. The hairlike marks at the bottom apparently covered the entire body except the tail.
U.S.S.R.

132
1981 SCIENTIFIC AMERICAN, INC
© 1981 INC
tion. The pterosaurs therefore should flown in the same way.
would have flown floating platform.
thus could fish from a floating
flight
have been less able to maneuver in flight More recent efforts to duplicate ptero­
ptero- When stomach contents have not
than a bird is. The larger pterodactyls flight have been disappointing.
saur flight been discovered, the best clue to an ani­
ani-
flapped their wings slowly at
probably flapped mal's
mal’s diet is usually its dentition and the
flight
takeoff and for restoring level flight
when that was necessary. Once airborne, T heof fossilized
he fossilized contents of the stomach
contents of the stomach
of the pterosaur (or at least the fos­
the pterosaur (or at least the fos-
structure of its jaw. On such evidence it
has been suggested that Anurognathus,
Anurognathus, a
an animal such as Pteranodon
Pteranodon may have sils
sils found
found inside
inside the
the fossilized
fossilized rib
rib cage)
cage) tiny pterosaur with a short face and peg­
peg-
remained aloft for long periods, rising show that several species of pterosaur like
like teeth,
teeth, ate
ate insects.
insects. Whether
Whether the
the ani­
ani-
on updrafts
updrafts near surface features such fish. How the fish
ate fish. fish were caught is not mal was maneuverable enough to catch
as cliffs and then riding weak air cur­
cur- clear. The skeleton of the pterosaur insects on the wing is debatable. Other
rents over land or water. probably could not have withstood
withstood the species clearly had different diets. The
The smaller pterosaurs were proba­
proba- stresses
stresses of
of diving
diving into
into water,
water, and
and the
the lower jaw of Pterodaustro
Pterodaustro has hundreds
fliers, able to flap
bly more active fliers, flap their body of the pterosaur makes it
it about as of teeth that resemble the baleen of a
wings more energetically. They none­
none- suitable as an unfolded
unfolded newspaper for whale. The animal may therefore have
theless were probably slower than a bird motion underwater.
underwater. fish-eating Pteran­
fed on plankton. The fish-eating Pteran-
of similar size. In 1956 Erich von Holst Pteranodon
Pteranodon and some other pterodac­
pterodac- odon was toothless.
odon
of the Max Planck Institute for Be­ Be- fished on the wing. A
tyls may have fished Quetzalcoatlus
Quetzalcoatlus may
may also
also have
have eaten
eaten
havioral Physiology at Seewiesen in pteranodon gliding slowly just above fish,
fish, but
but where
where itit would
would have
have caught
caught
West Germany constructed a model of the surface of a body of water could them is a problem. There is no evidence
a'rhamphorhynchoid
a rhamphorhynchoid from rice paper, have dipped its beak into the water to to suggest that there were large perma­
perma-
balsa wood and aluminum. Powered by fish. The narrow profile
pluck out a fish. profile of nent bodies of water near the habita­
habita-
flapped its
rubber bands, the model flapped the beak would meet with little resist­
resist- tions of Quetzalcoatlus.
tions of Quetzalcoatlus. Moreover,
Moreover, the
the
up and down about 40 times with
wings up fish could then have been
ance. The fish rock
rock in
in which
which the
the fossils of Quetzalcoat­
fossils of Quetzalcoat-
a wingbeat of two to three times per sec­
sec- gulped down on the spot. Alternatively Ius
[us were found are almost devoid of fish
ond, then glided to a soft landing. The Pteranodon
Pteranodon could
could have carried fish in
carried the fish remains.
remains. The
The rocks
rocks do,
do, however,
however, bear
bear
flight was described as elegant-not
elegant—not at its
its throat
throat sac,
sac, a
a pouch
pouch that
that resembles
resembles the
the many traces of burrowing animals. In
all,
all, for
for example,
example, like
like the
the jerky
jerky flight
flight of
of a
a pouch of a pelican. The fossil remains of addition the presence of masses of fos­
fos-
bat. The wings of the model were broad­
broad- fish have been found in a fossil impres­
fish impres- silized logs in the area suggests period­
period-
er,
er, however,
however, than
than those
those of
of a
a pterosaur,
pterosaur, sion of the sac. Some investigators be­
be- flooding. Perhaps a monsoon
ic ancient flooding.
and
and so it is unlikely that the pterosaur lieve pterosaurs could land on water and was responsible. All of this raises the

LARGEST FLYING ANIMAL ever

ever to inhabit the earth
earth is thought to color) were found. It is calculated, however, that the wingspan of the
have been tbe
the pterosaur Quetzalcoatlus northropi, wbose
whose remains animal
animal must
must have
have been
been 1111 to
to 12
12 meters.
meters. The humerus (dark color) is
The humerus is
were discovered in Texas in 1971.
1971. Only incomplete wing bones (light the bone that appears at the top of the photograph on the hext"
next page.

135

©1981
© 1981 SCIENTIFIC AMERICAN, INC
possibility
possibility that Quetzalcoarlus employed
that Quetzalcoatlus employed times
times the
the earth's
earth’s present
present land
land masses
masses voke
voke such
such catastrophes
catastrophes as as volcanic
volcanic ex­
ex—
its
its slender
slender beak
beak toto probe
probe for
for mollusks
mollusks or
or were
were all
all more
more or or less
less in
in contact.
contact. Later
Later plosions
plosions or
or the
the lethal
lethal radiation
radiation of
of aa su­
su-
arthropods
arthropods living
living in
in shallow
shallow flood
flood ba­
ba- the
the sea
sea floors
floors spread
spread and
and the
the continents
continents pernova
pernova near
near the
the solar
solar system.
system. The
The new­
new—
sins.
sins. That
That wo
woulduld mean,
mean, of
of course,
course, that
that moved
moved apart.
apart. The
The pterosaur
pterosaur popula­
popula- est
est hypothesis
hypothesis is is that
that an
an asteroid
asteroid struck
struck
Quetzalcoatlus foraged
Quetzalcoatlus foraged on
on the
the ground.
ground. tions
tions became
became isolated
isolated and
and evolution
evolution ac­
ac- the
the earth
earth at
at the
the end
end of
of the
the Mesozoic
Mesozoic and
and
Since
Since the
the pterosaurs
pterosaurs were
were reptiles,
reptiles, it
it centuated
centuated their
their diversity.
diversity. Large
Large species
species gave
gave rise
rise to
to a
a dust
dust clo ud that
cloud that enveloped
enveloped
might
might be
be ssurmised
urmised that
that they
they were
were ovipa­
ovipa- evolved
evolved onon several
several continents.
continents. the
the earth
earth for
for several
several years,
years, dimming
dimming the
the
rous.
rous. Indeed,
Indeed, purported
purported pterosaur
pterosaur eggs
eggs As
As the
the Mesozoic
Mesozoic drew
drew to
to a
a close
close the
the light
light of
of the
the sun.
sun. The
The forms
forms ofof life
life that
that
were
were discovered
discovered in
in England
England more
more than
than aa land
land masses
masses of
of the
the earth
earth approached
approached survived
survived onon land
land were
were either
either small
small oror
century
century ago.
ago. On
On the
the other
other hand,
hand, the
the pel­
pel— the
the positions
positions they
they have
have today.
today. It
It was
was aa warm-blooded
warm-blooded or or both.
both. The
The mammals,
mammals,
vis
vis of
of a
a pterosaur
pterosaur is
is not
not similar
similar to
to that
that time
time of
of widespread
widespread flooding
flooding inin the
the low­
low- it
it is
is imagined,
imagined, survived
survived by
by eating
eating seeds.
seeds.
of
of aa bird.
bird. In
In particular
particular the
the passageway
passageway lying
lying areas
areas of
of the
the continents.
continents. The
The total
total It
It counts
counts in
in favor
favor of
of this
this hypothesis
hypothesis that
that
for
for eggs
eggs would
would have
have been
been relatively
relatively land
land surface
surface was
was temporarily
temporarily reduced
reduced to
to the
the end
end of
of the
the Mesozoic
Mesozoic inin some
some places
places
constricted.
constricted. TheThe problem
problem would
would have
have perhaps
perhaps half
half of
of what
what it
it is
is at
at present.
present. is
is marked
marked in in the
the geological
geological record
record by
by a
a
been
been worsened
worsened because
because a
a pterosaur
pterosaur egg
egg For
For example,
example, aa wide
wide inland
inland seaway
seaway cut
cut thin
thin layer
layer of
of clay
clay in
in which
which the
the concen­
concen-
would
would no
no doubt
doubt have
have been
been large
large to
to ac­
ac- North
North America
America in in two
two from
from north
north to
to tration
tration of
of exotic
exotic chemical
chemical elements
elements such
such
commodate
commodate the
the folded
folded wings
wings of
of the
the de­
de- south.
south. Pterosaurs
Pterosaurs then
then achieved
achieved their
their as
as iridium_is
iridium.is curiously
curiously high.
high.
veloping
veloping animal.
animal. It
It also
also seems
seems likely
likely greatest
greatest distribution:
distribution: they
they extended
extended both
both The
The consequences
consequences of of a
a collision
collision of
of an
an
that
that the
the mother
mother would
would have
have had
had to
to fly
fly to
to north
north and
and south
south of
of the
the Equator
Equator to to lat­
Iat- asteroid
asteroid with
with the
the earth
earth would
would surely
surely
search
search for
for food
food throughout
throughout the
the gesta­
gesta- itudes
itudes of
of about
about 70 degrees.
degrees. Mountain
Mountain have
have disposed
disposed of
of the
the last
last of
of the
the ptero­
ptero—
tion,
tion, in
in spite
spite of
of her
her additional
additional weight.
weight. ranges,
ranges, however,
however, were
were rrising,
ising, and
and when
when sa urs. But
saurs. But another,
another, less
less dramatic
dramatic expla­
expla-
Perhaps
Perhaps the
the young
young were
were born
born alive
alive in
in an
an sea-floor
sea-floor spreading
spreading subsided
subsided at at the
the very
very nation
nation alsoalso seems
seems possible.
possible. The The fossil
fossil
immature
immature state
state of
of development.
development. Pro­
Pro- end
end of
of the
the Cretaceous,
Cretaceous, the the ocean
ocean basins
basins record
record clearly
clearly shows
shows that
that the
the pterosaurs
pterosaurs
longed
longed nurture
nurture would
would then
then have
have been
been deepened.
deepened. TheThe sea
sea level
level therefore
therefore fell,
fell, were
were inin decline
decline during
during the
the last
last few
few mil­
mil—
necessary.
necessary. Masses
Masses of
of fossil
fossil conifer
conifer nee­
nee— and
and the
the flooded
flooded land
land reemerged.
reemerged. These
These lion
lion years
years ofof the
the Cretaceous.
Cretaceous. Moreover,
Moreover,
dles
dles found
found with
with the
the remains
remains of
of Quetzal­
Quetzal- changes
changes altered
altered the
the climates
climates and and the
the it
it is
is thought
thought thethe climatic
climatic changes
changes at at the
the
coatlus sp.
coat/us sp. may
may be
be the
the remnants
remnants of
of nests.
nests. habitats
habitats toto which
which the
the pterosaurs
pterosaurs were
were end
end ofof the
the Cretaceous
Cretaceous included
included decreas­
decreas—
accustomed.
accustomed. Extinction
Extinction gradually
gradually re­ re- ing
ing world
world temperatures
temperatures and and increasing
increasing
rom thethe geographical
geographical distribution
distribution of duced
duced their
their numbers
numbers until
until only
only Quetzal­ seasonal
seasonal variation
variation inin the
the weather.
weather. TheThe
P
From of Quetzal-
the
the fossils
fossils and
and the
the age
age of
of the
the rocks
rocks inin coat/us remained.
coatlus remained. Then
Then itit too
too died
died o ut.
out. earth
earth thus
thus would
would have
have beenbeen stormy.
stormy.
which
which thethe fossils
fossils are
are fo und one
found one may
may sup­
sup- The
The disappearance
disappearance of of the
the pterosaurs
pterosaurs The
The g usty winds
gusty winds ofof the
the storms
storms would
would
pose
pose thethe pterosa
pterosaursurs first
first appeared
appeared in in coincided
coincided with
with that
that of
of many
many forms
forms of of have
have been
been disastrous
disastrous for
for creatures
creatures such
such
what
what is is now
now southern
southern Europe.
Europe. FromFrom marine
marine plankton,
plankton, mollusks
mollusks andand reptiles.
reptiles. as Pteranodon, which
as Pteranodon, which were
were adapted
adapted for for
there
there they
they spread
spread into
into Asia,
Asia, Africa,
Africa, In­
In- On
0n land
land the
the dinosaurs
dinosaurs disappeared.
disappeared. soaring
soaring in in light,
light, steady
steady winds.
winds. Their
Their only
only
d ia and
dia and North
North America
America by by the
the end
end of
of the
the Indeed,
Indeed, no
no animal
animal heavier
heavier than
than about
about chance
chance would
would seem
seem toto have
have been
been to
to im­
im-
1urassic
Jurassic and and reached
reached South
South America
America 23 kilograms
23 kilograms survived
survived the
the Cretaceo us.
Cretaceous. prove
prove their
their aerodynamic
aerodynamic capabilities
capabilities by by
and
and A ustralia in
Australia in the
the Cretaceous.
Cretaceous. Their
Their The
The reasons
reasons for
for the
the mass
mass extinction
extinction are
are red ucing their
reducing their size.
size. The
The remains
remains of of Quet­
Quet-
dispersal
dispersal waswas facilitated
facilitated byby the
the relative
relative widely
widely debated.
debated. Early
Early ideas
ideas have
have given
given zalcoarlus suggest
zalcoatlus suggest that
that thethe pterosaurs
pterosaurs
lack
lack ofof geographical
geographical barriers.
barriers. In
In those
those way
way to
to explanations
explanations that
that sometimes
sometimes in-
in- evolved
evolved in
in the
the opposite
opposite direction.
direction.

COMPARISON OF BONES of Quetzalcoatlus northropi with those

Quetzalcaatlus lIorthropi us of an animal designated Quetzalcoatlus sp. in the absence of proof
proof
of smaller, similar pterosaurs found nearby contributed to the recon­
recon- that it was the young of the species lIorthropi.
northropi. The bone is .24 meter
meter
struction of Quetzalcoatlus. The bone at the top is the left humerus of long. The fossils of Quetzalcoatlus sp. are sufficiently
sufficiently complete for it
northropi. It is .54 meter
Q. lIorthropi. meter long. The bone below it is the right humer- to be said that the wingspan of the smaller animal was six meters.

1136
36

©1981
© 1981 SCIENTIFIC AMERICAN, INC