HOW PTERODACTYLES MAY HAVE ORIGINATED
Ornithosauria have many characters inseparably blended together which
are otherwise distinctive of Reptiles, Birds, and Mammals, and
associated with peculiar structures which are absent from all other
animals. They are not quite alone in this incongruous combination of
different types of animals in the same skeleton. Dinosaurs, which were
contemporary with Ornithosaurs, approximate to them in blending
characters of Birds with the structure of a Reptile and something of a
Mammal in one animal. If an Ornithosaur is Reptilian in its backbone, in
the articular ends of each vertebra having the cup in front and ball
behind in the manner of Crocodiles, Serpents, and many Lizards, a
Dinosaur like Iguanodon, which had the reversed condition of ball in
front and cup behind in its early vertebrae, may be more Mammalian than
Avian in a corresponding resemblance of the bones to the neck in hoofed
Mammals. But while Pterodactyles are sometimes Mammalian in having the
head of the thigh bone moulded as in carnivorous Mammals and Man, the
corresponding bone in a Dinosaur is more like that of a Bird. And while
the Pterodactyle shoulder-girdle is often absolutely Bird-like, that
region in Dinosaurs can only be paralleled among Reptiles.
Such combinations of diverse characters are not limited to animals which
are extinct. There were not wanting scientific men who regarded the
Platypus of Australia, when first sent to Europe, as an ingenious
example of Eastern skill, in which an animal had been compounded
artificially by blending the beak of a Bird with the body of a Mammal.
Fuller knowledge of that remarkable animal has continuously intensified
wonder at its combination of Mammal, Bird, and Reptile in a single
animal. It has broken down the theoretical divisions between the higher
Vertebrata, demonstrating that a Mammal may lay eggs like a Reptile or
Bird, that the skull may include the reptilian characters of the malar
arch and pre-frontal and post-frontal bones, otherwise unknown in
Mammals and Birds. The groups of Mammals, Birds, and Reptiles now
surviving on the earth prove to be less sharply defined from each other
when the living and extinct types are considered together. But in
Pterodactyles, Mammal Bird and Reptile lose their identity, as three
colours would do when unequally mixed together.
This mingling of characteristics of different animals is not to be
attributed to interbreeding, but is the converse of the combination of
characters found in hybrid animals. It is no exaggeration to say that
there is a sense in which Mammal, Bird, Reptile, and the distinctive
structures of the Ornithosaur, have simultaneously developed from one
egg, in the body of one animal.
The differences between those vertebrate types of animals consist
chiefly in the way in which their organisation is modified, by one
strain of characters being eliminated so that another becomes
predominant, while a distinctive set of structures is elaborated in each
class of animals. The earlier geological history of the higher
Vertebrata is very imperfectly known, but the evidence tends to the
inference that the older representatives of the several classes
approximate to each other more closely than do their surviving
representatives, so that in still earlier ages of time the distinction
between them had not become recognisable. The relation of the great
groups of animals to each other, among Vertebrata, is essentially a
parallel relation, like the colours of the solar spectrum, or the
parallel digits of the hand. It was natural, when only the surviving
life on the earth was known, to imagine that animals were connected in a
continuous chain by successive descent, but Mammals have given no
evidence of approximation to Birds; and Birds discover no evidence that
their ancestors were Reptiles, in the sense in which that word is used
to define animals which now exist on the earth. When the variation which
animals attain in their maturity and exhibit in development from the egg
was first realised, it was imagined that Nature, by slow summing up and
accumulation of differences which were observed, would so modify one
animal type that it would pass into another. There is little evidence to
support belief that the changes between the types of life have been
wrought in that way. The history of fossil animals has not shown
transitions of this kind from the lower to higher Vertebrata, but only
intermediate, parallel groups of animals, analogous to those which
survive, and distinct from them in the same way as surviving groups are
distinct from each other. The circumstance that Mammals, Birds, and
Reptiles are all known low down in the Secondary epoch of geological
time, is favourable to the idea of their history being parallel rather
than successive. Such a conception is supported by the theory of
elimination of characters from groups of animals as the basis of their
differentiation. This loss appears always to be accompanied by a
corresponding gain of characters, which is more remarkable in the soft,
vital organs than in the skeleton. The gain in higher Vertebrates in the
bones is chiefly in the perfection of joints at their extremities; but
the gain in brain, lungs, heart, and other soft parts is an elaboration
of those structures and an increase in amount of tissue.
The resemblances of Ornithosaurs to Mammals are the least conspicuous of
their characters. Those seen in the upper arm bone and thigh bone are
manifestly not derived from Mammals. They cannot be explained as
adaptations of the bones to conditions of existence, because there is no
community of habit to be inferred between Pterodactyles and Mammals, in
which the bones are in any way comparable.
Other fossil animals show that a fundamentally Reptilian structure is
capable of developing in the Mammalian direction in the skull, backbone,
shoulder-girdle, hip-girdle, and limbs, so as to be uniformly Mammalian
in its tendencies. This is proved by tracing the North American Texas
fossils named Labyrinthodonts, through the South African Theriodonts,
towards the Monotremata and other Mammalia. Just as those animals have
obliterated all traces of the Bird from their skeletons, Birds have
obliterated the distinctive characters of Mammals. The Ornithosaur has
partially obliterated both. With a skull and backbone marked by typical
characters of the Reptile, it combines the shoulder-girdle and
hip-girdle of a Bird, with characters in the limbs which suggest both
those types in combination with Mammals.
The bones have been compared in the skeleton of each order of existing
Reptiles, and found to show side by side with their peculiar characters
not only resemblances to the other Reptilia, but an appreciable number
of Mammalian and Avian characters in their skeletons. The term
"crocodile," for example, indicates an animal in which the skeleton is
dominated by one set of peculiar characters. Crocodiles retain enough of
the characteristics of several other orders of reptiles to show that an
animal sprung from the old Crocodile stock might diverge widely from
existing Crocodiles by intensifying what might be termed its dormant
characters in the Crocodile skeleton. Comparing animals together bone by
bone it is possible to value the modifications of form which they put
on, and the resemblances between them, so as to separate the inherited
wealth of an animal's affinities with ancestors or collateral groups,
from the peculiar characters which have been acquired as an increase
based upon its typical bony possessions or osteological capital. There
is no part of the Pterodactyle skeleton which is more distinctly
modified than the head of the upper arm bone, which fits into the socket
between the coracoid bone and the shoulder-blade. The head of the
humerus, as the articular part is named, is somewhat crescent-shaped,
convex on its inner border, and a little concave on its outer border,
and therefore unlike the ball-shaped head of the upper arm bone in Man
and the higher Mammals. It is much more nearly paralleled in the little
group of Monotremata allied to the living Ornithorhynchus. In that sense
the head of the humerus in a Pterodactyle has some affinity with the
lowest Mammalia, which approach nearest to Reptiles. The character might
pass unregarded if it were not found in more striking development in
fossil Reptiles from Cape Colony, which from having teeth like Mammals
are named Theriodontia. In several of those South African reptiles the
upper arm bone approaches closer to the humerus in Ornithosaurs than to
Ornithorhynchus. Such coincidences of structure are sometimes dismissed
from consideration and placed beyond investigation by being termed
adaptive modifications; but there can be no hope of finding community of
habit between the burrowing Monotreme, the short-limbed Theriodont, and
the flying Pterodactyle which might have caused this articular part of
the upper arm bone to acquire a form so similar in animals constructed
so differently. If the resemblance in the humerus to Monotremes in this
respect is not to be attributed to burrowing, neither can the crescent
form of its upper articulation be attributed to flight; for in Birds the
head of the bone is compressed, but always convex, and Bats fly without
any approach to the Pterodactyle form in the head of the humerus. This
apparently trivial character may from such comparisons be inferred to be
something which the way of life of the animal does not sufficiently
account for. These deepest-seated parts of the limbs are slow to adapt
themselves to changing circumstances of existence, and retain their
characters with moderate variation of the bones in each of the orders
or classes of animals. It therefore is safer to regard Mammalian
characters, as well as the resemblances which Pterodactyles show to
other kinds of animals, as due to inheritance from a time when there was
a common stock from which none of these animals which have been
considered had been distinctly elaborated.
A few characters of Ornithosaurs are regarded as having been acquired,
because they are not found in any other animals, or have been developed
only in a portion of the group. The most obvious of these is the
elongated wing finger; but in some genera, like Dimorphodon, there is
also a less elongation of the fifth digit of the foot, and perhaps in
all genera there is a backward development of the first digit of the
hand, which is without a claw, and therefore unlike the clawed digit of
a Bat. An acquired character of another kind, which is limited to the
Cretaceous genera, is seen in the shoulder-blade being directed
transversely outward, so that its truncated end articulates by a true
joint with the early vertebrae of the back, and defended the cavity
inclosed by the ribs by a strong bony external arch. And finally, as the
animals later in time acquire short tails, and relatively longer limbs,
the bones of the back of the hand, termed metacarpals, acquire greater
and distinctive length, which is not seen in the long-tailed types like
Rhamphorhynchus.
These and such-like acquired characters distinguish the class of animals
from all groups with which it may be compared, and mark the possible
limits of variation of the skeleton within the boundary of the order.
But no further variation of these parts of the skeleton could make a
transition to another order of animals, or explain how the
Pterodactyles came into existence, because the characters which separate
orders and classes of animals from each other differ in kind from those
which separate smaller groups, named genera and species, of which the
order is made up. The accumulation of the characters of genera will not
sum up into the characters of an order or class.
In making the division of Vertebrate animals into classes the skeleton
is often almost ignored. Its value is entirely empirical and based upon
the observed association of the various forms of bones with the more
important characters of the brain and other vital organs. What is
understood as a Mammalian or Avian character in the skeleton is the form
of bone which is found in association with the soft vital organs which
constitute an animal a Mammal or a Bird.
The characters which theoretically define a Mammal appear to be the
enormous overgrowth of the cerebral hemispheres of the brain by which
the cerebrum comes into contact with the cerebellum, as among Birds.
This character distinguishes both groups of animals from all Reptiles,
recent and fossil. But in examining the mould of the interior of the
brain case it is rare to have the bones fitting so closely to the brain
as to prove that the lateral expansion below the cerebrum and cerebellum
is formed by the optic lobes of the brain. Otherwise the brain of a
Pterodactyle might be as like to the brain of Ornithorhynchus as it is
like that of a Bird (Fig. 19). But it is precisely in this condition of
arrangement of the parts of the brain that the specimens appear to be
most clear. The lateral mass of brain in specimens of Ornithosaurs from
the Lower Secondary rocks appears to be transversely divided into back
and front parts, which may be thought to correspond to the structures in
a Mammal brain named _corpora quadrigemina_, but to be placed as the
optic lobes are placed in Birds, and to have relatively greater
dimensions than in Mammals. No evidence has been observed of this
transverse division of the optic lobes of the brain in Pterodactyles
from the Chalk and Cretaceous rocks, and so far as the evidence goes
this part of the brain was shaped as in birds, but rather smaller.
The brain is the only soft organ in which a Mammalian character could be
evidenced. The uniformity in character of the brain throughout the group
in Mammals is remarkable, in reference to the circumstance that the
reproduction varies in type; the lowest, or Monotreme division, being
oviparous. If there is no necessary connexion between the Mammalian
brain and the prevalent condition under which the young are produced
alive, it may be affirmed also that there is no necessary connexion
between the form of the brain and the form of the bones, since the brain
cavity in Theriodont reptiles shows no resemblance to that of a Mammal,
while the bones are in so many respects only paralleled among
Monotremata and Mammalia. The variety of forms which the existing
Mammalian orders of animals assume, shows the astonishing range of
structure of the skeleton which may coexist with the Mammalian brain.
And therefore we are led to the conclusion that any other fundamental
modification of brain--such as distinguishes the class of Birds--might
also be associated with forms and structures of the skeleton which
would vary in similar ways. In other words, if for convenience we define
a Mammal by its form of brain, structure of the heart and lungs, and
provision for nutrition of the young, without regard to the covering of
the skin, which varies between the scales of a pangolin and the
practically naked skin of the whale--a bird might be also defined by its
peculiar conditions of brain and lungs, without reference to the
feathered condition of the skin, though the feathered condition extends
backward in time to the Upper Secondary rocks, as seen in the
Archaeopteryx.
The Avian characters of Pterodactyles are the predominant parts of their
organisation, for the conditions of the brain and lungs shown by the
moulds of the brain case and the thin hollow bones with conspicuous
pneumatic foramina, give evidence of a community of vital structures
with Birds, which is supported by characters of the skeleton. If any
classificational value can be associated with the distribution of the
pneumatic foramina as tending to establish membership of the same class
for animals fashioned on the same plan of soft organs, the evidence is
not weakened when a community of structures is found to extend among the
bones to such distinctive parts of the skeleton as the sternum,
shoulder-girdle, bones of the fore-arm and fore-leg; for in all these
regions the Pterodactyle bones are practically indistinguishable from
those of Birds. This is the more remarkable because other parts of the
skeleton, such as the humerus and pelvis, show a partial resemblance to
Birds, while the parts which are least Avian, like the neck bones, have
no tendency to vary the number of the vertebrae, in the way which is
common among Birds, following more closely the formula of the seven
cervical vertebrae of Mammals.
It would therefore appear from the vital community of structures with
Birds, that Pterodactyles and Birds are two parallel groups, which may
be regarded as ancient divergent forks of the same branch of animal
life, which became distinguished from each other by acquiring the
different condition of the skin, and the structures which were developed
in consequence of the bony skeleton ministering to flight in different
ways; and with different habit of terrestrial progression, this extinct
group of animals acquired some modifications of the skeleton which Birds
have not shown. There is nothing to suggest that Pterodactyles are a
branch from Birds, but their relation to Birds is much closer, so far as
the skeleton goes, than is their relation with the flightless Dinosaurs,
with which Birds and Pterodactyles have many characters in common.
On the theory of elimination of character which I have used to account
for the disappearance of some Mammalian characters from the
Pterodactyle, that loss is seen chiefly in the removal of the parts
which have left a Reptilian articulation of the lower jaw with the
skull, and the articulation of the vertebrae throughout the vertebral
column by a modified cup-and-ball form of joint. The furculum of the
Bird is always absent from the Pterodactyle. No specimen has shown
recognisable clavicles or collar-bones. Judged by the standard of
existing life, Pterodactyles belong to the same group as Birds, on the
evidence of brain and lungs, but they belong to a different group on
account of the dissimilar modifications of the skeleton and apparent
absence of feathers from the skin.
The most impressive facts in the Pterodactyle skeleton, in view of these
affinities, are the structures which it has in common with Reptiles.
Some structures are fundamental, like the cup-and-ball articulation of
the vertebrae, which is never found in birds or mammals. Although not
quite identical with the condition in any Reptile, this structure is
approximately Lizard-like or Crocodile-like in the cup-and-ball
character. It shows that the deepest-seated part of the skeleton is
Reptile-like, though it may not be more Reptilian than is the vertebral
column of a Mammal, if comparison is made between Mammals and extinct
groups of animals known as Reptiles, such as Dinosaurs and Theriodontia.
The orders of animals which have been included under the name Reptilia
comprise such different structural conditions of the parts of the
skeleton which may be termed reptilian in Ornithosaurs, that there is
good reason for regarding the cup-and-ball articulation as quite a
distinctive Reptilian specialisation, in the same sense that the
saddle-shaped articulation between the bodies of adjacent vertebrae in a
bird is an Avian specialisation. From the theoretical point of view the
Ornithosaur acquired its Reptilian characters simultaneously with its
Avian and Mammalian characters.
There is nothing in the structure of the skeleton of the Dinosauria, to
which Ornithosaurs approximate in several parts of the body, which would
help to explain the cup-and-ball articulation of the backbone, if the
Flying Reptile were supposed to be an offshoot from the carnivorous
Dinosaurs.
The elimination of Reptile characters from so much of the skeleton, and
the substitution for them of the characters of Birds and Mammals, would
be of exceptional interest if there had been any ground for regarding
the flying animal as more nearly related to a Reptile than to a Bird.
But if the evidence from the form of the brain and nature of the
pneumatic organs seen in the limb bones accounts for the Avian features
of the skeleton, the Reptilian condition of the vertebral column helps
to show a capacity for variation, and that the fixity of type and
structure, which the skeleton of the modern Bird has attained, is not
necessarily limited to or associated with the vital organs of Birds.
The variation of the cup-and-ball articulation in the neck of a
Chelonian, which makes the third vertebra cupped behind, the fourth
bi-convex, the fifth cupped in front, and the sixth flattened behind,
shows that too much importance may be attached to the mode of union of
these bones in Serpents, Crocodiles, and those Lizards which have the
cup in front; for while in Lizards the anterior cup, oblique and
depressed, is found in most of its groups, the Geckos show no trace of
the cup-and-ball structure, and in that respect resemble the Hatteria of
New Zealand.
If, therefore, the cup-and-ball articulation of vertebrae in
Ornithosauria has any significance as a mark of affinity to Reptiles, it
could only be in approximation to those living Reptiles which possess
the same character, and would have it on the hypothesis that both have
preserved the structure by descent from an earlier type of animal. This
hypothesis is negatived by the fact that the cup-and-ball articulation
is unknown in the older fossil Reptiles.
Although the articulation for the lower jaw with the skull in
Ornithosaurs is only to be paralleled among Reptiles, the structure is
adapted to a brain case which is practically indistinguishable from that
of a Bird, except for the postorbital arch.
The hypothesis of descent, therefore, becomes impossible, in any
intelligible form, in explanation of distinctive character of the
skeleton. The hypothesis of elimination may also seem to be
insufficient, unless the potential capacity for new development be
recognised as concurrent, and as capable of modifying each region of the
skeleton, or hard parts of the animal, in the same way that the soft
organs may be modified. From which we infer that all structures, which
distinguish the several grades of organisation in modern
classifications, soft parts and hard parts alike, may come into
existence together, in so far as they are compatible with each other, in
any class or ordinal division of animals.
Although the young Mammal passes through a stage of growth in which the
brain may be said to be Reptilian, there is no good ground for inferring
that Mammal or Bird type of skeleton was developed later in time than
that of Reptiles. The various types of Fishes have the brains in general
so similar to those of Reptiles that it is more intelligible for all the
vertebrate forms of brain to have differentiated at the same time, under
the law of elimination of characters, than that there should be any
other bond of union between the classes of animals.
If we ask what started the Ornithosauria into existence, and created the
plan of construction of that animal type, I think science is justified
in boldly affirming that the initial cause can only be sought under the
development of patagial membranes, such as have been seen in various
animals ministering to flight. Such membranes, in an animal which was
potentially a Bird in its vital organs, have owed development to the
absence of quill feathers. Thus the wing membrane may be the cause for
the chief differences of the skeleton by which Ornithosaurs are
separated from Birds, for the stretch of wing in one case is made by the
skin attached to the bones, and in the other case by feathers on the
skin so attached as to necessitate that the wing bones have different
proportions from Ornithosaurs.
It is a well-known observation that each great epoch of geological time
has had its dominant forms of animal life, which, so far as the earth's
history is known now, came into existence, lived their time, and were
seen no more. In the same way the smaller groups of species and genera
included in an ordinal group of animals or class have abounded, giving a
tone to the life of each geological formation, until the vitality of the
animal is exhausted, and the species becomes extinct or ceases to
preponderate. This process is seen to be still modifying the life on the
earth, when some kinds of animals and plants are introduced to new
conditions. Plants appear to wage successful war more easily than
animals. The introduction of the Cactus in some parts of Cape Colony has
locally modified both the fauna and flora, just as the Anacharis
introduced into England spread from Cambridge over the whole country,
and became for many years the predominant form of plant life in the
streams. The Rabbit in Australia is a historic pest. Something similar
to this physical fertility and increase appears to take place under new
circumstances in certain organs within the bodies of animals, by the
development of structures previously unknown. A familiar example is seen
in the internal anatomy of the Trout introduced into New Zealand, where
the number of pyloric appendages about the stomach has become rapidly
augmented, while the size and the form of the animal have changed. The
rapidity with which some of these changes have been brought about would
appear to show that Nature is capable of transforming animals more
rapidly than might have been inferred from their uniform life under
ordinary circumstances. Growth of the vital organs in this way may
modify the distinctive form of any vital organ, brain or lungs, and thus
as a consequence of modification of the internal structures due to
changes of food and habit, bring a new group of animals into existence.
And just as the group of animals ceases to predominate after a time, so
there comes a limit to the continued internal development of vital
structures as their energy fails, for each organ behaves to some extent
like an independent organism.
Under such explanations of the mutual relations of the parts of animals,
and groups of animals, time ceases to be a factor of primary importance
in their construction or elaboration. The supposed necessity for
practically unlimited time to produce changes in the vital organs which
separate animals into great orders or classes is a nightmare, born of
hypothesis, and may be profitably dismissed. The geological evidence is
too imperfect for dogmatism on speculative questions; but the nature of
the affinities of Ornithosaurs to other animals has been established on
a basis of comparison which has no need of theory to justify the facts.
It is not improbable that the primary epoch of time, even as known at
present, may be sufficiently long to contain the parent races from which
Ornithosaurs and all their allies have arisen.
In thus stating the relation of Ornithosaurs to other animals the Flying
Reptile has been traced home to kindred, though not to its actual
parents or birthplace. There is no geological history of the rapid or
gradual development of the wing finger, and although the wing membrane
may be accepted as its cause of existence, the wing finger is powerfully
developed in the oldest known Pterodactyles as in their latest
representatives.
Pterodactyles show singularly little variation in structure in their
geological history. We chronicle the loss of the tail and loss of teeth.
There is also the loss of the outermost wing digit from the hind foot as
a supporter of the wing membrane. But the other variations are in the
length of the metacarpus, or of the neck, or head. One of the
fundamental laws of life necessitates that when an animal type ceases to
adapt its organisation and modify its structures to suit the altered
circumstances forced upon it by revolutions of the earth's surface its
life's history becomes broken. It must bend or break.
The final disappearance of these animals from the earth's history in the
Chalk may yet be modified by future discoveries, but the Flying Reptiles
have vanished, in the same way as so many other groups of animals which
were contemporary with them in the Secondary period of time. Such
extinctions have been attributed to catastrophes, like the submergence
of land, so that the habitations of animals became an area gradually
decreasing in size, which at last disappeared. It appears also to be a
law of life, illustrated by many extinct groups of animals, that they
endure for geological ages, and having fought their battle in life's
history, grow old and unable to continue the fight, and then disappear
from the earth, giving place to more vigorous types adapted to live
under new conditions.
The extinct Pterodactyles hold a relation to Birds in the scheme of life
not unlike that which Monotremata hold to other Mammals. Both are
remarkable for the variety of their affinities and resemblances to
Reptiles. The Ornithosauria have long passed away; the Monotremes are
nearing extinction. Both appear to be supplanted by parallel groups
which were their contemporaries. Birds now fill the earth in a way that
Flying Reptiles never surpassed; but their flight is made in a different
manner, and the wing is extended to support the animal in the air,
chiefly by appendages to the skin.
If these fossils have taught that Ornithosaurs have a community of soft
vital organs with Dinosaurs and Birds, they have also gone some way
towards proving that causes similar to those which determined the
structural peculiarities of their bony framework, originated the special
forms of respiratory organs and brain which lifted them out of
association with existing Reptiles.
These old flying animals sleep through geological ages, not without
honour, for the study of their story has illuminated the mode of origin
of animals which survive them, and in cleaving the rocks to display
their bones we have opened a new page of the book of life.
APPENDIX
The best public collections of Ornithosaurian remains in England are
in the British Museum (Natural History); Museum of Practical Geology,
Royal College of Surgeons; the University Museum, Oxford; Geological
Museum, Cambridge; and the Museum of the Philosophical Society at
York.
Detailed descriptions and original figures of the principal specimens
mentioned or referred to may be found in the following writings:--
H. v. Meyer, _Reptilien aus dem Lithograph_. _Schiefer_. 1859. Folio.
v. Quenstedt, _Pterodactylus suevicus_. 1855. 4to.
Goldfuss, _Nova Acta Leopold_. XV.
v. Munster, _Nova Acta Leopold_. XV.
A. Wagner, _Abhandl. Bayerischen Akad._, vi., viii.
Cuvier, _Annales du Museum_, xiii. 1809.
" _Ossemens fossiles_, v. 1824.
Buckland, _Geol. Trans._, ser. 2, iii.
R. Owen, _Palaeontographical Society_. 1851, 1859, 1860, 1870, 1874.
K. v. Zittel, _Palaeontographica_, xxix. 1882.
T. C. Winkler, _Mus. Teyler Archives_. 1874, 1883.
Oscar Fraas, _Palaeontographica_, xxv. 1878.
Anton Fritsch, _Boehm. Gesell. Sitzber_. 1881.
R. Lydekker, _Catalogue of Fossil Reptilia in British Museum_ I. 1888.
O. C. Marsh, _Amer. Jour. Science_. 1882, 1884.
S. W. Williston, _Kansas University Quarterly_. 1893, 1896.
E. T. Newton, _Phil. Trans. Royal Soc._ 1888, 1894.
H. G. Seeley, _Ornithosauria_. 8vo. 1870.
" _Annals and Mag. Natural Hist._ 1870, 1871, 1890, 1891.
" _Linn. Society_. 1874, 1875.
" _Geol. Mag._ 1881.
Felix Pleininger, _Palaeontographica_. 1894, 1901.
INDEX
A
Abdominal ribs, 85, 154
Accumulation of characters, 220
Acetabulum, 95
Acquired characters, 219
Adjacent land, 136
Air cells, 10, 48
Albatross, 23, 36, 176
Alligator, brain, 53;
pelvis, 98
American Greensand, 185
-- ornithosaurs, 87, 126
Amphibia, 4, 191
Anabas, 17
Anacharis, 227
Anchisaurus, 199
Angle of lower jaw, 75
Ankle bones, 103, 195, 207
Anomodonts, 192
Ant-eater of Africa, 142;
India, 40;
South America, 40, 185
Apteryx, lungs, 48;
pelvis, 95
Aquatic mammals, 141
Aramis, scapular arch, 113
Archaeopteryx, 58, 76, 104, 130, 197, 211
Aristosuchus, 129, 190, 205, 209
Armadillo, 40, 141
Articulation of the jaw, 12, 75
Ashwell, 177
Atlantosaurus, 202
Atlas and axis, 80, 81
Aves, 190
Avian characters, 220, 222
B
Backbone, 78, 84
Banz, 148
Barbastelle, 25
Barrington, 177
Barton, 177
Bat, 38, 110, 197;
sternum of, 107;
metacarpus, 128
Bavaria, 156, 185
Beak, horny, 74, 178
Bear, skull of, 12;
femur, 100
Bel and the Dragon, 15
Belodon, 202
Bird, 80, 110, 120
-- resemblances, 63, 65, 71, 95, 102, 108, 113, 119, 120, 211
Bird-reptile, 188
Bird wing, 128, 130
Birds in flight, 22;
with teeth, 76
Black-headed bunting, 47
Blainville, D. de, 30, 193
Blood, temperature of, 56
Bohemia, 34
Bonaparte, Prince Charles, 30
Bones of birds, variation in, 41
-- of reptiles, variation in, 42
-- about the brain, 69
-- in the back, 84
Bone texture, 59, 209
Bonn Museum, 32, 85, 156
Brain and breathing organs, 55
Brain cavity, in birds and reptiles, 52;
in mammals, 221, 226;
in Solenhofen pterodactyles, 54, 220
Brazil, 34
Breathing organs, 8
Bridgewater Treatise, 143
British Museum, 133, 183
Brixton, Isle of Wight, 55, 174
Buckland, Dean, 143, 148, 231
Burrowing limb, 38
C
Cactus, 227
Calamospondylus, 203
Cambridge Greensand, 33, 89, 176
-- Museum, 177
Camel, 83
Campylognathus, 68, 71, 135;
size of, 149
Canary, 47
Carnivorous dinosaurs, 129
Carpus, 122
Caudal fin, 91, 161
-- vertebrae, 89, 92, 203
Ceratodus, 4, 5, 9, 17
Ceratosaurus, 203, 204
Cervical rib, 81
Cetacea, 40
Cetiosaurus, 198, 203
Chalinolobus, 25
Chalk, pterodactyles in, 136;
of Kansas, 103, 132
Chameleon, 17, 51, 70;
scapula, 112;
sternum, 107
Chameleonoidea, 191
Cheek bones, 178
Chelonia, 86, 112, 193
Chesterton, 177
Chlamydosaurus, 21
_Chrysochloris capensis_, 121
Classification, 192;
on pelvis characters, 195;
of dinosaurs, 198
Clavicles, 111, 112
Claw, 105, 116, 183, 208
Coelurus, 203, 209
Coldham Common, 177
Collar bone, 111
Collini, 27
Comparison with dinosaurs, 198;
of pelvis, 204, 206;
of skulls, 192, 199, 201
Cope, Professor, 31, 34
Coracoid, 109, 112, 113
Cordylomorpha, 191
Cormorant, 70, 174;
sternum, 108
Corpora quadrigemina, 221
Crisp, Dr., on pneumatic skeleton, 47
Crocodile, characters of, 217;
heart, 56;
lung, 9;
shoulder-girdle, 111;
skull, 46;
vertebrae, 79
Crocodilia, 190
Curlew, 68
Cuvier, 1, 27, 28, 54, 76, 77, 130, 231
Cycnorhamphus, 70, 94, 171, 173, 204
_Cycnorhamphus Fraasii_, 80, 96, 169
-- _suevicus_, 169, 170
Cypselus, 42
D
_Dacelo gigantea_, 63
Darwin, 3
Davy, Dr. John, 142
Deuterosaurus, 97
Dicynodon, 200
_Dicynodon lacerticeps_, 71
Digits, of ostrich, 23;
of pterodactyle, 128
Digits with claws, 130;
foot bones in, 105
Dimorphodon, 63, 64, 66, 67, 73, 74, 83, 90, 102, 113, 143, 192, 194,
199, 201, 206
Dinosauria, 6, 77, 84, 87, 95, 129, 144, 198, 209
Dinosaurs from Lias, 135, 192;
from Elgin, 201, 207;
Stuttgart, 202;
Trias dinosaurs, 199, 200
Diopecephalus, 168
Diving birds, 23, 83, 102
Dolichosauria, 191
Dolphin, 107
Doratorhynchus, 173
Dorygnathus, 74, 148
Dragons, 3, 15, 17
Drumstick bone, 103, 195
Duck, 22, 83
E
Echidna, 75, 76, 95, 100
Edentata, 185
Edentulous beak, 153
Eichstaedt, 32
Elephant, head of, 46
Enumeration of characters, 223, 225
Ephesus, winged figure, 16
Epiphysis to first phalange, 123
Exocoetus, 18
Extinctions, 129
Eye hole, 144;
sclerotic bones in, 65
F
Farren, William, 34
Femur, 100
Fibula, 102, 183, 206
Fifth outer digit, 132;
in foot, 145
Figure from temple at Ephesus, 16
First phalange, 151
Fish-eating crocodile, 137
Flight, organs of, 17;
in bats, 25
Flying limb, 38
Flying fishes, 18, 57;
foxes, 26;
frogs, 19, 197;
gecko, 21, 24;
lizards, 20;
reptiles, 37, 46;
squirrel, 24
Foot, 104;
digits in, 105, 146
Fore leg, 102, 206
-- limb, 38, 107, 116, 120
Four claws, 147
Fox, Rev. W., 55, 174
Fraas, Professor Oscar, 172, 231
Frigate bird, vertebrae of, 86, 174
Frog, lungs of, 8
Furculum, 114
G
Gaudry, Professor A., 31
Gavial, 136
Gecko, 21, 23
Genera, comparison of, 192
Geological distribution, 186
Gills, 4
Giraffe, 38, 39
Glossy starling, 47
Golden eagle, 120
-- mole, 121
Goldfuss, 30, 231
Granchester, 177
Great ant-eater, 40, 185
Guillemot, 102
Gull, 22
H
Haarlem, Teyler Museum at, 32
Habits, probable, 134, 176, 198
Hairless skins, 141
Hand in mammals, 38
Harston, 177
Haslingfield, 177
Hastings, 174
Hatteria lung, 9, 27;
brain, 53;
skull, 70, 77;
ribs, 86;
a reptile type, 13
Head, characters of, 76
Heidelberg Museum, 32, 54, 159
Herpetomorpha, 191
Heron, 65, 174
Hesperornis, 76
Hind foot, 104, 135
-- limb, 93, 99, 159, 206
Hip-girdle in whale tribe, 39, 159
Homoeosauria, 191
Horningsea, 177
Horse, metacarpus of, 127;
vertebrae of, 79
Humerus, 46, 117, 217
Huxley, Professor, 31, 89, 154, 188
Hyo-mandibular arch, 13
Hypothesis of descent, 226
Hyrax, 101
I
Ichthyornis, 76
Ichthyosaurus, 6, 191
Iguanodon, 209;
pelvis, 206
Ilium, 93, 95, 96, 98, 204
Instep, 105, 207
Inherited characters, 217
Interclavicle, 111
Ischium, 93, 96, 203, 204
Isle of Wight, 174
J
Jaw, in birds, 12;
in fishes, 13;
in mammals, 12;
in reptiles, 13;
in pterodactyles, 63;
suspension of, 11, 74, 76
-- lower, 75
K
Kansas, Chalk of, 72, 103, 115;
University Museum of, 181
Kelheim, 32
Keuper, 33
Kimeridge Clay, 132
Kingfisher, 63
Kiwi, 23
L
Labyrinthodontia, 191
Lachrymal bones, 67
Laramie rocks, 34
Largest ornithosaur, 133
Lateral vacuities in skull, 147
Lawrence in Kansas, 181
Lengths of bones, 146
Lepidosiren, 17
Lias, 33
Lithographic Slate, 35, 156
Lizards, 20, 21, 27, 123
Llama, neck of, 79, 83
Loach, swim bladder of, 52
Lower jaw, 12, 74, 76, 149
Lumbar vertebrae, 89
Lungs, 47;
in apteryx, 48;
in chameleon, 51;
in ostrich, 49;
in reptiles, 8, 9, 51
Lydekker, R., 160, 169, 231
Lyme Regis, 33
M
Macrocercus, palate of, 71
Malar bone, 67
Mallard, 22
Mammal, 8, 12, 24, 79, 53, 95
Mammalia, 38, 141
Mammalian characters, 12, 220
Mammoth, 141
Manis, 40, 57, 142
Manubrium of sternum, 108, 109, 183
Marrow bones in a bird, 134
Marsh, Professor O. C., 31, 72, 90, 115, 121, 131, 140, 160, 165,
180, 181, 210, 231
Marsupial, 70, 94, 99
Megalosaurus, 129, 198
Merganser, 108
Merry-thought, 114
Metacarpus, 116, 124, 126, 128, 130
Metatarsal bones, 104, 207, 208
Meyer, Hermann von, 31, 45, 46, 85, 105, 108, 121, 160, 192, 231
Moa of New Zealand, 35
Mole, humerus, 38;
sternum, 107
Monotremes, 70, 94, 111, 121, 185, 218
Mososaurus, 77
Movement of the leg, 101
Mugger, 137
Munich Museum, 32, 159
Munster, von, 231
Muschelkalk, 184
Museum, 32, 156, 231, 159;
Natural History, 133, 231
Myrmecophaga, 185
N
Names of genera, 183
Natural History Museum, 38, 231
Neck, 79;
in Dimorphodon, 145;
in Giraffe, 39;
in Llama, 79;
in Pterodactyles, 80;
in Whales, 39
Newton, E. T., 55, 70, 158, 160, 201, 232
New Zealand Bat, 25
-- -- Hatteria, 68
Niobrara rock, 183
Nostril, bones round the, 62;
small, 147
Notarium, 87, 115
Nothosauria, 192
Nusplingen, 32
Nyctodactylus, 115, 180
O
Obliteration of characters, 216
Opercular bones, 13
Ophidia, 52, 191
Optic lobes, 53, 221
Organs of flight, 17
Ornithischia, 190, 198
Ornithocephalus, 166
Ornithocheirus, atlas and axis, 81;
brain, 55, 69;
carpus, 124;
cervical vertebra, 83, 179;
claw phalange, 129;
coracoid, 109;
femur, 100;
pelvis, 98;
pubic bones, 194;
sternum, 109;
shoulder-girdle, 115;
remains, 176;
teeth, 74, 76;
absence of teeth, 138
_Ornithocheirus machaerorhynchus_, 139;
_microdon_, 139
Ornithocheiroidea, 193
Ornithodesmus, neck bones, 173, 175;
coracoid, 109, 116;
dorsal vertebrae, 86;
remains of _O. latidens_, 173;
_O. sagittirostris_, 175
Ornithomorpha, 189
Ornithorhynchus, 40, 53, 95, 117
Ornithosauria, 30, 31, 50, 52, 58, 72, 89, 95, 104, 108, 125, 132,
133, 143, 187, 190, 192, 216
Ornithostoma, 66, 69, 72, 180;
lower jaw, 75, 76;
pelvis, 98;
sternum, 110;
phalange, 122;
size, 133;
skull, 181, 182
Ornithosuchus, 201
Orycteropus, 96
_Ossa innominata_, 93
Ossified ligaments, 150
Ostrich, 23, 45, 49, 113, 129
Owen, Sir R., 31, 36, 46, 48, 110, 117, 143, 172, 176, 180, 231
Owl, 46, 53
Oxford Clay, 33, 156
-- University Museum, 154
Ox, vertebra of, 79;
metacarpus, 127
P
Palate, bones of, 71
Pangolin, 142
Pappenheim, 32
Parallel groups, 215
Parrot, 71
Patagial membranes, 227
Pelican, 174
Pelvis, 88, 94-98, 151, 195, 202, 204, 206
Penguin, 41, 42, 104, 176
Periophthalmus, 17
Peterborough, bones from, 113, 156
Phalanges, 129, 132;
wing finger, 155
Phillips, Professor John, 155
Pigeon, 119
Platydactylus, 21
Platypus, 214
Plesiosaurus, 6, 73, 75, 93, 189
Pleininger, 149, 232
Pneumatic foramina, 45, 83, 88, 132, 209
Pond, Mr., 34
Porcupine, 40
Porpoise, 38, 73, 141, 200
Premaxillary bones, 77, 200, 205
Prepubic bones, 94, 96-98, 194, 204, 205
Protorosauria, 192
_Ptenodracon brevirostris_, 64, 99, 167, 169, 192
Pterodactyle aspects, 35;
avian characters, 222;
beak, 200;
brain, 53;
coracoid, 113;
discovery, 27, 33;
foot, 104;
fore limb, 117;
history in Germany, 31, 148;
hand, 130;
hind limb, 100;
long tails, 156;
palate, 71;
sacrum, 89;
short tails, 165;
size, 35, 133;
sacrum, 89;
skull, 192;
teeth, 73;
vertebrae, 80
Pterodactyles from Kansas Chalk, 177, 181
-- from Lias Clay, 135, 147, 152
-- from Neocomian Sand, 176
-- from Oxford Clay, 155
-- from Purbeck beds, 173
-- from Solenhofen Slate, 156, 158
-- from Stonesfield Slate, 153, 158
Pterodactylia, 30, 165, 193, 199
_Pterodactylus antiquus_, 167;
_brevirostris_, 99, 167, 169;
_crassirostris_, 156;
_dubius_, 87, 96, 97, 203;
_elegans_, 169;
_Fraasii_, 169;
_grandipelvis_, 87, 90;
_grandis_, 102, 167, 169;
_Kochi_, 12, 61, 87, 90, 168, 169;
_longirostris_, 28, 90, 96, 101, 103, 105, 167, 169;
_micronyx_, 105, 169;
_rhamphastinus_, 183;
_scolopaciceps_, 105, 166;
_spectabilis_, 83;
_suevicus_, 169
Pterodermata, 194, 199
Pteroid bone of first digit, 121
Pteromys, 24
Pterosauria, 187, 193
Pterygoid bones, 72, 147
Pythonomorpha, 191
Q
Quadrate bone, 12, 68, 77
Quenstedt, 231
R
Rabbit, 227
Radius, 119, 120
Redshanks, 22
Relation between head and tail, 157, 193
Reptile, 6, 79, 80
Resin, 136
Restorations--
Campylognathus, palate of, 71
Dimorphodon, 143, 147, 164
Ornithocheirus, 164
Ornithostoma, 164, 183
Ptenodracon, 167
Pterodactylus, 29, 30
Rhamphocephalus, 164
Rhamphorhynchus, 161, 164
Scaphognathus, 163
Rhacophorus, 19
Rhaetic beds, 184
Rhamphocephalus, 113, 136, 153
Rhamphorhynchus, 118, 192;
foot, 104;
hind limb, 99;
pelvis, 95;
sacrum, 88;
skull, 54, 63-6, 69;
sternum, 108;
tail, 91;
teeth, 73;
tibia and fibula, 103;
web-footed, 105
_Rhamphorhynchus curtimanus_, 163;
_hirundinaceus_, 163;
_longimanus_, 164;
_phyllurus_, 91, 165
Rhinoceros, 40, 141
Rhopoladon, 97
Rhynchocephala, 192
Roc, 36
Rochester, 136
Running limb, 38
Ryle, Bishop, 17
S
Sacrum, 87, 88
St. George, 15
St. Ives, 156
Sarcorhamphus, 102
Saurians, 27
Saurischia, 190, 195, 198, 199
Sauromorpha, 191, 192
Sauropsida, 188
Sauropterygia, 192
Scaphognathus, 64, 85, 140, 152, 192, 212
_Scaphognathus crassirostris_, 73-5, 83
Scapular arch, 111, 113
Scelidosaurus, 135
Sclerotic circle, 65
Seals, 41
Sedgwick, Professor Adam, v, 46
Shillington, 77
Shoebill, 67
Shoe-shaped prepubic bones, 204, 205
Short-tailed pterodactyles, 165, 193
Shoulder-girdle, 107, 111, 114, 115, 183
Siberia, 141
Simultaneous origin of characters, 214, 224
Skin covering, 40, 41, 58, 139, 140
Skulls, 68
Sloth, 112
Snipe, 47, 68
Solenhofen Slate, 28, 32, 88, 153, 156
Soemmerring, 29
South African reptiles, 188, 208, 216
Spotted fly-catcher, 47
Squamosal bone, 12, 13
Sternal ribs, 110
Sternum, 107, 158
Stonesfield Slate, 33, 88, 153
Structures common to reptiles, 224
Stuttgart Museum, 32, 172, 203
Swanage, 172
Swan, neck of, 80, 113
Swift, 50
Swimming limb, 38
Synotus, 25
Syrinx, 48
T
Tail, description of, 90;
in Cretaceous Pterodactyles, 193
-- long, 156;
short, 166;
in Dimorphodon, 145;
in Ornithocheirus, 179
Tanystrophoeus, long vertebrae in, 79
Tarsal bones, 102, 207
Tarso-metatarsus, 128
Teeth, 73, 137, 138;
in porpoise, 40
Temperature of blood, 56
Temporal arches, 68
-- bone, 12
-- fossa, 67
Teredo, 137
Texas fossils, 216
Thecospondylus, 209
Theriodont pelvis, 97
-- reptiles, 75;
of Russia, 96, 97;
of South Africa, 96, 117
Theropsida, 188
Thigh bone, 100, 206, 211
Three claws, 146, 197
Tibia, 102, 195;
in Iguanodon, 207
Toothless mammals, 40
-- pterodactyles, 138, 181;
beak of pterodactyles, 150
Transition from reptiles to birds, 211
Tree frogs, 21
Trias dinosaurs, 199
Triceratops, pelvis of, 204
Trout, 139;
of New Zealand, 228
Tuatera, 13
Tuebingen Museum, 32
Tundras, 141
Tunny, 57
Turtles, neck bones, 79
U
Ulna, description of, 119
Uncinate process of ribs, 85
Unlimited time, 228
Upper arm bone, 117
-- Greensand, remains in, 136
-- Lias of Whitby, 147
-- Oolites, 185, 195
V
Variation of bones in mammals, 38
-- in Pterodactyles, 229
Variation of bones in vertebrae, 225
Vertebrae, caudal, 89, 92, 203
-- cervical, 173, 179, 203
-- dorsal, 86
Vertebral articulation, 82, 224
-- column, 78
Vulture, neck vertebrae of, 80;
tibia and fibula of, 102
Vomer, 147
Vomerine bones, 72
W
Wagler, 29
Wagner, Andreas, 30, 148, 231
Walker, J. F., 54
Wealden beds, Pterodactyles in, 55, 84;
bones in, 135, 136, 173
Weight of Pterodactyle, 106
Whinchat, 47
Whitby, 33, 135
Williston, Professor W. S., 75, 82, 92, 98, 105, 110
Willow-wren, 47
Wing finger, 116, 130, 133, 151, 178, 197
-- membrane, 32, 121, 140, and frontispiece
-- metacarpal, 123;
in Dimorphodon, 151;
in Ornithostoma, 184;
in bats, 131
Wings of Dragons, 16
Winkler, T. C., 231
Woodwardian Museum, 34
Wood-wren, 47
Wrist bones, 122
Wuertemberg, 33
Y
Yale College Museum, 32
York Museum, 34, 176
Z
Zittel, Karl von, 31, 157, 165, 231
Zygomatic arch, 67
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