Comparative anatomy of

vertebrate skeleton
Bone
inorganic components of bone comprise 60% of the dry weight
(largely calcium hydroxy-appetite crystals) & provide the compressive
strength of bone. The organic component is primarily collagen, which
gives bone great tensile strength.
provides support and movement via attachments for soft tissue and
muscle, protects vital organs, is a major site for red marrow for
production of blood cells, and plays a role in the metabolism of
minerals such as calcium and phosphorus.
 There are two basic structural types of
bone, compact and spongy. Compact bone
forms the outer shell of all bones and also
the shafts in long bones. Spongy bone is
found at the expanded heads of long bones
and fills most irregular bones
Bone formation begins with a blastema
(any aggregation of embryonic
mesenchymal cells which will differentiate
into tissue such as muscle, cartilage, or
bone). These cells then develop into either
FIBROBLASTS or OSTEOBLASTS. Fibroblasts
form collagen; osteoblasts form bone cells.
Together, these form MEMBRANE BONE
(bone deposited directed in a blastema).
Intramembranous ossification
is the process of membrane bone formation. This process give rise to:
bones of the lower jaw, skull, & pectoral girdle
dentin & other bone that develops in the skin
vertebrae in some vertebrates (teleosts, urodeles, & apodans)
Endochondral ossification
is the process in which bone is deposited in
pre-existing cartilage, &
such bone is called REPLACEMENT
BONE.
Skeletal elements:
Dermal skeleton
skin of most living vertebrates has
no hard skeletal parts but dermal
bone elements are usually present
in the head region
early vertebrates (ostracoderms)
had so much dermal bone they
were called 'armored fishes'

after ostracoderms, fish continued

to develop much bone in skin but
that bone has become 'thinner'
over time
Endoskeleton

Somatic - axial & appendicular

skeletons
Visceral - cartilage or bone
associated with gills & skeletal
elements (such as jaw cartilages)
derived from them
Dermal bone of fishes:
Basic structure includes lamellar (compact) bone, spongy
bone, dentin, &, often, a surface with a layer of enamel-like
material
Evolutionary 'trend' = large bony plates giving way to smaller,
thinner bony scales
Ancient armor - not found on living fish
Ganoid scales - found only on Latimeria (coelocanth) & sturgeons
Placoid scales - elasmobranchs (diagram to the right; pulp cavity >
dentin layer > enamel)
Ctenoid & Cycloid scales - modern bony fish
Tetrapods - retain dermal elements in the skull, jaws, &
pectoral girdle
1 = lamellar bone, 2 = spongy bone, 3 = dentin, 4 = enameloid,
& 5 = fibrous plate (collagen)
 Somatic skeleton = axial skeleton (vertebral column,
ribs, sternum, & skull) + appendicular skeleton
Vertebral column:
Vertebrae - consist of a centrum (or body), 1 or 2 arches, plus various processes
Amphicelous
concave at both ends
most fish, a few salamanders (Necturus), & caecilians
Opisthocoelous
convex in front & concave in back
most salamanders
Procelous
concave in front & convex in back
anurans & present-day reptiles
Acelous
flat-ended
mammals
Heterocelous
saddle-shaped centrum at both ends
birds
Vertebral arches:

Neural arch - on top of

centrum
Hemal arch (also called
chevrons) - beneath centrum
in caudal vertebrae of fish,
salamanders, most reptiles,
some birds, & many long-tailed
mammals
Vertebral processes:
projections from arches & centra
some give rigidity to the column, articulate
with ribs, or serve as sites of muscle
attachment
Transverse processes - most common type of
process; extend laterally from the base of a
neural arch or centrum & separate the epaxial
& hypaxial muscles
Diapophyses & parapophyses - articulate with
ribs
Prezygapophyses (cranial zygapophyses) &
postzygapophyses (caudal zygapophyses) -
articulate with one another & limit flexion &
torsion of the vertebral column
 Vertebral columns:
Cartilaginous fishes
do not have typical fish vertebral columns
vertebrae include neural arches (cartilaginous
dorsal plates) & dorsal intercalary plates are
located between successive arches
Teleosts
well-ossified amphicelous vertebrae
notochord persists within each centrum (but
constricted)
neural arch associated with each centrum & hemal
arches in tail (caudal) vertebrae
Chondrosteans (sturgeons & paddlefish) &
modern lungfishes
incomplete centra
notochord is not constricted
cartilage deposited in notochord sheath provides
structural support
Vertebral columns of tetrapods
Cervical region
Amphibians- single cervical vertebra; allows little head movement
Reptiles - increased numbers of cervical vertebrae (usually 7) & increased
flexibility of head
Birds - variable number of cervical vertebrae (as many as 25 in swans)
 Mammals - usually 7 cervical vertebrae

Reptiles, birds, & mammals - 1st two cervical

vertebrae are modified & called the atlas & axis
atlas - 1st cervical vertebra; ring-like (most of
centrum gone); provides 'cradle' in which skull
can 'rock' (as when nodding 'yes')
axis - 2nd cervical vertebra

Transverse foramen (#6 in above caudal

view of a cervical vertebra)
found in cervical vertebrae of birds &
mammals
provides canal for vertebral artery & vein
 Dorsal region
Dorsals - name given to
vertebrae between cervicals &
sacrals when all articulate with
similar ribs (e.g., fish,
amphibians, & snakes)
Crocodilians, lizards, birds, &
mammals - ribs are confined to
anterior region of trunk
thoracic - vertebrae with ribs
lumbar - vertebrae without ribs

 Sacrum & Synsacrum

sacral vertebrae - have short transverse
processes that brace the pelvic girdle &
hindlimbs against the vertebral column
Amphibians - 1 sacral vertebra
Living reptiles & most birds - 2 sacral vertebrae
Most mammals - 3 to 5 sacral vertebrae
Sacrum - single bony complex consisting of
fused sacral vertebrae; found when there is
more than 1 sacral vertebra (see examples
below):

Synsacrum
found in birds
produced by fusion of last thoracics, all
lumbars, all sacrals, & first few caudals
fused with pelvic girdle
provides rigid support for bipedal locomotion

 Caudal region
Primitive tetrapods - 50 or more caudal
vertebrae
Present-day tetrapods
number of caudal vertebrae is reduced
arches & processes get progressively shorter (the
last few caudals typically consist of just cylindrical
centra as shown below)
Anurans - unique terminal segment called
the urostyle (section of unsegmented vertebral
column probably derived from separate
caudals of early anurans)
Birds - last 4 or 5 caudal vertebrae fused to
form pygostyle (see drawing above)
Apes & humans - last 3 to 5 caudal vertebrae
fused to form coccygeal (or tail bone)

Ribs - may be long or short, cartilaginous or bony;

articulate medially with vertebrae & extend into
the body wall
A few teleosts - have 2 pair of ribs for each centrum of trunk
(dorsal rib separates epaxial & hypaxial muscles)
Most teleosts - ventral ribs only
Sharks - dorsal ribs only
Agnathans - no ribs
Tetrapods - ribs usually articulate with vertebrae in moveable joints
(see above drawing)
Early tetrapods - ribs articulated with every vertebra from the
atlas to the end of the trunk
Later tetrapods - long ribs limited to thoracic region
Thoracic ribs - most composed of a dorsal element
(vertebral rib) & a ventral element (sternal rib)
Sternal rib - may be ossified (birds) or remain
cartilaginous (mammals); usually articulate with
sternum (except 'floating ribs')
Uncinate processes - found in birds; provides rib-cage
with additional support
 Sternum - strictly a tetrapod structure &,
primarily, an amniote structure.
Amphibians - no sternum in
early amphibians &, among
present-day amphibians,
only anurans have one
Amniotes
sternum is a plate of cartilage
& replacement bone
sternum articulates with the
pectoral girdle anteriorly &
with a variable number of ribs
SKULL
Consist of:
1 - neurocranium (also called endocranium or primary braincase)
2 - dermatocranium (membrane bones)
3 - splanchnocranium (or visceral skeleton)
 Neurocranium
1 - protects the brain
2 - begins as cartilage that is partly or entirely
replaced by bone (except in cartilaginous
fishes)
Cartilaginous stage:
neurocranium begins as pair of parachordal &
prechordal cartilages below the brain
parachordal cartilages expand & join; along with the
notochord from the basal plate
prechordal cartilages expand & join to form
an ethmoid plate
Cartilage also appears in the
olfactory capsule (partially surrounding the olfactory
epithelium)
otic capsule (surrounds inner ear & also develops into
sclera of the eyeball)
 Completion of floor, walls, & roof:
Ethmoid plate - fuses with olfactory capsules
Basal plate - fuses with otic capsules
Further development of cartilaginous neurocranium =
development of cartilaginous walls (sides of braincase)
&, in cartilaginous fishes, a cartilaginous roof over the
brain
Cartilaginous fishes - retain a cartilaginous neurocranium (or
chondrocranium) throughout life
Bony fishes, lungfishes, & most ganoids - retain highly
cartilaginous neurocranium that is covered by membrane bone
Cyclostomes - the several cartilaginous components of the
embryonic neurocranium remain in adults as more or less
independent cartilages
Other bony vertebrates - embryonic cartilaginous neurocranium
is largely replaced by replacement bone (the process of
endochondral ossification occurs almost simultaneously at
several ossification centers)
DERMATOCRANIUM - lies superficial to
neurocranium & forms:
1 - bones that form the roof of the brain & contribute
to the lateral walls of the skull
2 - bones of the upper jaw
3 - bones of the palate(s)
4 - opercular bones
 Bones of the upper jaw
Pterygoquadrate (palatoquadrate) cartilage - 1st upper jaw that
vertebrate embryos develop
Cartilaginous fishes - palatoquadrate is the only upper jaw that develops
Bony vertebrates - the palatoquadrate becomes covered with dermal bones
(premaxillae & maxillae) that make up the adult upper jaw

 Palatal bones - the floor on which the brain rests is at the same time
the roof of the oral cavity in fishes & amphibians (primary
palate)Sharks - cartilaginous
bony vertebrates - membrane bones form
Birds, mammals, & some reptiles - a secondary ('false')
palate develops creating a horizontal partition that separates the oral
cavity into nasal & oral passages. The secondary palate is formed
from processes of the premaxillae, maxillae, and palatines.

 Opercular bones
Operculum = fold of the hyoid
arch that extends back over the
gill slits in holocephalans &
bony fishes
Tetrapods - no vestiges of
opercular bones remain
VISCERAL SKELETON or Splanchnocranium
skeleton of the pharyngeal arches
Fishes - skeleton of the jaws & gill arches
Tetrapods - skeleton modified for new functions
 Fish visceral skeleton - consists of 7 sets of paired cartilages in the 7
visceral arches & a series of mid-ventral cartilages (basihyal &
basibranchials) in the pharyngeal floor
Bony fishes
visceral skeleton resembles that of sharks except that bone is added
caudal ends of the cartilaginous pterygoquadrate undergo
endochondral ossification & become the quadrate bones. The
remainder becomes the palatine & pterygoid bones. The posterior tip
of Meckel's cartilage becomes an articular bone.
Jaw suspension of fishes
The jaw-hyoid complex of fishes requires bracing against some support to
function effectively, and the nearest one is the neurocranium
(endocranium).
Types of suspensions:
autostyly (below left) - hyomandibula play no role in bracing the jaws (lungfish &
tetrapods)
amphistyly (below middle) - jaws & hyomandibula both braced directly against the
braincase (extinct sharks)
hyostyly (below right) - mandibular cartilage is braced against the otic capsule; jaws
braced against hyomandibula (sharks & present-day bony fishes)
Tetrapods
With life on land (& pulmonary respiration), the visceral skeleton
underwent substantial modification. Some structures were lost &
others remained to perform new functions.
 Pterygoquadrate (palatoquadrate) cartilage = embryonic upper jaw
cartilage
Amphibians, reptiles, & birds - posterior end undergoes endochondral
ossification & becomes the quadrate (which articulates with the articular
bone of the lower jaw)
Mammals - dentary (lower jaw) articulates with the squamosal of skull
(quadrate separates from the rest of the palatoquadrate & becomes the incus
of the middle ear)
 Meckel's cartilage
Reptiles - largely ensheathed by dermal bones
Birds & mammals - few or no remnants in adult lower jaw (&, in mammals,
the articular, formed by ossification of the tip of Meckel's cartilage, projects
into the middle ear cavity & becomes the malleus)
 Arch II = Hyomandibular cartilage:
Sharks - interposed between quadrate region &
otic capsule
Tetrapods - no longer articulates with quadrate &
ossifies to become part of the stapes (columella)
Arches III ---> V become part of hyoid
apparatus
Arches VI & VII - not present in tetrapods
Hyoid apparatus of tetrapods
consists of a body & 2 or 3 horns (cornua)
anchors tongue, provides attachment for some
extrinsic muscles of larynx, & is site of
attachment of muscles that aid in swallowing

Appendicular skeleton
consists of pectoral & pelvic girdles plus skeleton of fins & limbs
Some vertebrates have no appendicular skeleton (e.g., agnathans, apodans,
snakes, & some lizards) & in others it is much reduced.

Pectoral girdles:
1 - brace for anterior appendages
2 - consist of membrane & replacement bones (in bony vertebrates)
3 - Early fishes had 3 replacement bones (coracoid, scapula, & suprascapula) and
a series of dermal bones (clavicle, cleithrum, supracleithrum, and post-temporal)
4 - Later bony fishes (ganoid fish) - tendency for reduction in number and size of
replacement bones
5 - Tetrapods - tendency for reduction in number of dermal bones
 Tetrapods - early ones had pectoral girdle similar to those of early
bony fishes, but lost posttemporal & acquired interclavicle (which
still occurs in several amniotes, e.g., alligator, birds, & monotremes)
Clavicle & coracoid - one or both typically brace scapula against
sternum (as in birds; below)
Scapula - present in all tetrapods with even vestiges of anterior limbs,
e.g., turtles & birds & mammals
brace posterior paired appendages
no dermal components (unlike pectoral girdle)
Fishes - pelvic girdle consists of 2 cartilaginous or bony plates (ischiopubic plates) that articulate with the pelvic fins
 Frogs & toads
ilia elongated & extend from sacral
vertebra to urostyle
joint between ilium & sacral
vertebra (sacroiliac) is freely
moveable (& moves when a frog
or toad jumps)
 Birds:
ilium & ischium expanded to
accommodate musculature
needed for bipedal locomotion
girdle is braced against lumbar &
sacral vertebrae
pubic bones are typically reduced
(long but thin); the limited pubic
symphysis provides a larger outlet
for eggs

 Mammals - ilium, ischium, and

pubis unite to form the
innominate bone (the 2
innominates = pelvic girdle)
 Limbs
Starting with amphibians, vertebrates typically have 4 limbs. However, some have lost one
or both pairs &, in others, one pair is modified as arms, wings, or paddles
typically have 5 segments:
Anterior limb
brachium (upper arm) - consists of humerus
antebrachium (forearm) - consists of radius & ulna
carpus (wrist) - consists of carpals
metacarpus (palm) - consists of metacarpals
digits - consist of phalanges
Posterior limb
femur (thigh) - consists of femur
crus (shank) - consists of tibia & fibula
tarsus (ankle) - consists of tarsals
metatarsus (instep) - consists of metatarsals
digits - consist of phalanges

 Arm & forearm

Upper arm = humerus
Forearm = radius & ulna
Manus (or hand)
Wrist - 3 rows of carpal bones:
proximal row = radiale, ulnare, intermedium, &
pisiform
middle row = 3 central carpals (centralia)
distal row = 5 distal carplas numbered 1 through 5
(starting on thumb, or radial, side)
Palm - metacarpals
Digits
each consists of a series of phalanges
general formula' (starting at thumb) = 2,3,4,5,3

Some vertebrates lack both pairs of limbs

caecilians (apodans)
most snakes
snake-like lizards
Some vertebrates have forelimbs only:
manatees & dugongs
dolphins (see diagram below)
cetaceans (vestigial elements may be embedded in body wall)
sirens (salamander)
Adaptive modifications of the
manus
1 - Flight
Birds - loss of digits & bones plus
fusion of some bones
Bats - 5 digits; elongated
metacarpals (II through V) &
phalanges support the patagium
Pterosaurs - 4th digit elongated
to support patagium
2- Swimming - increase in number,
& size, of phalanges
 3 - Terrestrial locomotion (walking
& running):
Plantigrade
flat-footed
all bones of manus and/or pes on the
ground
amphibians, most reptiles (see
alligator photo above), & some
mammals (insectivores, monkeys,
apes, humans, & bears)
Digitigrade
1st digit is reduced or lost
manus & pes are elevated
rabbits, rodents, & many carnivores
Unguligrade
reduced number of digits
walk on tips of remaining digits
claws become hooves

 4 - Grasping
opposable thumb
saddle joint at base of thumb where it meets palm
thumb at wider angle from index finger
strong thumb muscles
Posterior limbs - bones are comparable to those of forelimbs except
that a patella (kneecap') develops in birds & mammals