Chapter 9: Skull and Visceral Skeleton

Skull
-Formed by three main parts :
1. Neurocranium
2. Dermatocranium
3. Splanchnocranium

Neurocranium
Primary Braincase
Endocranium or Chondrocranium
Protects the brain and certain special sense
organs
Partly or wholly replaced by bone (except
cartilaginous fishes)
Gnathostomes Cartilaginous Brain case - Start
as several independent cartilages that later
expand and unite.

- the expanding basal plate unites with

the otic capsule, lateral to the hindbrain
1. Hypophyseal Fenestra 2 plates (midline)
- accommodates hypophysis and
internal carotid arteries.
-later reduced to a pair of foramina
-transmitting the arteries
2. Tectum (cartilaginous roof) above the brain
-has 1 or 2 prominent fenestrae
- primitive condition in some craniates
3. Foramen Magnum largest structure
- rear wall of neurocranium
*Sclerotome ( Epimeric Mesoderm) - Mesenchyme that
forms parachordal cartilages, occiput and excapsulate
the inner ear
*Neural crest Mesenchyme that forms the rest of the
neurocranium

Cartilaginous Stage

Neurocranium Prechordal and Parachordal

Cartilages
Parachordal Parallel the anterior end o notochord
- beneath midbrain and hindbrain
- incorporated with single, broad,
cartilaginous basal plate

Prechordal Anterior to notochord

- underneath the forebrain
- expand and unite across midline of
anterior ends
- Ethmoid plate
Sense Capsules (Cartilage)
Olfactory (Nasal) capsule
Partially surrounding the olfactory
epithelium
Incomplete (arteriorly) accesss for
water or air
Otic capsule
Completely surrounding the otocyst
-developing inner ear
Optic capsule
Forms around the retina
Sclerotic coat of the eyeball
Usually fibrous (mammals)
Cartilaginous/bony plates often form
within the sclerotic coat
Not fused with neurocranium
Allows free/independent
movement of the eyeball

Completion of floor, walls and roof

Ethmoid Plate expands and unites anteriorly

with olfactory capsules

Cartilagionous Neurocrania of Adult Craniates

4. Agnathans
o Olfactory capsule
o Otic capsules
o Basal plate
o Notochord (not fused with basal plate)
*Roof above brain is unchondrified and fibrous
5. Cartilaginous Fishes
o Chondrocranium Adult cartilaginous
braincase formed by embryonic
components
o Fully developed walls and presence of
posterior occipital wall
o Unossified fenestra
o Otic capsules fused into post lateral
walls of braincase
o Olfactory capsules firmly united
anteriorly to otic capsules
o Hypophysis cradled in a cartilaginous
pocket
o Sella Turcica Cartilaginous pocket
beneath the brain
o Rostrum (Neurocranium) projects
forward beyond olfactory capsules
o Occipetal Condyle immovable
articulation between occipital region of
neurocranium
o Endolymphatic fossa Depression: two
pairs of foramina
o Endolymphatic and Perilymphatic ducts
housed by endolymphyatic fossa
o Endolymphatic duct opens on to the
surface of the head
*TOTALLY CARTILAGINOUS ADULT SKULL only in
chondrichthyes

Chapter 9: Skull and Visceral Skeleton

* Most basal tetrapods have no ethmoidal ossification
centers
Neurocranial Ossification Centers
- Occurs more or less simultaneously at numerous
separate ossification centers.
-Four Universal group
1. Occipital
2. Sphenoid
3. Ethmoid
4. Otic

Occipital Centers
- Cartilage around the foramen magnum
- may be replaced by as many as 4 bones.
Basioccipital bone produced by endochondral
ossification centers ventral to foramen magnum
- Underlying the hindbrain
Exoccipital bones (2) produced by centers in
the lateral walls of foramen magnum
Supraoccipital bone above foramen
-In mammals The 4 bones fuse to form one
occipital bone.
-In other Tetrapods, neurocranium articulates with 1st
vertebra via two occipital condyles.
-In Stem amphibians, a single condyle borne chiefly
on basioccipital bone.
Sphenoid Centers
Basisphenoids ossification of
embryonic cartilaginous neurocranium
underlying the midbrain and pituitary
gland
-In mammals:
Sphenoid elements may separate or unite to
form a single sphenoid bone with wings
Presphenoid bone- ossifies anterior to the
basiosphenoid bone
Bony platform basioccipital + sphenoid
bones that underlies the brain
Side wall in the sphenoid region above
basiosphenoid
o Formed by additional presphenoid
ossification
Alisphenoid forms lateral ossification of the
sphenoid region
-In Archosaurs (Crocodiles and Dinosaurs):
Laterosphenoid bone forms lateral
ossification of the sphenoid region
Orbitosphenoid bone separate interorbital
septum
Ethmoid Centers
-lies immediately anterior to the sphenoid
-ethmoid plate and olfactory (nasal) capsules
-In tetrapods, ethmoid tends to remain cartilaginous

Mesethmoid bones chief ossification

center in amniotes
- septum of birds and mammals
- Birds anterior portion of interorbital
septum
- Reptiles and Birds one or more scroll like
turbinal bones (Conchae) in the walls of
nasal passages
- Mammals Cribiform plate perforated by
Olfactory foramina transmit bundles of
olfactory nerve fibers from the olfactory
epithelium to the brain.
-Anurans Sphenethmoid sole bone
arising in the ethmoidand sphenoid regions
- Ectethmoid develops in the lateral
walls of the nasal passageways of
Sphenodon
*Not all cartilages in the nasal passageways of the
amniotes are derived from the ethmoid ossification
centers.
Otic Centers
-cartilaginous otic capsule surrounding the
membranous labyrinth is replaced by :
o Prootic bones
o Opisthotic bones
o Epiotic bones
-In Birds and Mammals
- prootic, opisthotic, and epiotics unite to form
one periotic or petrosal bone may unite with
squamosal (membrane bone) temporal bone
-In non-avian Reptiles
- opisthotics fuse with exoccipitals
*6 ossification centers have been describes in the
human fetus otic capsule.

Dermatocranium
-Membrane bones
-How it began? (page 167)

Basic Structure
o Bones that form above and alongside the
brain and neurocranium(roofing bones)
o Dermal bones of the upper jaw (marginal
bones)
o Dermal bones of the primary palate
o Opercular bones
Hypotheses:
1. Ray-finned and lobe-finned fishes share via their
common ancestor parietal and postparietal
bones with the frontal being unique to the later
sarcopterygians.
2. Roofing bones evolved independently and are
not homologous

Roofing bones

Chapter 9: Skull and Visceral Skeleton

-early pattern seen in rhipsidian fishes inherited
almost intact by labyrinthodonts
-protective shield over brain and special sense
organs
-openings are for the nares, paired eyes and a
median or parietal eye.
-In Rhipidistians, a series of pared and unpaired
scalelike bones extended along the middorsal
line from the nares to the occiput, overlying the
brain, olfactory capsules and any other
components of the neurocranium.
- In Labyrinthodonts, the unpaired bones were
missing. A series of paired nasals, frontals,
parietals and postparietals(dermoccipetals) took
their place.
-In modern amphibians, the postparietals
disappeared as independent bones, may have
coalesced with endochondral supraoccipitals.
-In fishes, amphibians and lizards, a parietal
foramen housing the median eye is still present.
o

Generalized skull:
Lacrimal related to the
nasolacrimal duct in amniotes
which drains excess fluid
tears.
Prefrontal
Postfrontal
Jugal (infraorbital)
Posterior angle of the skull
Intertemporal
Supratemporal
Tabular
Lower angle of the skull
Squamosal
quadratojugal

Dermal bones of the upper jaw

o Palatoquadrate cartilages the only
upper jaws that cartilaginous fishes
develop embryonic precursors of the
upper jaw of bony vertebrates.
o Premaxilla and maxilla tooth-bearing
dermal bones that overlay or sheath the
cartilages : margins of the
dermatocranuim.
Primary Palatal bones
- the roof of the oropharyngeal cavity of fishes
and the oral cavity of basal tetrapods.
o In sharks, cartilaginous- floor of
neurocranium where the brain rests.
o In bony Vertebrates, membrane bones
are applied to the underside of the
neurocranium and any part of the
palatoquadrate cartilages (upper jaw
cartilages)
o Rhipidistian fishes and early tetrapods,
palate membrane bones were:

unpaired parasphenoids
beneath the ethmoid region.
Paired palatines, ectopterygians
and pterygoids laterally
Invested the
palatoquadrate cartilage
Internal nares pierced the palate
anteriolaterally
Primary palate is still present in
tetrapods (with modifications)
Those with secondary palate ,
the primary palate remains in
the roof of the nasal
passageway

Opercular bones
-flap of tissue that arises as an outgrowth of the
hyoid arch and extends caudad over the fill slits
o Membranous in holocephalans
o Absent in elasmobranchs
o Stiffened by squamous plates of
dermal bone in bony fishes.
Most constant: Large opercular,
smaller operculars overlying the
site of articulation of the upper
and lower jaws, superoculars
and interoperculars.
Basal bony fishes: one or mor
gular bones lie in the opercular
membrane in the floor of the
opercular chamber
- in Ray-finned fishes,
Branchiostegal rays located in
caudally directed flapsBranchiostegal membranes
No operculum/gulars/vestiges in
tetrapods

Neurocranial-Dermatocranial Complex of Bony Fishes

Basal Actinopterygians
*Sturgeons:
-Neurocranium cartilaginous for life
-Endochondral Ossification only in the otic
capsuls and sphenoid bone walls of orbit
*Spoonbills
-unusual bill is extension of the cartilaginous
rostrum
*Polypterus
- retains more primitive features of paleozoic
fished than sturgeons and spoonbills.
-well ossified neurocranium overlaid In the
middorsal line by paired nasals, parietals and
postparietals.
-premaxilla and maxillae overlie the
endochondral palatoquadrate bones of the first
visceral arch
-preopercular bone overlies the site of
articulation of the upper and lower jaws

Chapter 9: Skull and Visceral Skeleton

-highly cartilaginous skull (Amia)

-Bony skull (gars)

-mesethmoid, epiotic and pterotic often

incorporate membrane bone along with the
usual replacement bone.
-branchiostegal rays in paired branchiostegal
membranes beneath the gill chamber take the
place of the more primitive gular bones of early
ray-finned fishes

Basal Neopterygians
-Dermal bones are grooved and pitted,
sculpturing effects of overlying skin
-skull bones or dermal scales
-cheek plates evidence that dermatocranium
may have been derived from ealier dermal
scales.

Teleosts
-highly specialized skull and architecturally
diverse
- combination of highly movable jaws and
palates.
-skull is compressed laterally and vaulted
dorsally
-fully ossified neurocranium except olfactory
capsules

Dipnoans
-scale-like dermal bones to few broad bony
plates
-largely cartilaginous neurocranium
-palate accommodates openings of the nasal
canal into the oral cavity just behind the mouth

Neurocranial-Dermatocranial Complex of Modern

Tetrapods