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NEUROCRANIUM (Endo/chondrocranium, primary braincase)

1. Protects brain and special sense organs
2. Starts as cartilage
3. Replaced by bone (whole or part only) EXCEPT for cartilaginous fish
4. Starts as several cartilage that unites as ONE.
5. PARACHORDAL STAGE develop underneath the brain. Cartilage
expand across the midline and unite
6. BASAL PLATE united notochord and PARACHORDAL STAGE
7. PRECHORDAL STAGE (trabeculae cranii) develop underneath the brain.
develop at the anterior of notochord and underneath the forebrain.
8. ETHMOID PLATE expanded Prechordal cartilages across midline @
anterioir ends
SENSE CAPSULES
1. Other cartilage formed in 2 locations namely;
a. Olfactory capsule perforated by foramina (transmit nerves and
vascular channels), partially surrounding olfactory epithelium coz
of access of air / water
b. Otic developing inner ear, surrounds the otocyst, perforated by
foramina (transmit nerves and vascular channels)
c. Optic around retina, later becomes eyeball sclerotic coat,, fibrous
in mammals, may have cartilaginous/bony plates (ancient trait),
DOES NOT FUSE WITH THE SKULL
WALLS, FLOOR, ROOF
1. Ethmoid plate + olfactory capsules (anteriorly)
2. Basal Plate + otic capsules (lateral with the hindbrain direction)
3. Ethmoid + Basal = floor where brain rests
4. Hypophyseal fenestra the midline where the ethmoid and basal cartilage
plates meet,
a. accommodates hypophysis and internal carotid arteries (en route to
brain).
b. Reduced to foramina pair (transmitting arteries)
5. Cartilaginous walls in the brain is also formed
6. Tectum a cartilaginous roof w/ prominent fenestrae, also formed.
7. Presence of cranial nerves, blood vessels found
8. Deposited cartilage with formed foramina
9. Foramen magnum largest foramina @ rear wall of neurocranium
10. Remodeling of neurocranial cartilage takes place as brain develops
11. Mesenchyne

gives rise to neurocranium from neural crest ventrad to optic stalks >
developing eyeballs connection to brain
gives rise to parachordal cartilage, occiput <?> , from schlerotome
the rest of mesenchyme for neurocranial formation is sourced from neural
crest

CARTILAGINOUS NEUROCRANIA OF ADULT CRANIATES

1. Living Agnathans
a. Neurocranium components independent throughout life
b. Olfactory capsule, otic capsule, basal plate, notochord (not fused
with basal plate) present
c. Roof of brain is unchondrified and fibrous
2. Cartilaginous fish
a. Model : Squalus acanthias
b. High water mark in development, Forms a boxlike adult
cartilaginous braincase CHONDROCRANIUM
c. CHONDROCRANIUM no more visible components, fully
developed wall,
posterior occipital wall for the first time
(gnasthosomes)
d. brain roof is fully cartilage, last part to chondrify is the
ROSTRUM (w.c may still be soft in younger specimens)
e. otic capsules completely fused with posterolateral walls of
braincase + olfactory capsules united anteriorly
f. visible notochord (seen as cephalad extending ridge) from the base
of foramen magnum (which has an occipital condyle at each side)
g. Occipital condyle immovable articulation bet. Occipital region of
neurcranium and 1st vertebrae
h. hypophysis cradled in cartilaginous pocket
i. sella turcica-beneath the brain
j. projects forward as ROSTRUM beyond olfactory capsules
k. Endolymphatic fossa depression at the posterodorsal aspect of
the neurocraniumm 2 pairs of foramina, houses the endolymphatic
(opens at surface) and perilymphatic ducts
3. BONY FISHES
a. Cartilaginous neurocranium throughout its life(chondrosteans,
gars, dipnoans)
b. Teleosts and tetrapods cartilage replaced by endochondrial bone
as it develops
NEUROCRANIAL OSSIFICATION CENTERS
1. occurs more/less simultaneously at numerous ossification centers

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2. actual no. varies per species
a. Occipital Centers
Basioccipital bone 1 or more centers @ ventral to
foramen magnum, underlying the hindbrain
2 Exoccipital lateral walls of foramen magnum
Supraoccipital above foramen,
1 Occipital bone fusion of 4 occipital elements
(mammals), 1 or 2 remain cartilaginous in amphibians
2 Occipital condyles articulates with the 1st vertebra,
single in amphibians (basiooccipital), birds and reptiles
b. Sphenoid center
o Basisphenoid bone ossifies under midbrain and pituitary gland,
anterior to basioccpita
o Presphenoid Mammals. Anterior to basissphenoid, with addl
ossification at the walls
o Laterosphenoid bone (archosaurs) lateral ossification
o Orbitosphenoid (archosaurs) separate interorbital septum
o Alisphenoid in some mammals, but from palatoquadrate cartilage
not neurocranium
o Sphenoid elements may form (single sphenoid) or separate
o Sella turcica pituitary gland rests
o No replacement bones develop above the brain
c. Ethmoid centers
Anterior to sphenoid
Ethmoid plate + nasal capsules
Remains cartilaginous in tetrapods
Ossification centers in amniotes:
Mesethmoid nasal septum of birds and mammals, turbinal
bones @ nasal passageways of reptiles and birds, cribiform
plate in mammals
Sphenethmoid anurans, sole bone arising from both
ethmoid and sphenoid bones
Ectethmoid nasal passageways of Sphenodon

d.

Not derived from ethmoid ossification - winglike alar and

small sesamoid cartilage (human nose)
Otic Centers

Replaced by the bones: prootic, opisthotic, epiotic

May unite with other bones

Periotic/perosal bone all 3 unite in birds and mammals,

Petrosal + Squamosal =Temporal bone

GENERALIZED DERMATOCRAUNIUM
1. Membrane bones of the skull
2. Fish have highly specialized structures
3. How It May Have Begun
a.
Earliest vertebrates (Ostracoderms were incased in dermal
armor)
b.
This armor goes through one or more cycles of expansion
and reduction
c.
Bony plates are found in the head- integral part of the
skull
d.
Neurocranium is the endoskeleton
e.
Dermal bone exoskeleton dermatocranium (living
fossils)
f.
Modern vertebrates no longer ossify at the
skin(mesenchyme to dermis) rather from subdermal mesenchyme.
Though the inductors may be the same.

4.

Basic Structurea. Read A Question in Homology (regards naming)

b. Roofing Bones

Early pattern in Rhipidistians inherited by

Labyrinthodonts.- protective shield to brain and special
sense organs, no unpaired bones, paired nasals, frontal
parietals, postparietals (disapperared in modern
amphibians)

paired and unpaired scale bones extends

Parietal foramen housing median eye still found in fish

amphi, lizards

Lacrimal from nasolacrimal duct (drain of excess fluid

and tears

Lacrimal, pre and postfront, jugal ring around the orbit

Posterior: Intertemporal, supratemporal, tabular,

squamosal, quadratojugal,

Labyrinthodonts longer snouts that rhipidistians (food)

c. Dermal bones of the Upper Jaw

Embroynic origin: Palatoquadrate cartilage

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Palatoquadrate cartilage > Overlaid with > Premaxillae and
maxilla (tooth bearing bones)

Primary Palatal Bones:

1. Roof of oropharyngeal cavity/ oral cavity of
tetrapods/ cartilaginous floor of neurocranium in
sharks
2. Bony vertebrates: membrane bones under
neurocranium and upper jaw cartilage
3. Rhipis. Fish and early tetrapods: unpaired
parasphenoid (beneath sphenoid), paired vomers
(ethmoid region), & paired palatines,
ectopterygoids and pterygoids
4. Internal nares pierced anterolaterally
5. May bear teeth in all extant basal vertebrates
6. Modified primary palate may still be present in
all tetrapods
Opercular Bones
1. Flap of tissue arises as an outgrowth of the hyoid
arch and extends caudad over the gill slits.
2. Membranous in holocephalans, absent in
elasmobranchs,
3. Stiffened plates of dermal bone in bony fishes:
Opercular, preopercular, subopercular,
interopercular
NEUROCRANIAL-DERMATOCRANIAL COMPLEX OF BONY
FISHES
Sturgeons and Spoonbills neurocranium cartilaginous all thru out
life, isolated sites of ossification
Spoonbill an extension of cartilaginous rostrum
Polypterus neurocranium is well ossified overlaid with paired
nasals, parietals, postparietals
Amia and Gars- little resemblance to modern fish skulls
Gars- cheek plates, bony NC
Amia highly cartilaginous neurocranium
Teleosts highly specialized, architecturally diverse (feeding
habits), highly maneuverable jaws and palates, many bones
(anatomic diversity), neurocranium is fully ossified except for
olfactory capsules, carp skulls have islands of cartilage on the
surface of the carp skull, dermal bone complement of the palate

reflects that of the ancestral bony fishes. , bones of the skill were
named accdg to location, shape, other characteristic THESE
BONES ARE NOT HOMOLOGUES.
Dipnoans appeared @Devonian, no burst of speciation
conservative skull change, dermatocranium of lungfish has
evolved from a large number of scalelike dermal bones to few
broad bony plates with cartilaginous neurocranium & palate that
accommodates openings of the nasal canal into oral cavity behind
the mouth.
NEUROCRANIAL-DERMATOCRANIAL COMPLEX OF
MODERN TETRAPODS
Amphibians skull is moderately ossified (platybasic),
incomplete neurocranium dorsally,
o rigidity for burrowing, replacement bones in anurans:
sphenethmoid, 2 prootics, 2 exoccipitals all with
condyles,
o Columella skeletal rod, middle ear ossicle that conducts
sound waves from eardrum to capsule
o Complex dermatocranium missing primitive temporal
bones from orbit exposing otic capsule, present:
squamosal, quadratojugal, premaxillae and maxillae
o Primary palate has been altered large palatal vacuities,
reducing palatines to transverse splinters, results to
protruding eyeballs
o Urodeles parasphenoid is broad, palatal vacuities
lacking orders
Nonavian Reptiles - little changed from labyrinthodonts
o well ossified neurocranium, single occipital condyle, large
complement of membrane bones, parietal foramen for
median eye
o stem reptile skulls: temporal fossae, partial/complete
secondary palate, (from birds)
o Adult alligator ossified neurocranium, 2 exocipital , 1
supraoccipital, basiooccipital with condyle for atlas,
laterosphenoids, ethmoids, otics that are overlaid with
squamosal, a dermatocranial bone,
o Crocodile largest number of membrane bones
o Turtles- enigmatic skull
Temporal Fossae

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o

Cavernous opening in the temporal region of som amniote

skulls bounded ny 1 or more bony arches
o Anapsid no arches temporal bone (stem reptilesturtles)
o Synapsida 1 lateral Temporal fossa surrounded by
postorbital squamosal and orbital bones, last 2
zygomatic arch (cheek bones in humans)
o Diapsid archosaurs, Spehenodon and snakes. 2 fossae
are present Upper is supratemporal, lower is zygomatic
arch.
o Modified diapsid skulls modern lizard , - lost part or all
of the arches leaving a cavernous void Allows cranial
kinesis
o Euryapsid extinct reptiles convergence in lineages 1
temporal fossa, that resembles dorsal fossa of diapsids
o Turtle skull enigma temporal region it is absent. Wide
vacuity, recession of bones
o Provides space and surfaces in functionally advantageous
positions for accommodating the powerful adductor
muscles >lower jaws
o Early reptiles confined adductor mandibulae > allows
only bite off of food
o Temporal fossa provided the space for adductor muscle to
shorten and thicken during contraction.
o Allows more leverage and anchorage ( maseter,
zygomatic arch) - chewing
Secondary Palates
o Horizontal partition that partially or completely divides
the primitive oral cavity displacing the internal nares
o Crocodilians Medially directed palatal processes meet at
the midline forming a long bony secondary palate
o Varying degrees of completion in diff. reptiles
o If secondary palate is incomplete > deep longitudinal
groove (Palatal fissure) bordered with Palatal Folds (also
in birds)
o Mammals extends all the way to the pharynx but caudal
portion is BONELESS (tugs tugs)
CRANIAL KINESIS / KINETISM

o
o
o
o
o
o
o

Birds
o

Movement of a functional component of a skull

independent of another componentby intracranial joints
between 2 components
May have evolved independently during vertebrate
evolution
Teleosts, Lizards, Snakes, birds have it, Early reptilians
too (mammalian ancestor)
Raise upper jaw & palate as a unit independent of the
neurocranium when they open their mouths.
Teleosts 2 sides move independently
for food-getting and food manipulation in the oral cavity
enables suction, scraping, unhinging of jaws
The palate and temporal region have become so modified
for feeding that the dermatocranial-splanchnocranial
complex hasbecome greatly disrupted

Skull reptilian but modified for flight, feeding habits

and larger brain
o Lost roofing bones
o Thin dermal bones
o Carinate skulls obliterated sutures
o 2 solid regions:
Solid bony box (neuro and dermatoC) rear,
houses the structures needed for input and
information processing
Front food procuring and handling
o Posterior component highly vaulted, bulging outward and
arching upward alongside the great enlarged brain
o Orbits maximal visibility
o Parietal foramen is closed
o Incomplete neurocranium but completely ossified, with
single occipital condyle
o mSkull modified diapsid
o Kinetic plate resembles that of squamates except for
ectopterygoids
o Parasphenoid is immobile, fused to the basisphenoid
o Not all bird skulls are equally kinetic
o Woodpeckers have minimal kinetism due to its use
Mammals
o Dentary as sole jaw bone

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o
o
o
o
o
o
o
o

o
o
o
o
o

o
o
o
o

o
o
o
o
o

Altered site of articulation of the lower jaw in the

braincase and the secondary palate
3 bones in the middle ear cavity
Synapsid
Increasingly domed cerebral hemisphere
Neurcranium is incomplete dorsally
Fontanels-soft spot in the neurocranium until ossification
Bregmatic bones small bones in the fontanel that may
later ossify
Single bregmatic bone may develop antiepileptic
bone accdg to Paracelsus served as valve to relieve
pressure in the cranial cavity
Floor of brain basiooccipital, basissphenoid and
presphenoid
Partial side walls exocippitals, alisphenoids, lateral
extensions of the presphenoids
Supraoccipital completes a bony ring that resembles the
verterbra
Ethmoid cartilage & bone olfactory epithelium
Otic capsules unite solidly to form petrosal bone sbeneath
the overgrown temporal lobes of themammalian cerebral
hemispheres
The sequence in mammals centrum like basio occipital
and basiospehnoid and presphenoid and their dorsal wings
resemble a series of 3 successive vertebrae that are
incomplete dorsally
Dermatocranium paired premaxillae, maxillae, jugals,
nasals, lacrimal and squamosals
Postparietal present in Homo erectus, Inca bone.
Premaxillae are not identifiable in adult human skulls
The zygomatic arch varies from massie to slender
depending on the magnitude of force exerted on it by the
masseter muscle
Temporal complex- consists of numerous components of
intramembranous and endochondral
Tympanic bulla tympanic (surrounds tympanic rings) &
entotympanic (cartilage replacement ossification)
Petrous portion ossified otic capsule
Mastoid portion endochondral origin new in mammals.
Petrotympanic bone tympanic + petrous portion united

o
o
o
o
o
o
o
o
o
o

Petrotympanic + squamosal = temporal bone

Hyoid arch + temporal bone = styloid process ( some
species only)
Air-filled cranial sinuses- found within the maxilla,
frontal, sphenoid, and ethmod bones
Front sinus of goats and sheep horns
Unpaired vomer base of the nasal septum (consist of
mesethmoid bone + cartilage)
Nasal process of the palatine lateral wall of the
nasopharynx
Pterygoids small winglike processes (origin of the same
name muscle)
Parasphenoid and ectopterygoids lost
2ndary palate palatal process of the premaxilla, maxilla
and palatine bones
3 pairs of turbinal bones (nasal conchae) in the medial
walls of the nasal passageways
2 lower conchae nasal epithelium
Superior concha- olfactory epithelium
Reptiles have 1 turbinal, birds have 2
Unique in mammals presence of ossified
posterior tips of the palatoquadrate and Meckels
cartilage in the middle ear cavity
No. of bones especially membrane bones tend to
be reduced due to evolution of tetrapods
Reduction is a fusion of adjacent embryonic
ossification centers, phylogenetic loss of
ossification centers, & obliteration of sutures in
young animals.
Membrane bones often unite with adjacent
replacement bones, giving rise to a single bone
with a dual history.

VISCERAL SKELETON
Splanchnocranium
Skeleton that develops within the pharyngeal arches
Fish skeleton of jaws and gills
Tetrapods- modified for land based functions
Come from neural crest, may be partly/wholly replaced by bone
First arch ensheathed by dermal bone. Only that arch

Sharks
o
o
o
o
o
o
o

o
Squalus acanthias generalized vertebrae, lacks bone
cartilage in each pharyngeal arch
median basihyal and basibranchial cartilage pharyngeal floor.
Skeleton of mandibular arch Palatoquadrate cartilage meet
dorsal, Meckels cartilage meet ventrally
Slender labial cartilage where lips are, extended from the
mouth
Paired Hyomandibular cartilage skeleton of the hyoid arch
dorsally and gill bearing ceratohyals
Hyostylic jaw hyomandibula is bound by ligaments to the otic
capsule and suspend the jaws and entire branchial skeleton from
the neurocranium

Jaw Suspension in Fish

o braced against some support
o Elasmobranchs and bony fish braced against otic capsule
o Posterior of Palatoquadrate omandibula (Hyostylic jaw
suspension/hyostyly)
o Amphistyly older sharks in which hyomandibular and
palatoquadrate is braced independently against the braincase
o Autostyly even older sharks, also (independently found) in
chimaeras and lungfishes. Palatoquadrate is ATTACHED to the
neurocranium and the hyomandibula plays no role in jaw
suspension
o Anatomic relationship of the palatoquadrate cartilage to the
hyomandibula and to the neurocranial-dermatocranium complex
related to the diet.
Bony Fish
o Resembles that of sharks in basic morphology
o Embryonic palatoquadrate and Meckels cartilage become invested
by membrane bones during development
o Hyoid skeleton more segments
o Embryonic cartilage of gill arches replaced by bone
o The posterior end of the palatoquadrate cartilage, articulate
thorughtout life with the posterior ends of the Meckels cartilage
o Refer to Figure 9.10
o Meckels cartilage ossify at their caudal ends to become articular
bones

o
o

Symplectic & Interhyal ossification centers in the

hyomandibular cartilage
Epihyal ossification center in the ceratohyal
Gill arch same as a shark. Has: hypo-, cerato-, epi- and
pharyngobranchial . middle 2 bear gills, borders covered with
denticles. Size and arrangement depends on diet.

Feeding and Arrangement in Bony Fish

o Early Jawed- wide mouths, upper jaw fused with braincase
o Modern teleosts large adductor muscle, additional array of
muscles, addl number of jaw muscles, mouth narrower and oval, >
jaws can be thrust forward for feeding
o Feeding by inertial suction creates forward suction (Petrotilapia
and predatory fish)
o Theory has been advanced that vertebrate jaws are former gill
arches that become modified, along with hyoid arch for predatory
feeding
Living Agnathans
o Like that of jawed fish
o Has V-shaped lingual cartilage horny teeth & rasping teeth,
operated by protractor & retractor muscles
o No evidence that cartilage related feeding apparatus is related to
ancestral visceral arches
o Rest of pharyngeal skeleton branchial basket
Tetrapods
o Model : Larval frogs, used for branchial respiration and anchorage
o 6 pairs of visceral cartilage
Hypobranchial arch gill arch cartilage meet ventrally
3rd, 4th, 5th regress
1st enlarges along with hypobranchial plate
2nd arch ceratohyal slender anterior horn/cornu
4th arch posterior horn
Fate of the Palatoquadrate and Meckels cartilage
o the quadrate remains as the site of articulation of the cranium with
the lower jaw in all nonmammalian tetrapods
o squamates/ birds kinetic mechanism
o quadrate bone mammals, incus, middle ear ossicle
o turtles and crocodilians continue to grow and prominent within
the adult mandible
o cartilage may become ensheathed by dermal bones

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Modern amniotes and mammals dentary only
Bony fish posterior end of meckels cartilage becomes ossified
(articular bone)
o Mammals becomes malleus (middle ear ossicle)
Expansion of the dentary and a new jaw joint in Mammals
o Expansion of the dentary brought in close to the squamosal (new
articulation site at the skull)
o Former 2 sites of articulation
Primitive against quadrate
New one- condyloid process of dentary against squamosal
Both are present in Eozostrodon
Eventually, the new one remained
Condyloid process of the mandible differed by demand of
feeding habits of mammals
Ear Ossicles from the Hyomandibula and Jaws
o Hyomandibular cartilage are interspersed bet. Quadrate cartilage
and otic capsule (inner ear)
o Autostylic jaw hyomandibular bone can be dispensed (dipnoans),
persisted in mammals > DISSOCIATED WITH THE
QUADRATE AND BECAME TYMPANIC MEMBRANE
o Otic capsule surrounded by portion of 1st pharyngeal pouch
(middle ear cavity)
o Hyomandibula FIRST MIDDLE EAR OSSICLE /
COLUMELLA/STAPES of tetrapods
o Therapsids - articulated with squamosal > freed articular and
quadrate bones
o Caudal end of Meckels cartilage malleus of mammals
o Incus of middle ear quadrate bone of synapsids
o C.Reichert 19th century, mammalian ear ossicles > jaws
o E. Gaupp 20th century, modified theory to include stapes from
hyoid archs dorsal tip
Amniote Hyoid
o consist of a body in the pharyngeal floor just anterior to the larynx
and of 2 or 3 horns/cornua in the pharyngeal walls
o Anurans from basibranchail and hypobranchial cartilages,
skeleton of the hyoid arches, and 1 of the larval gill-bearing arches
o Amniotes from homologous anlagen
o 2nd arch cartilage anterior horns
o 3rd and 4th additional horns. (4th not all the time)
o
o

o
o
o
o

o
o
o
o

Entoglossus lizards and bones , elongated process (body forward

to the darting tongue)
Male lizard gular pouch
Snakes no hyoid, branchial skeleton is vestigial
Woodpeckers entoglossus + caudal horns + tongue = impaling
grubs
Base of entoglossus 2 long flexible caudal horns loop
all the way to the LORE space between eyes and bill
continue forward to the nasal canal, elastic recoil of horns
returns tongue to mouth after impaling food
Mammals cranial arch at second arch, caudal horns @ 3rd
Greater horns dogs and cats (longer cranial) composed
of 4 segments
Tympanohyal (dorsalmost)
Rabbits anterior horns are shorter (lesser horns)
o Tympanohyal represented by styohyal
bone
Humans anterior horns = ceratohyals
o Lesser horns unossified stylohyoid (
epi- and styohyal segments)
o Styloid process tympanohyal +
temporal bone
Hyoid anchors the movable tongue of the tetrapods
Skeleton for the buccophayngeal pressure pump respiration anurans
Provides attachment for some extrinsic muscles of the larynx
The associated hypobranchial and branchiometric muscles of
amniotes approachthe hyoid in many directions. These muscles
stabilize the hyoid in a single position
Laryngeal Skeleton
Cricoid and artytenoid cartilage replacing bones
Mammals throid cartilage in addition from 4th and 5th
pharyngeal arch

Perspective
For feeding and branchial respiration transmission of
sounds