SKELETAL SYSTEM Bony fishes, lungfishes, and most ganoids

- gives the vertebrate body shape - retain highly cartilaginous neurocranium that is covered by membrane
- supports its weight bone
- offers a system of levers that together with muscles produces
movement Other bony vertebrates
- protects soft parts such as nerves, blood vessels, and other viscera - embryonic cartilagious neurocranium is largely replaced by
replacement bone (the process of endochondral ossification occurs
TWO TYPES OF SKELETAL SYSTEM almost simultaneously at several ossification centers)
1. Exoskeleton - primarily an embryonic structure serving as as scaffold for the
- formed from or within the integument developing brain and as a support for the sensory capsules
- the dermis giving rise to bone and the epidermis to keratin
2. Endoskeleton Chondocranium Embryology
- forms deep within the body from mesoderm and other sources, 1. Condensations of head mesenchyme form elongate cartilages next
not directly from the integument to the notochord
- tissues contributing to the endoskeleton include fibrous 2. Forms cartilage at 6 pairs of isolated regions
connective tissue, bone, and cartilage a) Parachordal (posterior pair)
- includes cranial skeleton/skull and postcranial skeleton b) Polar (middle pair, only in some vertebrates)
- examples of postcranial skeleton are vertebral column, limbs, c) Trabeculae (anterior pair)
girdles, and associated structures such as ribs and shells d) The occipital
e) The otic, optic, and nasal capsules
CRANIAL SKELETON/ CRANIUM 3. As development proceeds, these cartilages fuse
- a composite structure formed of three distinct parts 4. The region between the nasal capsules formed by the fusion of the
anterior tips of the trabeculae is the ethmoid plate
1. Splanchnocranium (yellow) 5. The parachordals grow together across the midline to form the
- which first arose to support pharyngeal slits in protochordates basal plate between the otic capsules
2. Chondocranium (blue) 6. The occipitals grow upward and around the nerve cord to form the
- underlies and supports the brain occipital arch
- is formed of endochondral bone or of cartilage, or both
3. Dermatocranium (pink)
- a contribution that in later vertebrates forms most of the outer
casing of the skull
- composed of dermal bones

Braincase
- collective term that refers to the fused cranial components
Collectively, all of these expanded and fused cartilages constitute
immediately surrounding and encasing the brain
the chondocranium
- dermatocranium, chondocranium, splanchnocranium
SPLANCHNOCRANIUM
Neurocranium
- used as an equivalent term for the chondocranium along with fused or - an ancient chordate structure
attached sensory capsules--the supportive nasal, optic, and otic - in vertebrates, it generally supports the gills and offers attachment
capsules for the respiratory muscles while some elements contribute to the
jaws and hyoid appratus
CHONDOCRANIUM or NEUROCRANIUM
Splanchnocranium Embryology
- the primitive cartilaginous skeletal structure of the fetal skull that
1. The splanchnocranium arises from neural crest cells
grows to envelop the rapidly growing embryonic brain
2. The neural crest cells migrate into the walls of the pharynx to form
- inclused the box that encloses the brain and the capsules surrounding
pharyngeal arches between the gills
the sense organs
3. Each arch can be composed of a series of up to five articulated per
- protects the brain
side:
- can vary greatly in different species, but in general, it is made up of
a) Pharyngobranchial
five components: the sphenoid, the mesethmoid, the occipitals, the
b) Epibranchial
optic capsules, and the nasal capsules
c) Ceratobranchial
d) Hypobranchial
Cartilaginous fishes (elasmobranchs)
e) Basibranchial elements
- does not ossify
- instead, it grows still farther upward and over the brain to complete
Pharyngeal arches of aquatic vertebrates usually are associated
the protective walls and roof of the braincase and it is retained
with their respiratory gill system, they are referred to as branchial
throughout life
arches or gill arches
- the expanded and enveloping chondocranium supports and protects
the brain within
 - dentary means tooth-bearing
- splenials, angular (posterior cornet of the mandible) and the
surangular (above)
- many of these bones wrap around the medial side of the mandible
and meet the prearticular and one or several coronoid to complete the
medial mandibular wall

Left and right mandible usuall meet anteriorly at the midline in a

mandibular symphysis

- lateral bones: dentary, splenials, angular, surangular

- medial bones: prearticular, coronoids

Jaws
- the branchial arches that support the mouth

Mandibular arch
- the first fully functional arch of the jaw
- the largest and most anterior of the modified series of arches
A. Palatoquadrate (dorsal)
- contributes to the upper jaw
B. Meckel’s cartilage or mandibular cartilage (ventral)
- contributes to the lower jaw
Hyoid arch
- the second arch
- most prominent component is the hyomandibula
- in mammals, the ventral portion is involved in support the tongue,
muscles used for swallowing and jaw movements

Branchial arches I-V

- associated with the gill apparatus
- in mammals, they eventually contribute to the larynx
OVERVIEW OF THE SKULL MORPHOLOGY
DERMATOCRANIUM
Braincase
- the dermal bone that is believed to be derived from the external armor - also known as neurocranium
of primitve fish - the upper and back part of the skull
- phylogenitically, these bones arise from the bony armor of the
- forms a protective case around the brain
integument of early fishes and sink inward to become applied to the - occipital bones form the end of the sphenoid platform
chondocranium and splanchnocranium - occipital bones are made up of:
- bony elements of the armor also become associated with the
A. Basiooccipital
endochondral elements of the pectoral girdle to give rise to the dermal B. Supraoccipital
components of this girdle C. Paired exoccipitals
- forms the sides and roofs of the skull to complete the protective bony
case around the brain In sharks
- forms most of the bony lining of the roof of the mouth, and encases - an elabortate cartilaginous case around the brain
much of the splanchnocranium
- teeth that arise within the mouth usually rest on dermal bones In fishes and tetrapods
- extensively ossified
Parts of Dermatocranium - a box with a platform of endoskeletal elements supporting the brain,
1. Facial series (pink) all encased in exoskeletal bones
- encircles the external naris and collectively forms the snout
- maxilla and premaxilla define the margins of the snout and usually Foramen magnum
bear teeth - close the posterior wall of the braincase
- nasal lies medial to the naris - large hole
2. Orbital series (light blue) - through which the spinal cord runs
- encircles the eye to define the orbit superficially
- lacrimal takes its name from the nasolacrimal (tear) duct Occipital condoyle
- prefrontal, postfrontal, and postorbital continue the ring of bones - a single or double surface produced primarily within the basiooccipital
above and behind the orbit - where articulation or where two bones come together of the skull
- jugal usually completes the lower rim of the orbit with the vertebral column
3. Temporal series (green)
- lies behind the orbit, completing the posterior wall of the braincase Jaws
- intertemporal, supratemporal, and tabular make up the medial part of - a pair of bones that form the framework of the mouth of vertebrate
the temporal series animals
- squamosal and quadratojugal complete the temporal series and form - usually containing teeth and including a movable lower jaw and an
the “cheek” upper jaw
4. Vault series (yellow) - function by moving in opposition to each other
- the vault or roofing bones run across the top of the skull and cover the - used for biting, chewing, and the handling of food
brain beneath
- frontal Upper jaw
- postparietal
- parietal occupying the center of the rood and defining the small In sharks
parietal foramen if it is present - the palatoquadtrate is fully functional in the jaws
5. Palatal series (orange)
- cover much of the rood of the mouth In bony fishes and tetrapods
- pterygoid is the largest and most medial - makes limited contributions to the skull through its two derivatives:
- vomer, palatine, and ectopterygoid may have teeth on any or all Epipterygoid - fuses to the neurocranium
palatal bones Quadrate - suspends the lower jaw, except in mammals
- parasphenoid
6. Mandibular series (violet)
- the dermal maxilla and premaxilla replace the palatoquadrate as the CRANIAL KINESIS
upper jaw - refers to the movement within the skull
- specifically movement between the upper jaw and the braincase
Lower jaw or mandible about joints between them

In sharks Kinetic skulls

- Meckel’s cartilage - having mobile joints between various parts of the skull
Ex: being able to unhinge the jaws
In most fishes and tetrapods - found in ancient fishes, bony fishes, especially teleosts, very early
- Meckel’s cartilage persists but is enclosed in exoskeletal bone of the tetrapods, most reptiles, birds, and early therapsid ancestors to
dermatocranium which also supports teeth mammals

In mammals Akinetic skulls

- the lower jaw consists of a single bone, the dermal dentary - no such movement between upper jaw and braincase
- the anterior tooth-bearing part of the dentary is its ramus - present in modern amphibians, turtles, crocodiles, and mammals
- jaw-closing muscles are inserted on the coronoud process, an upward (except rabbits)
extension of the dentary

Hyoid or hyoid apparatus

Kinetic skulls Advantages
- a ventral derivative of the splanchnocranium behind the jaws
- provides a way to change the size and configuration of the mouth
- bones which suspend the tongue and larynx
rapidly
- consists of a main boy, the corpus, and extensions, the cornua
(horns)
Ex: in fishes and other vertebrates that feed in water, rapid kinesis
- elements of the hyoid apparatus are derived from the ventral parts of
creates a sudden reduction of pressure in the buccal cavity so that the
the hyoid arch and from parts of the first few branchial arches
animal can suck in a surprise prey (suction feeding)
In larval amphibians
- also allows tooth bearing bones to move quickly into stratefic
- the branchial bars persist but form a reduced hyoid apparatus that
positions during rapid feeding
supports the floor of the mouth and functional gills
- in many venomous snakes, linked bones along the sides of the skull
In adult amphibians
can rotate forward
- the gills and the associated part of the hyoid apparatus are lost,
although elements persist within the floor of the mouth usually to
Ex: the venomous viper erects the maxillary bone bearing the fang and
support the tongue
swing it from a folded position along its upper lip to the front of the
mouth, where it can more easily deliver venom into prey
In fish
- supports the floor of the mouth
Akinetic skulls Advantages
- loss of kinesis in mammals leaves them with an akinetic skull, which
In mammals
allows infants to suckle easily
- the distal end of the hyoid horn fuses with the otic region of the
- juvenile and adult mammals can chew firmly with sets of specialized
braincase to form the styloid process
teeth that work accurately from a secure, akinetic skull

SKULL FUNCTION AND DESIGN

Functions
- protects and supports the brain and its sensory receptors
- may house cooling equipment to cool the brain during sustained
activity or during a rise in environmental temperature
- in many active terrestrial mammals, the nasal epithelium lining the
nasal passages dissipates excess heat by evaporation as air moves
across this moist lining
- the skull of many animals also supports the voice box and
occasionally serves as a sound resonator to deepen or amplify an
animal’s call

Feeding in Water
- the higher viscosity of water presents both problems and opportunities
for the animal feeding in water
Problem: feeding in water poses a disadvantage in that water easily - the highly kinetic skull of snakes allows great freedom of jaw
carries shock or pressure wave (bow waves) immediately in front of the movement
predator approaching its food - in many vertebrates, swallowing involves mastication (the chewing of
food)
Opportunity: when a vertebrate quickly gulps water into its mouth, the - within animals, mastication has had a profound influence upon skull
viscosity of the water drags along the prey as well --suction feeding design, producing an akinetic skull with precise tooth occlusion and
only two replacement sets of teeth, a secondary palate, large
Suspension feeding jaw-closing musculature, and changes in lower jaw structure
- the capture and ingestion of food particles that are suspended in
water SKELETAL SYSTEM: The Axial Skeleton
- most suspension feeders are benthic (bottom-dwelling) organisms
and herbivorous/detritus feeding style Introduction
- two structural component combine to define the long axis of the
Mechanism: vertebrate body, offer sites for muscle attachments, prevent
1. Captured particles are usually smaller than the pores of the filter telescoping of the body, and support much of the weight
2. They may collide directly with the filter or because of their inertia,
they deviate from the streamline to collide with the mucus-covered Notochord
surface of the filter - a long, continuous rod of fibrous connective tissue wrapping a core of
3. Upon impact, the particles cling to the sticky mucus and are rolled fluid or fluid-filled cells
up in mucous cords, and then are passed by cilia into the digestive tract
Verterbral column
Least common: - consists of a discrete but repeating series of cartilaginous or bony
- a Sieve can be used to strain suspended particles larger than the elements
pores of the sieve The notochord is phylogenetically the oldest of the two structural
- as the stream of water passes through the sieve, the particles are components, but it tends to give way to the vertebral column, which
held back and then collected from the face of the selective filter assumes the role of body support in most later vertebrate
- this method is rare among animals because the relatively large
particles filtered tend to plug and foul the sieve Basic Components
Ex: buccal cirri (amphioxus); gill rakers (fishes) - vertebrae
When the filter is clogged, these fishes can clean the material by a kind - ribs
of cough or quick expansion of the gill arches - sternum
- gastralia
Suction feeding
- a method of ingesting a prey item in fluids by sucking the prey into the Vertebrae
predator’s mouth - first components of the vertebra to appear were the dorsal and
- a highly coordinated behavior achieved by the dorsal rotation of the ventral arches that rested upon the notochord
dermatocranium, lateral expansion of the suspensorium, and the - the dorsal arches, neural and interneural (intercalary) arches,
depression of the lower jaw and hyoid protected the neural tube
- the buccal cavity expands rapidly, pressure drops, and food is - the ventral arches, hemal and interhemal arches, enclosed the
aspirated into the mouth blood vessels
- the second component is the formation of two centra: an
In fishes and larval salamanders intercentrum (hypocentrum) and a pleurocentrum
- excess water, gulped in with the food, exits via gill slits at the back of - the bases of the ventral arches expand to form these centra where
the mouth they meet the notochord
- the centra served to anchor and support these arches
In adult salamanders, frogs, and aquatic vertebrates
- gills slits are absent, so excess water entering the mouth reverses
its direction of flow to exit via the same route

Feeding in Air
- terrestrial feeding in most amphibians and many lizards requires a
projectile tongue

Lingual feeding
- when an animal uses an oral muscular organ, most notably the
tongue that’s normally enlarged for foraging and hunting and acts as a
tool to obtain certain food items that take up a significant part in the
animal’s diet

Prehension
Centrum
- a method by which the animal rapidly grasps the prey with its jaws or
- each centrum constitutes the body of the vertebra
the process of seizing or grasping or otherwise getting food into the
Aspondydly - centra may be absent
mouth
Monospondyly - one body
- in such animals, the jaws are prey traps, designed to snare the
Diplospondyly - two centra per segment
unwary
- the pleurocentrum predominates and becomes the body of each
- does not always involve the jaws
vertebral segment
- jaws are used secondarily to help hold the struggling victim or to
- the intercentrum becomes the intervertebral cartilage (disk) of the
deliver a killing bite
amniote vertebral column
Ex: birds of prey snatch quarry with their talons; mammalian predators
often use claws to catch and then control intended prey

Swallowing
- the act of passing food from the mouth, by way of the pharynx (or
throat) and esophagus, to the stomach
- in suspension feeders, the food-laden cords of mucus are swept by
synchronized ciliary action into the esophagus
- other animals usually swallow prey whole or in large pieces
- in suction feeders, they rapidly expand the buccal cavity repeatedly to
work the captured prey backward into the esophagus
- in terrestrial vertebrates, they use the tongue to reposition the food
bolus and work it toward the back of the mouth
- in snakes, they swallow a relatively large animal by stepping the
tooth-bearing bones over the dispatched prey
Vertebral Types Intervertebral disk
- in adults, it is a pad of fibrocartiliage who gel-like core, the nucleus
1. Aspidospondyly pulposus, is derive from the embryonic notochord
- all elements are separate specifically the three arch elements - found only in mammals, in whom they reside between successive
(intercentrum, pleurocentrum, and neural arch) surfaces of adjacent centra
- the term means “cut-up spine”, a reference to the numerous separate - in other groups, it is called and intervertebral cartilage or body
parts that constitue each vertebral segment
2. Holospondyly Intervertebral ligament
- all vertebral elements in a segment are fused into a single piece - joins the rims of adjacent centra
- important in controlling the stiffness of the vertebral column when it
flexes

Axial column
- the centra chained , making a chain of vertebrae
- shapes of surfaces at the articular ends of the centra affect the Ribs
properties of the vertebral column and the way in which forces are - struts that sometimes fuse with vertebrae or articulate with them
distributed between vertebra - provide sites for secure muscle attachment, help suspend the body,
form a protective case around viscera (rib cage), and sometimes serve
General Centra Shapes as accessory breathing devices
1. Acoelous
- centra with flat ends and especially suited to receive and
distribute compressive forces within the vertebral column
- flex about a point on their rims, tending to stretch the centrally
located dorsal nerve cord
2. Amphicoelous
- if each surface is concave and the centrum is designed to allow
limited motion in most directions
- *function is the same as that of acoelous
3. Procoelous
- centra is concave anteriorly and convex posteriorly
- centra eliminate this potentially damaging stretching tendency
with ball-and-socket ends that establish a centrally located point of
rotation instead of one at the rims
4. Opisthocoelous
- the reverse shape, concave posteriorly and convex anteriorly
- *function is the same as that of procoelous In fishes
5. Heterocoelous - there are two sets of ribs with each vertebral segment, a dorsal and a
- centra that bear saddle-shaped articular surfaces at both ends ventral set
- centra, opposite, saddle-shaped surfaces fit together, allowing
extensive lateral and dorsoventral rotation In tetrapods
- ventral ribs are lost and the dorsal ribs, persists to become the trunk
ribs of terrestrial vertebrates
- ribs of primitive tetrapods are bicipital, having two head that articulate
with the vertebrae
- the ventral rib head, or capitulum, articulates/joints with the
parapophysis, a ventral process on the intercentrum
- the dorsal head, or tuberculum, articulates/joints with the
diapophysis, a process on the neural arch

True ribs
- ribs that meet ventrally with the sternum
- consist of two jointed segments

Vertebral (costal) rib

- a proximal segment articulated with the vertebrae

Sternal rib
- a distal segment that is usually cartilaginous and meets the sternum
 - the sternum may consist of a single bony plate or several elements in
False ribs series
- ribs that articulate with each other but not with the sternum - fishes, turtles, snakes, and many limbless lizards lack a sternum
- absent in the first fossil tetrapods, but it is present in modern
Floating ribs amphibians
- those false ribs articulating with nothing ventrally

In urordeles
- the sternum is a single midventral sternal plate grooved along its
anterior borders to receive the ventral elements of the shoulder girdle,
the procoracoid plate
IN REPTILES
In anurans
Turtles - a single element, the xiphisternum, often tipped with the xiphoid
- have no cervical ribs, and the ribs of the trunk are fused with the cartilage, lies posterior to the pectoral girdle and in some, a second
costal plates of the carapace element, the omosternum capped by the episternal cartilage, lies
- the two sacral ribs are not fused with the carapace anterior to the girdle

Snakes
- have a long, curved ribs beginning at the second vertebra and
continuing far into the tail
- there is no sternum for ribs to attach to; however, the ventral ends of
the ribs have ligamentous connections with integumentary scutes
- these ribs participate in locomotion

IN BIRDS
- in the thoracic region, the first several ribs are floating ribs, followed
by true ribs that articulate with the sternum

Cervical ribs
- reduced and fused to the vertebrae
In flying birds
- the massive flight muscle arise from a large sternum that bears a
Uncinate processes
prominent ventral keel, the carina
- projections that extend posteriorly from proximal rib segments
- present in some floating and most true ribs
Carina
- like the rib cage, it generally offer sites of attachment for respiratory
- provides additional surface for muscle attachment
and shoulder muscles

In mammals
- the sternum consists of a chain of ossified elements in series, the
sternebrae
- the first, manubrium and last, xiphisternum, bear a cartilaginous or
bony xiphoid process

Sternum
- a midventral skeletal structure that offers a site of origin for chest
muscles
- also secures the ventral tips of true ribs to complete the protective
chondrified or ossified rib cage

Rib cage
- consists of ribs and sternal elements that embrace the viscera
- size and shape changes in the rib cage also act to compress or
expand the lungs, promoting ventilation
 Gastralia Terrestrial environment
- also known as abdominal ribs
- a separately derived set of skeletal elements located posterior to the Problem: Gravity
sternum in some vertebrates - when remaining in place, the tetrapod’s body either rests on the
- of dermal origin ground between sprawled legs, or it is suspended between pairs of
- restricted to the sides of the ventral body wall between sternum and legs, as in most mammals
pelvis and do not articulate with the vertebrae - the pairs of legs function as abutments/brace that support the body
- common in some lizards, crocodiles, and Sphenodon between them
- serve as an accessory skeletal system that provides sites for muscle - the vertebral column serves as a bridge between the support posts,
attachment and support for the abdomen the legs, and suspends the body from it

In turtles
- the plastron is a composite bony plate form the floor of the shell
- it consists of a fused group of ventral dermal elements, including
contributions from the clavicles (epiplastrons) and interclavicle
(entoplastron) as well as dermal elements from the abdominal region
(possibly the gastralia)

Regional Specialization in Tetrapod Columns

- in tetrapods, the trunk becomes further differentiated anteriorly into

the neck, or cervical region, and posteriorly into the hip, or sacral
region
- there is further differentiation of the trunk into the chest, or the
thoracic region, and into the area between the thorax and the hips,
the lumbar region

In fishes
- the vertebral column is differentiated into two regions: an anterior
trunk region and a posterior caudal region
- the relatively undifferentiated vertebral column of fishes reflects the
fact that it is not used to support the body
- support comes generally from the buoyancy of the surrounding water
FORM AND FUNCTION - the vertebral column mainly offers sites of attachment for the
swimming musculature
Fluid environment - it serves as a mechanical replacement of the notochord and
- an organisms does not depend primarily on the endoskeletal allows lateral flexibility for swimming
framework for support, instead, the body takes advantage of its
buoyancy in the surrounding water

For an active aquatic organism, two problems are uppermost

1. The drag on the body as it slips through a relatively dense

medium, water.
In tetrapods
The answer is streamlining, contouring of the body to reduce drag
- the vertebral column supports the body agains gravty and receives
forces
and transmits the propulsive forces that limbs generate during
- the general body shapes of fast-swimming fishes are streamlined
locomotion
- this shape improves the performance of fishes as they meet common
- diverse functional demands are placed on the vertebral column, so we
physical demands while traveling through a medium that resists
might expect to find delineation of specialized regions
passage
1. Sacral vertebrae - for locomotion and support
2. Orientation in three-dimensional space
2. Cervical vertebrae - to increase mobility of the head
3. Thoracic vertebrae - for external respiration
- any streamlines body has a tendency to tip and deviate from its line
4. Lumbar vertebrae - remaining segments anterior to the sacral
of travel, rotating about its center of mass
vertebrae
- in fishes, these perturbations are countered, by stabilizing fins
appropriately positioned along the body
- the body of a fish can deviate from its intended line of travel in three
ways:
A. A roll rocks the fish about its long axis
B. A yaw swings it from side to side
C. A pitch bucks it up and down about its center of mass

In snakes
- they have the longest columns with as many as 400 or more and
almost no regional specialization
In anurans - most mammals have three to five
- shortest columns
Sacrum
In birds and turtles - single bony complex formed when the sacral vertebrae of mammals
- only cervical and caudal segments are flexible ankylose
- most of the trunk vertebrae are being rigidly fused to the synsacrum in
birds, and carapace in turtles Synsacrum
- adult complex formed when the last thoracic vertebra, all lumbars,
The craniovertebral junction and neck vertebrae sacrals, first few caudals and the ribs of modern birds ankylose

In amphibians Tail Vertebrae

- the first and only cervical vertebra of modern amphibians lacks - numbered 50 or more
processes - in modern ones, the number is much reduced and highly variable
- the absence of processes and the nature of the amphibian
craniovertebral join permits limited dorsoventral rocking of the skull Tail end
- arches and processes become progressively shorter and rudimentary
In amniotes
- the first two vertebrae are atlas and axis
- the first vertrbra, or atlas, is ringlike bone and lacks centrum
- the centrum of the atlas has become the odontoid process of the
second vertebra, or axis
In bony proatlas, resembling a neural arch is interposed between the
skull and atlas in crocodilians Urostyle
- caudal vertebrae in anurans
In birds - develops from a continuous elongated perichordal cartilage at the
- cervical vertebrae are flexibly articulated to give the head great base of the larval tail, and it grows and ossifies after the tail is lost at
freedom of movement and reach when a bird preens its feathers or metamorphosis
probes for food
Pygostyle
Heterocelous centra - in birds
- combined with the atlas-axis complex and the largest number of - the skeleton of the visible part of the tail
cervical vertebrae in any tetrapod (12 commonly to 25 in swans) enable - develops as a 4 separate cartilaginous centra
some birds to turn their heads 180 degrees to the rear, enlarging the
arc of their horizon to that of a complete circle Coccyx
- in apes and humans
- “tailbone:
- the centra of the coccyx are still identifiable, but the last one is a mere
nodule of bone

In turtles
- uniquely flexible neck
- ball-and-socket joint in the procelous centra enables the entire head
and neck to be retracted into the shell

In mammals
- always have seven cervical vertebrae

Sacrum and Synsacrum

Sacral vertebrae
- bear stout transverse processes to bear the thrust of the pelvic girdle
- amphibians have one sacral vertebral
- living reptiles have two