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Anamniote Skeletons

Anamniote Skeletons

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INTRODUCTION<br />

Dermal bones are derived from dermal scales that sank deep & joined the internal skeleton. These bones form as connective<br />

tissue membranes that later ossify. Endochondral or replacement "elements" begin as cartilage & may ossify or remain cartilaginous<br />

in the adult.<br />

The axial skeleton elements are on the midline or mid-sagittal axis of the body. The axial skeleton includes the notochord,<br />

vertebrae, ribs & median fins. The notochord is a centrally located, gel-filled rod that supports the body of a developing embryo. In<br />

most vertebrates, vertebrae replace it, partly or completely. The vertebral body or centrum is the large solid disk that takes the<br />

compressive forces during body movement. Some centra are biconcave (amphicoelous) or flat (acoelous). In fishes, amphicoelous<br />

vertebrae are tightly held together with sheathing of connective tissue. Other centra have rounded cavities on the anterior (procoelous)<br />

or posterior (opisthocoelous) surface with a corresponding rounded opposite side that articulates with adjacent vertebrae. The<br />

vertebrae enclose the spinal cord with neural (or vertebral) arches. Caudal vertebrae have hemal arches that enclose caudal arteries<br />

& veins in the tail. All vertebrae have several areas of muscle attachment via neural spines, transverse processes & hemal spines.<br />

Tetrapod vertebrae have zygapophyses (zygo= yoke) to support the body & keep the trunk from sagging. The zygapophyses<br />

allow vertebrae to articulate with each other, and limit the total range & direction of body movement. The pre-zygapophyses are on the<br />

anterior side of vertebra & have articulating facets that face upward. The post-zygapophyses are on the posterior side & have<br />

articulating facets that face downward. Post-zygapophyses fit on top of pre-zygapophyses.<br />

Fishes have 2 regions in vertebral column: trunk vertebrae have ribs and caudal vertebrae form the tail. <strong>Anamniote</strong> tetrapods<br />

have 4 regions in vertebral column. They have 1 cervical or neck vertebra, called the atlas. This vertebra has large facets (prezygapophyses)<br />

that articulate with the paired occipital condyles seen in living amphibians. Tetrapods have a single sacral vertebra<br />

that attaches to the pelvic girdle. In frogs, the caudal vertebrae are fused into a rod-like urostyle.<br />

A sternum is present only in tetrapods, and even then may be missing or very small in a few species. The sternum acts as a<br />

brace to attach to support the rib cage or an additional brace for the pectoral girdle. Fishes have unpaired, median fins located along the<br />

mid-sagittal axis. These include dorsal, anal & caudal fins. Fish may have 1 or more dorsal fins. Caudal fins vary in design.<br />

Heterocercal caudal fins are present in the Chondrichthyes and some primitive Actinopterygians. In these tails, the caudal vertebrae<br />

extend into the dorsal & larger lobe of the caudal fin. Some Actinopterygians have a hemi-homocercal design where the vertebrae just<br />

turn dorsally at the very tail tip & do not extend far into the tail fin. The tail fin doesn’t appear to be asymmetrical, but the asymmetry can<br />

be seen in the skeleton & in the body profile. Most living Actinopterygians have a homocercal caudal fin that forms both a symmetrical<br />

profile & symmetrical caudal fin.<br />

The appendicular skeleton refers to the elements that form the paired, laterally placed appendages for the paired front & hind<br />

fins or limbs. The pectoral girdle of fishes is a combination of dermal & endochondral elements. The dermal pectoral girdle bones of<br />

ray-finned fish form an arc just behind the opercular bones of the skull. These dermal bones include the cleithrum, clavicle & other<br />

bones that attach the pectoral girdle to the skull. The sharks lack these dermal bones as they lack all dermatocranium in the skull as<br />

well. Tetrapods lose many dermal bones when the pectoral girdle detaches from the skull. Thus in tetrapods, the head moves<br />

independently of the legs. Some tetrapods retain the clavicles as paired bones that extend from the sternum towards the humerus. One<br />

new dermal pectoral girdle bone, the interclavicle, appears in Sarcopterygians & some tetrapods. This is an unpaired bone that may be<br />

in the midline between the paired clavicles.<br />

The endochondral elements of the pectoral girdle are called scapulocoracoid or coracoscapula, if they don’t ossify. In<br />

derived Actinopterygians (e.g. Perch) these ossify into a ventrally positioned anterior coracoid (procoracoid) & a more dorsally<br />

positioned scapula. The tetrapods may add a suprascapula dorsal to the scapula. Amniotes gained a new element, the posterior<br />

coracoid (metacoracoid). In<br />

Fish pelvic girdles are suspended in muscle: ischiopubic or pubioischiatic plates. In derived Actinopterygians this girdle &<br />

its associated fins are anterior in the body. This position may improve the braking ability of those fins. In some ray-finned fish, the pelvic<br />

girdle attaches to the pectoral girdle but is still ventral to the pectoral fins. All tetrapods (with legs) have 3 bones that form the pelvic<br />

girdle. The pubic bone or pubis is directed forward & down & remains cartilaginous in many anamniotes. The ischium faces<br />

downward & posteriorly. The ilium is the new bone in the tetrapod pelvis. It is oriented dorsally & attaches the pelvic girdle to the sacral<br />

vertebra.<br />

The median & paired fins of fishes contain the same types of support elements. The most proximal support elements are<br />

called basal pterygiophores & they are typically larger and fewer in number than the more distal radial pterygiophores. Different<br />

materials in fish support the fin membranes. Ceratotrichia are made of keratin & support the fins of sharks, skates & rays.<br />

Lepidotrichia are tiny, overlapping dermal scales in Actinopterygian fins. Actinotrichia are bony spines that may be present at the<br />

anterior edge of a fin or replace lepidotrichia throughout an entire fin.<br />

The paired limbs of tetrapods (with limbs) comprise a standard pattern with 1 bone, followed by a pair of bones then many<br />

bone in series. The forelimb has a humerus; radius & ulna; carpals; metacarpals; phalanges. The hind limb has a femur; tibia & fibula;<br />

tarsals; metatarsals; phalanges.

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