Lecture 10 The Dermatocranium and Axial Skeleton

KK Chapter 7,8; H&G Chapter 8,9

The dermatocranium changes relatively little from early bony fishes through to mammals. Many of the changes involve simplification by loss of bones, especially the back of the skull, in the transition from fish to tetrapod. (Fish have no necks, tetrapods do!) However, the dermatocranium changes more among teleost fishes where adaptations for feeding cause extensive modifications. We will not cover these changes. We will start by looking at the skull of Amia, a primitive bony fish, and follow the changes to mammals.

Amia KK 7.23, H&G 8.11

Amia has a skull typical of early Osteichthyes, and not unlike Crossopterygii. Many of the dermatocranium elements are homologous with those of tetrapods. Early tetrapods lack the opercular bones and the bones connecting the pectoral girdle to the head and in that location they have instead a flexible neck. The bones shaded in pink are part of the pectoral girdle. The red line marks the part of the skull that is lost with the gills.

Dermal Bones of a Tetrapod

TopView Palate KK 7.10 H&G 8.6 The major difference relative to Amia is the loss of bones at the back of the head. This trend will continue with loss of postparietals and temporals, most of the orbital series, and simplification of the lower jaw.

Early Amphibian (KK 7.37, H&G 8.12)

Early Reptile (KK 7.33)

Turtle (KK 7.37, H&G 8.15)

The dermal skull roof changes relatively little in tetrapods, except for appearance of openings in the temple. In many mammal and birds the temple opening converges with the orbit.

What are euryapsids?

KK 7.34,H&G 8.18

Temporal Openings in Tetrapods

KK 7.35, H&G 8.17 Frontal sections.

Temporal openings allow for muscles closing the jaws to bulge, and provide better points of attachment. Note the zygomatic arch. Some mammals have a sagittal and/or nuchal crest on the top or back of the head to provide even better attachment points.

The temple opening in humans. Humans are among the mammals (apes, artiodactyls) where the temple opening is not confluent with the orbit.

From Internal Nares to Secondary Palate

KK 7.27 early reptile (Parareptilia)

KK 7.57, H&G 8.19

Choanate vertebrates (Sarcopterygii, tetrapods) have a connection between the external naris and the mouth. In mammals, this connection between respiratory and digestive pathways moves to the back of the mouth because of the hard and soft palate.

The Secondary Palate

KK 7.58, H&G 8.19

The secondary palate is formed by folds in the marginal bones of the palate (premaxilla, maxilla and palatine). The space between the original and secondary palate in mammals contains turbinate bones. The secondary palate is also found in turtles and crocodiles. Why did this evolve? And what are the implications for the evolution of endothermy? Palatine fissures?

Review: the skull of the opossum, Didelphis KK 7.51

The Axial Skeleton

KK Chapter 8, H&G Chapter 9

Parts of the Axial Skeleton

Notochord Vertebral Column Ribs, sternum, gastralia Median Fins (caudal, dorsal, anal)

Embryological origins notochord, schlerotome and mesenchyme

The Notochord
found in all Chordata found in all vertebrate embryos, plus adults of Agnatha, Placodermi, Acanthodii, Holocephali, some Actinopterygii (not teleosts), many Sarcopterygii, and the first Amphibia mechanically important for swimming

Centra in bony fish provide the same mechanical properties as the notochord.
KK 8.19

intervertebral ligament

Fish with notochords have bony or cartilaginous pieces (neural or hemal arches or spines) associated with the notochord. Fish with vertebrae tend to have biconcave (amphicoelous) centra. Tetrapod centra tend to be acoelous (platyan); more stout. Procoelous or opisthocoelous centra may be found where more flexibility is required, e.g. tail or neck.

KK 8.4, H&G 9.2 KK 8.1, H&G 9.4

Vertebrae of Fish
KK 8.6

The ribs are between the muscle blocks (myomeres) of the fish trunk. The dorsal ribs follow the horizontal septum between the epaxial and hypaxial muscles.

Development of the Vertebral Column

Vertebrae arise from the scherotome, and begin as two elements per segment. In most cases they fuse to make a single centrum, but do so in an intersegmental fashion. Where they remain separate, they are called pleurocentrum and intercentrum.

KK 8.10 & 8.13, H&G 9.3

Evolution of the vertebra in the Amphibia

A labyrinthodont with equal interand pleurocentra (above). In the lineage leading to amniotes, the pleurocentra dominate. KK 8.24, 8.26. H&G 9.7

Caudal Fins
The tail fin is supported by flattened neural and especially hemal arches.

The tail of fishes with homocercal tails begin as heterocercal.

KK 8.20,H&G 9.14

The impact of terrestrial life on the axial skeleton

Centra expand, fuse and replace the notochord. Intervertebral joints and processes (zygapophyeses) develop that restrict or facilitate movement depending on the location

Processes to increase the leverage of muscles

Connection to limb girdles. More flexible relationship with the head via cervical vertebrae + atlas (and axis).

Tetrapod Vertebrae
KK 8.7, H&G 9.1 Zygapophyses resist the tendency of the trunk to sag, and ribs now have two attachments to the vertebral column and meet at the ventral mid-line (sternum). Tetrapods do not have dorsal ribs, but their ribs often have two heads and attach to the sternum.

Regional Specialization of Vertebrae

cervicals - small and flexible, with ribs small or absent thoracics - ribs, no hemal arches, reach sternum in tetrapods lumbars - no ribs, very stout sacrals - with heavy sacral ribs attaching to pelvic girdle caudals - flexible, with ribs small or absent. May be hemal arches, intercentra.

Regional Specialization of Vertebrae

Fishes: thoracics and caudals only Amphibia: atlas, 1 cervical, thoracics, 1 sacral, and caudals Reptilia: atlas, axis, cervicals, thoracics, 2+ sacrals, caudals Aves: atlas, axis, cervicals, thoracics, synsacrum, pygostyle Mammalia: atlas, axis, cervicals, thoracics, lumbars, 3+ sacrals, caudals

Atlas and Axis

In amniotes, the first two vertebrae (atlas and axis) are specialized for nodding the head (occipital condyles articulate with the atlas) and rotating the head (atlas rotates on the axis on the axis). The latter joint is actually between the pleurocentrum of the atlas, now fused to the axis, and the intercentrum of the atlas.

KK 8.27, H&G 9.10

Bull Skeleton