&

-
- Chapter 3: The Vertebrate Story -

=
Two Vertebrate Innovations
Vertebrate Column
more muscles to attach
Cranium (Craniates)
& specialized organs
cephalization (clustering of
-
involves concentrating neurons into a brain at one end of the animal
and evolving sensory organs at that same end
-

arthropods , annelids and chordates undergo cephalization

,

>
major regions : forebran) most anterior),
All vertebrate brains consist of 3
-

midbrain and windbrain (most posterior). The most posterior part of the
,

hindbrain is the medulla

oblongata.
Evolution of Early Vertebrates
Step 1 : Prevertebrates
-

Suspension -
feeders
-

deployed only cilia to produce the food-bearing current that entered the pharynx
-

incipient predator
Step 2 : Agnathans
-

Jawless fish
-

includes the jawless vertebrates ,

like hagfishes and lampreys
Step Gnathostomes
3 :

became raptorial and suction feeders

-

active predators
-

evolution of jaws
-

with the evolution of jaws it marked the next step for vertebrates to become
,

effective predators
 Vertebrate Classification
Agnathans (Jawless Vertebrates)
The first vertebrates were agnathans, lacking jaws but possessing the vertebral
column. They also possess a cartilaginous skeleton. They include extinct groups
and living agnathans like:

• Myxinoidea (Hagfish): Primitive living vertebrate, eel-like, and scavengers.

• Petromyzontida (Lampreys): Parasites and filter-feeders.

Early Vertebrate Fossils

• Conodonts: Early vertebrate-like organisms with tooth-like elements.
• Ostracoderms: Early armored jawless fish, including:
Pteraspidomorpha: Heavily armored fish.
Osteostracans (characterized by their bony armor and paired eyes, often
displaying a flattened body), Anaspids (known for their smooth, scaleless skin
and streamlined shape, lacked bony plates and had prominent eyes), and
Thelodonts (featured tooth-like scales and a distinct head structure, often
resembling early jawed vertebrates):
Various other early agnathan groups with distinct features.

Overview of Agnathan Evolution

• In general, they paved the way for the development and evolution of jawed
vertebrates.
• Bones were first evolved in ostracoderms, basal to gnathostomes.
• Characterized by a lack of jaws, paired fins, and a bony skeleton, relying
instead on a cartilaginous structure and a notochord.
• The evolution of jaws in gnathostomes is believed to have arisen from
modifications of the first pair of gill arches in ancestral agnathans.
Gnathostomes (Jawed Vertebrates)
The evolution of jaws, which allowed for more diverse feeding strategies. Led
also to the development of paired fins. Went also from detritus feeder to
active predation.

Evolved by modification of skeletal rods. Gnathostomes include:

• Placoderms: Extinct, heavily armored jawed fish with the presence of

head shield
• Chondrichthyes: Cartilaginous fish, including:
Elasmobranchii: Sharks and rays, with the development of tooth
replacement. They are predators and planktivores
Holocephali: Chimaeras, with distinct head morphology and development of
operculum (an extended flap of skin).

Teleostomi (Bony Fish and Related Forms)

• a large group that encompasses the acanthodians (sister group of the
bony fishes) and their tetrapod derivatives.
• Acanthodians: Spiny fish with both primitive and derived traits. Their gill
slits were covered by a body operculum. Its dermal armor reduced to scales
across the body surface. Have a partially ossified internal skeleton.
• Osteichthyes: Bony fish, with gas-filled swim bladder for buoyancy.
Presence of bone in the endoskeleton (with exception to sturgeons, paddle
fishes and lung fishes). Split into:
Actinopterygii: Ray-finned fish that is internally supported by slender,
endoskeletal lepidotrichia (rays). Muscles are located within the body wall that
controls fin movements.
Sarcopterygii: Lobe-finned fish or fleshy-finned fish, which gave rise to
tetrapods. Its muscles are located outside of the body wall along the
projecting fin. Were once known as Choanichthyes in recognition of external
nostrils opening internally to the mouth through holes termed choanae.
Overview of Fish Phylogeny

• Fish are the most diverse group of vertebrates, with multiple evolutionary
branches.

• All modern fishes, except cyclostomes, belong to either Chondrichthyes

(cartilaginous fishes) or Osteichthyes (bony fishes).

• Ostracoderms, already featured significant bone structures, while later groups

showed varying trends in ossification, with chondrichthyans losing bone and
osteichthyans gaining it in their endoskeletons.

• Ray-finned fishes have dominated aquatic environments since the mid-Paleozoic,

while fleshy-finned fishes eventually gave rise to tetrapods.

• They are pivotal in the vertebrate evolution, with them being the first to
develop jaws and fins.

• It led to the evolution of tetrapods.

Tetrapod Evolution
• means “four-footed”
• characterized by Chridium (muscular limb with well defined joints and digits)
• evolved from Sarcopterygii and transitioned from water to land.
Tiktaalik Characteristics of Fish and Tetrapod
• exhibits both

/
Fish Tetrapod
fish and • scales • neck
tetrapod • fins • ribs
characteristics • gills and lungs • fin skeleton
• vertebrate • flat skull
transition from • eyes on top of
water to land the skull
 • Primitive Tetrapods: Early four-limbed vertebrates that still retained aquatic
traits.
• Labyrinthodonts:
-Early amphibians with complex tooth structure.
-Transition between within Sarcopterygians and terrestrial tetrapods.
-Direct ancestral of modern day amphibians.
• Lissamphibia (Modern Amphibians)
-Arise within the labyrithodont radiation.
-Includes frogs, salamanders, and caecilians.
-The egg lacks shell and amniotic membranes.
-Undergoes metamorphosis.
-Presence of paired lungs but maybe reduced or absent in some salamanders.

Includes three groups:

Urodela (Caudata): Salamanders; having paired limbs and a long tail. In
primitive salamanders, fertilization occurs externally, while in more advanced
groups, males produce a spermatophore that females collect to facilitate internal
fertilization of their eggs within the reproductive tract. This process enhances
sperm transfer and ensures successful fertilization.
Salientia (Anura): Frogs and toads, with fertilization is mostly external. Eggs
are mostly laid in water or moist locations. The tadpole larvae is a striking
characteristic of frogs.
Gymnophiona (Apoda): Limbless amphibians or “no feet (apodans)”. All are
restricted to tropical habitats, where they live in burrowing lifestyle. The caecilian
skull is solid and compact.
Common Characteristics of Living Amphibians
• cutaneous respiration- mucous glands to keep skin moist
• auricular operculum- extra ear used to transmit signals
• pedicellate teeth- calcified root and crown but separated by non-calcified tissue

LEPOSPONDYLS
Distinct Features:
• Lepospondyls differ from labyrinthodonts primarily through skeletal
characteristics, including solid vertebrae that are fused onto a single, spool-
shaped centrum
• reduced skull bones
• Paired limbs were small and ossification was reduced
• The tail of some species was quite long
Habitat and Lifestyle: While primarily terrestrial, some lepospondyls were aquatic,
showing varied adaptations.

AMNIOTES
• Definition:
Amniotes bear embryos with extraembryonic membranes, typically in shelled
eggs.
Fossil animals rarely provide direct evidence of reproductive style; phylogenetic
studies help establish relationships.

• Two Major Lineages:

Sauropsida: Includes birds, dinosaurs, modern reptiles, and diverse Mesozoic
forms.
Subdivisions: Parareptilia and Eureptilia.
Synapsida: Monophyletic lineage leading to therapsids and modern mammals.
Skull Fenestration
• Temporal region of the skull is a key indicator for evolutionary relationships.
• Skull Types
Anapsid: No temporal openings; found in primitive amniotes and turtles.
Synapsid: One pair of temporal openings; associated with mammalian ancestors.
Diapsid: Two pairs of temporal openings; includes pterosaurs and dinosaurs.
Euryapsid: A modified diapsid skull with loss of the lower temporal opening;
found in some Mesozoic marine reptiles (plesiosaurs, ichthyosaurs).
Stem-Amniotes
The diadectomorphs, Diadectes, are considered the most likely sister group
to amniotes.
These primitive tetrapods emerged in the Late Carboniferous and show
some evidence of herbivory.
Diadectomorphs are key transitional forms between non-amniote and
amniote tetrapods.
They share affinities with amphibians and are linked to anthracosaurs.
The relationships among early amniotes are still under investigation.
Diadectomorphs may be classified within anthracosaurs or potentially revisited as
part of the old term Cotylosauria.
Anthracosaurs:
Historically, anthracosaurs have included various late non-amniote groups
and are thought to be closely related to or ancestral to amniotes.
The term "Cotylosauria" refers to a basal group of amniotes, meaning "stem
reptiles," from which all later amniote groups evolved.

Key Terms to Remember:

• Diadectomorphs: Primitive, herbivorous tetrapods.
• Anthracosaurs: Group believed to be closely related to amniotes.
• Cotylosauria: Old term for the basal group of amniotes.