VERTEBRATES: Evolution and Diversity
The earliest ancestors of vertebrates were plate-skinned fishes known as
OSTRACODERMS. (from the Greek ostraco meaning "shell" and
derm meaning "skin"). This was a diverse array of armored fishes
that lacked a lower jaw (mandible) or paired fins (remember the
lancelets! Do they have paired fins? What is the advantage of paired
appendages?).
One group of these fish (OSTEOSTRACANS) evolved paired pectoral fins just
behind the head. A separate group (ANASPIDS) closely resembled modern-day lampreys.
Vertebrates, as we already have discussed, are either
- AGNATHANS - lacking a lower jaw
- GNATHOSTOMES - having a mouth with a lower jaw
Only the latter group is monophyletic, as jawed vertebrates all show
similar embryogeny with respect to jaw formation. (The jaw bones develop
from portions of the cartilaginous arches that once formed the support of
the anterior gill slits.) The jawless
vertebrates, however, showing the primitive condition, may have had
several different ancestors.
The Devonian period saw the rise of the first jawed fishes, the PLACODERMS, all of which are now extinct. Another
group, the ACANTHODIANS which lived at the
same time as the placoderms, may have included the ancestors of all modern
bony fishes.
Let's meet the Vertebrates, Class by Class
SUPERCLASS AGNATHA - The Jawless Fishes
-
Class Myxini - The Hagfishes
(You are responsible for all the information on hagfishes on the first
page linked above.)
-
Class Petromyzontiformes - The Lampreys
(You are responsible for all the information on lampreys on the first
page linked above.)
SUPERCLASS GNATHOSTOMATA - The Jawed Vertebrates
I. Fishes
Vertebrate jaws evolved from the skeletal supports of the anterior pharyngeal gill
slits. The posterior slits were retained as respiratory structures.
-
Chondrichthyes - The Cartilaginous Fishes (sharks, skates,
rays)

Actinopterygii - The Ray-finned Fishes
- These are the most numerous of vertebrates, both in species (about 30,000) and
individuals.
- Vast diversity in form, function and natural history
- All bony fishes were once placed in "Class Osteichthyes",
which was determined to be polyphyletic. That traditional class has
since been divided into three (putatively)
monophyletic taxa:
- The lungfishes are believed to share a more recent common ancestor
with tetrapods than do other groups of fish.
Fun Fish Facts
Skeleton composed primarily of bone
ectothermal poikilotherms
Tail is primitively homocercal (see what happens if you don't come to
class?)
Skin covered with dermal scales that shed readily.
Skin copiously supplied with mucous glands
Fins may be either medial and unpaired, or lateral and paired
Primitively, fins have rays made of cartilage or bone
Toothy, terminal (i.e., at the end of the body) mouth
Gill arches covered by a bony operculum
Swim bladder, derived from the digestive tract, may or may not be open
to the pharynx
Heart is two chambered (one ventricle (caudal) and one atrium (cranial)
Brain differentiated, with small olfactory lobes and cerebrum; large
optic lobes and cerebellum (what does this tell you about what fish do
best, and what they might not be quite as good at?)
Most species dioecious, a few hermaphroditic; some change sex during
the lifespan, depending on environmental circumstances
The Amazing Swim Bladder
Many pelagic fish (but not all) have this gas-filled sac found dorsal
to the digestive tract
In some species, it is connected by the pneumatic duct to the
esophagus or pharynx. (These fish gulp or burp air to control the amount
of gas in the swim bladder.)
In other species, the swim bladder is not connected to the outside
world. This can be a problem! What happens to the bladder when the fish
goes deeper (HINT: higher pressure)? What happens when the fish goes more
shallow?
- In these species, the gas gland collects gasses dissolved in the
blood stream and secretes them into the swim bladder
- The resorptive area removes gas from the bladder
- These processes take time, so species without a pneumatic duct
generally cannot make drastic changes in swim depth quickly.
These are the four-legged vertebrates, derived from a specialized group of
shallow-water fishes.
A monophyletic lineage of
tetrapods must include these ancestral,
extinct relatives.
Tetrapoda: Class Amphibia
The amphibians made first landfall during the Devonian (400 mya). The
first tetrapod had come ashore, with those defining characteristics:
- four limbs connected to the trunk
- each limb ending in five digits
There are three main Orders:
- Order Gymnophiona - The Caecilians
- Order Caudata - Salamanders & Newts
- Order Anura - Frogs & Toads
Modern Caudates (salamanders and newts) most closely resemble the
ancestral amphibians, in that they retain a tail. But all three extant
orders of amphbians are highly derived in their own ways, with each
species showing amazing specializations in both morphology and behavior.
Fun Amphibian Facts
ectothermal poikilotherms
Primitively, larvae are aquatic, with gills
Skin is scaleless, and serves a major respiratory function: must remain moist
Skin is well supplied with glands, including poison glands. Some species far more poisonous
than others.
large mouth with very small teeth
Mostly insectivorous/carnivorous (ambush predators)
Heart is three-chambered: One ventricle, and two atria. Oxygenated
blood enters via one atrium, and unoxygenated through the other: both are
mixed in the ventricle, which then pumps the blood to the rest of the
body. This means that the oxygenation system isn't perfect!
Separate sexes
- internal fertilization in caecilians and salamanders
- external fertilization in most frog species
The Amniota
includes
- Class Reptilia
- Class Mammalia
The main homology linking the amniotes together is the Amazing Amniotic
Egg
All amniotes share this character, though it is highly derived
in some groups (e.g., mammals).
There is some argument as
to the proper classification of amniotes, but they are generally
divided into three main groups on the basis of skull morphology:
- Anapsida - No temporal opening behind the eye orbits
- Synapsida - Single temporal opening behind the eye orbits
- Diapsida - Two temporal openings behind the eye orbits
Today's living remnants of these three lineages:
- Anapsids - turtles
- Synapsids - mammals
- Diapsids - lizards, snakes, crocodilians, and birds
Of greatest note here is the reorganization of what we used to call "Class
Reptilia." This has traditionally included the animals that are now known
to be Anapsids (turtles) and Diapsids (dinosaurs, birds, crocodilians,
snakes, lizards, and tuataras), so "Class" Reptilia as it is usually
constructed is polyphyletic, and is now in the process of being reorganized.
Anapsida - Turtles and their relatives
- Anapsids are characterized by an external bony shell
consisting of a dorsal CARAPACE and a
ventral PLASTRON. The shell is
fused to the vertebrae and ribs, and is an integral part of the
skeleton.
- Turtles lack teeth, but have sharp cutting edges to the maxilla (upper
jaw) and mandible (lower jaw).
- Turtles may be marine, fresh water, or terrestrial.
- Here's a nice
gallery of a few of the many turtle species.
Diapsida -
Dinosaurs, Birds, Crocodilians, Tuataras, Snakes, and Lizards
- a word about lizards (Order Squamata, Suborder Sauria)...
- a word about snakes (Order Squamata, Suborder Serpentes)...
amphisbaenids (Order Squamata, Suborder Amphisbaenia) "worm lizards" - found extensively in South America
and tropical Africa, these are burrowing legless lizards that
superficially seem to resemble earthworms due to convergence)
crocodilians (Order Crocodilia)- sister taxon to birds and dinosaurs
"Traditional" Reptile characteristics:
Birds are Reptiles?
Birds are perhaps the most derived of all Diapsids
Birds began as feathered reptiles derived from a group of small,
carnivorous dinosaurs known as therapods
The most famous of these is Archaeopteryx.
Feathers are homologous to scales/scutes of crocodilians.
Some secondarily flightless birds (ratites) have evolved where
selective pressures were appropriate.
Diagnostic Characteristics of Birds
- diapsid skull with fused bones
- upper maxilla and lower mandible fused, covered with keratinous
sheath that forms the toothless BILL (or "beak")
- saurischian pelvis (see pp 350-351 of your text!)
- long neck, compared to other reptilian descendants
- forelimbs (primitively) modified for flight; some (ratites) have
secondarily lost flight capability
- epidermal scales modified to form FEATHERS on the body. On the
legs, the scales are comparable to those of their reptilian cousins'
- ear pinna (the part that sticks out!) lost or rudimentary
- no sweat glands
- oil gland at the base of the tail
- fully ossified (bony) skeleton with air spaces in the bones.
- highly developed nervous system with complex sense organs
- tri- or tetra-chromatic vision plus oil droplets in the cone cells
- four-chambered heart
- nucleated red blood cells (unlike yours!)
- homeothermic and endothermic
- respiration by means of slightly elastic lungs connected to AIR
SACS nestled among the viscera and bones.
- NO DIAPHRAGM (unlike you!)
- Voice box near distal end of trachea, just before the bronchi of
the lungs. In perching songbirds (Order Passeriformes, commonly
called "passerines"), this is modified to form the SYRINX,
which enables these birds to vocalize complex songs.
- Ureters of renal system open into CLOACA. There is no bladder,
and urine is excreted primarily as URIC ACID.
- Internal fertilization
- Egg shell highly calcified and hard; large yolk.
- Young may be altricial or precocial
- Sex determination is via the female parent:
- male = ZZ sex chromosomes
- female = ZW sex chromosomes
- (in mammals, male = XY and female = XX sex chromosomes)
Birds are a diverse group comprised of 28 orders.
Vast diversity of form and behavior, with some of the most complex mating
behaviors known in the animal kindgom. (A word about LEKs)
Many species are MIGRATORY. (See pp 371-373 of your text)