THE EVOLUTION OF VERTEBRATES
The Vertebrate Story
Introduction
To understand the evolution of vertebrate form and function we must recognize that both external and internal factors are important.
External factors (such as predators, climate, and competition) are fighting against internal capabilities.
Internal designs are integrated into a functional individual animal and if design changes, it must do so without serious disruption of the whole organism.
The internal construction sets boundaries to allowable change in external form.
Morphological Concepts:
Homology: two or more features that share a common ancestor, i.e., a bird’s wing and a mole’s arm
Analogy: features with a similar function, i.e., the wings of bats and bees
Homoplasy: features that look alike, dolphin and turtle flippers
Each of these three concepts are separate contributors to animal design
1. Similar function in similar habitat may produce convergence of form (analogy),
2. Common ancestry can carry shared and similar structure (homology), and
3. Sometimes these collectively can result in similarity of appearances (homoplasy).
Biological Function: refers to the action or property of a part as it functions in the organism
Biological Role: refers to how the part is used in the environment during the course of the animal’s life
Biological functions (and roles):
• insulation (thermoregulation)
• aerodynamic contouring (flight)
• display during courtship (reproduction)
To determine function, we need to use physiological and anatomical studies, as well as use tools such as electromyograms, X-rays, etc.
To determine biological roles, we need to observe the animal in the real world and see how it operates ---- ecomorphology.
Preadaptation: this means that a structure or behavior has the necessary form and function before the role arises. This does not mean that the trait arises in anticipation of some future role.
What is a Chordate???
All chordates share the following four characteristics at some point in their development:
• pharyngeal gill slits - a series of openings that connect the inside of the throat to the outside of the "neck". These are often, but not always, used as gills.
• dorsal hollow nerve cord - a bundle of nerve fibers which runs down the "back". It connects the brain with the lateral muscles and other organs.
• notochord - cartilaginous rod running underneath, and supporting, the nerve cord.
• post-anal tail - an extension of the body past the anal opening
What is a Vertebrate?
It has all of the chordate characters plus:
Vertebrates are characterized by a vertebral column: that is, a variable number of endoskeletal elements aligned along the notochord (green) and flanking the spinal cord (yellow).
Origin of Vertebrates:
1. emergence of the chordate body plan
2. origin of bone
3. origin of paired sensory structures
Origin of Gnathostomes:
1. origin of jaws
2. origin of teeth
3. origin of paired fins
4. origin of swim bladder in osteichthyes
Origin of Tetrapods:
1. origin of limbs in amphibians
2. origin of the amniotic egg
Diversity of Chordates: Subphylum Vertebrata
Vertebrates may be characterized by 12 general derived characteristics. You should become very familiar with these traits.
2. Two pairs of jointed locomotor appendages, which can include fins (pectoral and anal/dorsal fins, as well as the forelimbs and hindlimbs)
3. Outer covering of protective cellular skin, which can be modified into special structures such as scales, hair, and feathers
4. Metamerism found in skeletal, muscular and nervous system. This includes ribs, vertebrae, muscles, and ganglia/peripheral nerves
5. Well-developed coelom or body cavity completely lined with epithelium (cellular tissue of mesodermal origin) that may be divided into 2 to 4 compartments
6. Well-developed internal skeleton of cartilage and/or bone, separated into axial skeleton (skull, vertebrae, ribs, sternum) and appendicular skeleton (girdles and appendages)
7. Highly developed brain enclosed by skull and nerve cord enclosed by vertebrae - these provide advanced neural structures that are highly protected from damage
8. Well-developed sense organs (eyes, ears, nostrils) located on the head (cephalization)
9. Respiratory system, including either gills or lungs, located closely to the pharynx or throat
10. Closed circulatory system with ventral heart and median dorsal artery
11. Genital and excretory systems closely related, utilizing common ducts and pathways
12. Digestive tracts with two major digestive glands
(liver and pancreas) that secrete into it
SUBPHYLUM: Vertebrata
Class: Myxini
Superclass AGNATHA
Class:
Conodontia*
Pteraspidomorphi*
Cephalospidomorphi
Superclass GNATHOSTOMATA
a) Grade Pisces
Class:
Placodermi*
Acanthodia*
Chondrichthyes
Sarcopterygii
Actinopterygii
b) Grade Tetrapoda
Class:
Amphibia
Reptilia
Aves
Mammalia
SUBPHYLUM: Urochordata (tunicates or sea squirts)
SUBPHYLUM: Cephalochordata (Branchiostoma)
SUBPHYLUM: Craniata
(Myxini plus Vertebrata)
Myxini - hagfishes
Cepahlospidomorphi - lampreys
- about 50 species still living
- 360+ million year fossil record
- 2 independent lineages
- lampreys are FW and oceanic animals
- parasitic and non-parasitic forms (latter only
live as an adult long enough to reproduce)
Chondrichthyes (chondro = cartilage + ichthyes
= fish)
2 subclasses with about 800 living species
- Elasmobranchii - sharks, skates and rays (5-7
pairs of gill arches)
- Holocephali - ratfish and chimeras
(4 pairs of gill arches, single external opening)
Elasmobranchs are subdivided into:
1. sharks (superorder Selachimorpha - 340 species)
2. skates and rays (superorder Batidoidimorpha -
425 species)
some give birth to live young, some are filter feeders
Holocephalens have a smooth epidermis, no ribs, large
eyes, upper jaw
fused to the braincase and a long flexible tail
(30 species)
- these fish are mollusk feeders and have broad
grinding teeth (ratfish)
Sarcopterygii ("lobe-finned")
2 living subclasses
Porolepimorpha and Dipnoi (lungfishes - 6 species)
Coelacanthimorpha (extinct except for the Coelacanth)
Actinopterygii ("ray-finned" fishes) (20,000+
species)
2 living subclasses
Chondrostei (sturgeons and bichirs)
Neopterygii (all teleosts)
Amphibia - amphibians = amphi + bios ("double
life")
2 subclasses
Labrynthodontia (extinct)
Lepospondylia
The latter group has about 3000 species.
3 orders
Urodela (Caudata) - salamanders
(tailed)
Salientia (Anura) - frogs/toads
(tail-less)
Gymnophiona (Apoda) - caecilians
(foot-less)
Reptilia (currently ~6,000 species)
4 subclasses
Anapsida - turtles
Parapsida - extinct aquatic dinosaurs
Synapsida - extinct dinosaurs ==> mammals
Diapsida - extinct dinosaurs, crocodiles, existing
lizards and snakes ==> birds
Aves -- feathered reptiles ???
25 orders with 8600 species
2 subclasses
Archaeornithes (extinct)
Neornithes (modern)
- at least 2 reptilian characters they retain are
their amniotic eggs and the scales on their legs
- almost every component of a bird’s anatomy is
geared for flight
- hollow, honeycombed bones (2 m wingspan frigate
bird weighs 4 oz)
- also get a reduction in some organs for example
only one ovary
- excellent vision, very adaptable beak
- modified digestive system (no teeth, gizzard instead)
- very efficient respiratory system
- large brains, complex behavior
Archaeopteryx lithographica - primitive bird with reptilian characters often considered a hoax
Chordates ---
Vertebrates ---pharyngeal gill slits dorsal hollow nerve cord notochord post-anal tail
Changes in the axial skeleton were important as animals went from a buoyant environment to one where the body was slung between the limbs
What was it like on land?
Warm and humid
Lots of vegetation:
scale trees
horsetails
Lots of invertebrates
insects
arachnids
fossorial - burrowing
cursorial - running
arboreal - in trees
aerial — flying or gliding
saltatorial - hopping
Heterodont dentition - many types of teeth
Homodont dentition - one type of tooth