Amniote Origins and Characteristics
Evolutionary Origins of Amniota
- The Transition from Amphibians: The evolutionary transition from amphibians to amniotes is characterized as being less dramatic than the preceding transition from fish to tetrapods.
- Ancestry: Most systematists conclude that the clade Amniota is derived from the anthracosaur amphibians.
- Gephyrostegus: This genus is noted because anthracosaurs like it were already very amniote-like in their general characters prior to the formal emergence of the Amniota group.
- Geographic Origin: The earliest known amniote fossils are located in the Joggins Formation of Nova Scotia.
- Geological Context:
* The Joggins Formation preserves the remains of an ancient swamp forest.
* The age of this formation is mid-Pennsylvanian, dating back approximately 300myo.
- Fossil Preservation Mechanisms:
* The formation contains many fossilized tree stumps that were preserved in situ (in their original place).
* Some of these stumps became hollow over time.
* Small animals entered these hollow stumps, died, and their remains were preserved within them.
Principal Early Amniote Taxa
- Hylonomus:
* Identified as the earliest known and one of the most primitive amniotes.
* It possessed an anapsid skull condition.
- Other Joggins Discoveries:
* The tree stumps yielded other amniote specimens, including a primitive synapsid and another anapsid similar to Hylonomus.
* Note: While these are the earliest known fossils, they are not considered the most structurally primitive amniotes.
- Carbonodraco: An early amniote found in Ohio.
Early Reptilian Lineages and Parareptiles
- Parareptiles (Stem Reptiles):
* This group evolved prior to the derivation of modern reptilian groups and is placed on the branch below the split between archosaurs and lepidosaurs.
* Pareiasaurs: A group of terrestrial species within the parareptiles.
* Example: Scutosaurus, a pareiasaur from the later Permian of eastern Europe.
* Milleretids:
* Example: Millerosaurus.
* Procolophonida:
* Example: Sclerosaurus.
- Mesosaurs:
* A primitive group of aquatic marine reptiles from the Permian age.
* Physical Characteristics: Narrow, elongate snouts with long, interlocking teeth.
* Classification: Historically considered to be part of the Parareptiles.
Distinctive Characteristics of the Amniota
- The Cleidoic Egg: A defining feature of amniotes, which contains four essential extraembryonic membranes:
1. Amnion
2. Chorion
3. Allantois
4. Yolk sac
- Amniote Integument (Skin): Differing significantly from amphibian skin, amniote skin is characterized by:
* A thicker, many-layered epidermis.
* Cornification via the deposition of abundant keratin.
* Integumentary thickenings known as scales, which are particularly dense and provide high levels of waterproofing.
* A significant reduction in the number of skin glands.
Keratin Types and Epidermal Dynamics
- Reptilian Keratins:
1. Alpha keratin: Characterized as pliable and supple; found throughout the skin and is the exclusive keratin type in the hinge regions between scales.
2. Beta keratin: Characterized as tough and hard; found abundantly in the scale areas of the reptile epidermis.
- Epidermal Growth and Shedding:
* Stratum Germinativum: This lower layer of the epidermis produces upper layers through mitosis.
* Turtles and Crocodilians: Mitosis occurs continuously at low levels (excluding hibernation periods). Skin cells are produced from below and shed from above in small patches.
* Lepidosauria (Lizards and Snakes):
* Possess two distinct cellular layers in the epidermis: an outer generation and an inner generation.
* Ecdysis: The outer generation is shed all at once. The inner generation is replaced from below following episodic periods of mitotic activity in the stratum germinativum.
Amniote Classification by Skull Morphology
- The main groups of amniotes are distinguished primarily by the arrangement of bones in the temporal region of the skull, specifically involving the postorbital and squamosal bones.
- Skull Conditions:
* Anapsid: No temporal openings (e.g., the primitive condition seen in Hylonomus).
* Emarginated Anapsid: A variation of the anapsid condition.
* Synapsid: A single temporal opening (includes early and late synapsid forms).
* Diapsid: Two temporal openings (includes early diapsids like Araeoscelis and late diapsids).
* Euryapsid: A specific skull configuration often found in certain marine reptiles.
* Modified Diapsid: Advanced configurations found in modern lizards and snakes.
Major Diapsid Lineages: Lepidosauria vs. Archosauria
- Diapsida Overview: Most animals categorized as reptiles are diapsids. They underwent an early radiation of lizard-like forms such as Araeoscelis and then split into two major lineages.
- Lineage 1: Lepidosauria
* Included Taxa: Lizards, Snakes, Amphisbaenians, and Tuataras (Sphenodon).
* Diagnostic Characters:
1. Transverse cloacal opening.
2. Periodic ecdysis (skin shed all at once).
3. Presence of autotomy planes in the tail (found at least in primitive members).
- Lineage 2: Archosauria
* Included Taxa: Crocodilians, Aves (birds), and Dinosauria.
* Diagnostic Characters:
1. Longitudinally oriented cloacal opening.
2. Continuous ecdysis (skin shed continuously in small clumps).
3. Lack of autotomy planes in the tail.