SCHOOL OF SCIENCE

ZOOL 322: PRINCIPLES OF VERTEBRATE ZOOLOGY

Course leader: Dr. Emily J. C.

Senior Lecture: Zoology
Department of Biological Sciences
University of Eldoret

Email: emilychemoiwa@yahoo.com
Mobile: 0722 931 952
Office D124
ZOOL 322: PRINCIPLES OF VERTEBRATE ZOOLOGY COURSE
DESCRIPTIONS
 Characterization of Mammalogy and Mammalian
Traits
 The evolutionary history of various mammalian
lineages

ZOOL 121 2019/2020

 Characterization of the orders and families of the extanct
mammals
 Morphological adaptations of mammals for feeding,

@DREJC
locomotion, reproduction, etc
 Physiological adaptation of mammals for homeostasis
and reproduction
 Behavioural adaptations of mammals for feeding,
homeostasis, reproduction
 Social Organization of mammals
  Early evolution of mammals
 The evolution of mammals has passed through many
stages since the first appearance of their synapsid

03/12/2023
ancestors in the Pennsylvanian sub-period of the late
Carboniferous period. ... The synapsid lineage became
distinct from the sauropsid lineage in the late

@Dr. EJC 2017

Carboniferous period, between 320 and 315 million
years ago.

READ ABOUT THE GEOLOGIC TIME

SCALE
ORIGINS OF A GEOLOGIC TIME SCALE-
USE THE LIST OF REFERENCES AND 3

THE PDF PROVIDED

03/12/2023 @Dr. EJC 2017
4
I don’t expect you to know this, but knowing
the order of the geological periods can help
you make sense of what we’ll be discussing.

What helped me was this little mnemonic.

Come Over Some Day, Might Play Poker. Three

Jacks Covers Two Queens.

~360 MYA

See http://geology.com/time.htm
 3 Living Groups of Mammals
Monotremes Metatherians

Eutherians
Monotremes Metatherians (Marsupials) Eutherians (Placentals)
Mo no tremes Metath erian s (Marsu p ials) Eu th erian s (Placen tals)

Depth represents relative time. Node - Divergence Event

Branch - Common Ancestor
 Amphibians

Mammals

Squamates
(Lizards & Snakes)

Synapsids Turtles

Crocodylians
Tetrapod Phylogeny

Amnion
Stem Amniotes

Dinosaur I
Evolution of Limbs

Birds

Dinosaur II
 Temporal Fenestrae

Anapsid Synapsid
Temporal fenestra
Naris Orbit Naris Orbit

Postorbital Postorbital
Squamosal Squamosal
 Synapsid Phylogeny

Ear ly The ra p s id s
"P e lyc o s a u r s "
Cy no d o n t s

Th e r ap s id s

~ 323 Ma
 “Pelycosaurs”
(Early synapsids)
Carboniferous (~323 MYA) and
persisted through Permian.

Some had a large dorsal sail

(thermoregulatory? Mate choice?)

Rather large (~ 3 meters)

Range of Ancestral Characters

Dimetrodon Weakly heterodont

Small temporal fenestra

Angular/articular in mandible

Quadrate/articular jaw joint

Two nares - no secondary palate

Single occipital condyle

 Synapsid Phylogeny

"P e lyc o s a u r s " Ear ly The ra p s id s

Cy no d o n t s

Th e r ap s id s Middle Permian (~270

Ma)
 Early Therapsids
Middle Permian (ca. 270 MYA)

Active and diverse (4 major lineages) Lycaenops

Dominant terrestrial life form* (significant

later)

Most went extinct during Permo-Triassic

extinction event

Mixture of Ancestral vs. Derived

Characters
Enlarged temporal fenestra

Deeply thecodont teeth

Partial, gradually evolving secondary palate

Sweeping changes to skull and jaw structure in

one lineage.
 Synapsid Phylogeny

Permo-Triassic Mass Extinction

Ear ly T h e ra p s id s
"P e lyc o s a u r s "
Cy n o d o n t s

Th e r ap s id s
 Cynodonts*: Advanced Theraspids
(*’dog teeth’)

Cynognathus

Evolution of mammalian characters

•Many transitional fossils

•Complete secondary palate
•Two occipital condyles
•Gradual enlargement of dentary /
shrinking of post-dentary bones
•Vast expansion of temporal fenestra
•Strongly heterodont dentition
 Cynodonts*: Advanced Theraspids
(*’dog teeth’)

Cynognathus

Evolution of mammalian characters • Very late Permian & survived

•Many transitional fossils
•Complete secondary palate
•Two occipital condyles
•Gradual enlargement of dentary /
shrinking of post-dentary bones
•Vast expansion of temporal fenestra.
•Strongly heterodont dentition
the P-T extinction
• Direct interaction with dinosaurs
• By late Triassic, they were small
and inconspicuous
• Extinction of dinosaurs (end of
Cretaceous) lead to radiation
SOME BROAD QUESTIONS IN MAMMALIAN
EVOLUTION
•What are the key cynodont groups, and how are they related?

•Which of the cynodont groups are ‘mammals’?

•Why and how did mammalian characters evolve?

 Probainognathus +

Tritylodonts+

Sinocodon+

Morganucodonts+ Monotremes

Docodonts+

Early Cynodonts
Haramiyids+

Triconodonts +

Multituberculates +
Simplified Cynodont Phylogeny (Following Huttenlocker et al. 2018)

Metatherians

Eutherians
 The Key-character
Approach.
Which bones comprise the jaw joint?

Dentary and Squamosal  Mammal

Quadrate and Articular  Non-mammalian cynodont

 Probainognathus +

Tritylodonts+

Sinocodon+

Morganucodonts+ Monotremes

Q-A
Docodonts+

Early Cynodonts
Haramiyids+

Triconodonts +

Multituberculates +

D-S

Metatherians
The Key-character Approach.

Eutherians
 Fossils with both jaw joints!
Probainognathus - Middle Triassic

D/S Jaw Joint

Q/A Jaw Joint

Image from http://www.palaeos.com/Vertebrates/Units/Unit420/420.300.html

Ventral View

D/S Joint

Q/A Joint
Diarthrognathus –Another late cynodont with both jaw joints.

Clearly, the key-character approach isn’t applicable.

 Shift to a ‘Suite-of-Characters’ approach…
(Feldhammer et al.)

1) D-S jaw joint

2) Strongly heterodont dentition

3) Molar surfaces complex, with wear facets. --Occlusion--

4) Alternate side chewing, implying complex jaw musculature

5) Well-developed inner ear region.

6) Small

7) Axial skeletal characters - dorso-ventral flexion, placement of ribs, etc.

 Probainognathus +

Tritylodonts+

Sinocodon+

Not a mammal
Morganucodonts+ Monotremes

Docodonts+

Early Cynodonts
Haramiyids+
Triconodonts +

Multituberculates +

Mammal
The Suite-of-characters Approach.

Metatherians

Eutherians
 Both approaches (‘Key character’, ‘Suite of Characters’) are
referred to as ‘Grade-based’ definitions.
Problems:

•Evolution is a continuum (many transitional fossils)

•Traits may evolve at multiple locations on a phylogeny

So, ideally, what makes for a useful and appropriate

classification?

1) Classifications should reflect evolutionary history.

2) Classifications should be stable.

3) Where these conflict, priority goes to evolutionary history.

 Reptilia

Crocodylians
Amphibians

Dinosaur II
Squamates

Dinosaur I
Mammals

Turtles

Birds
Archosauria

Reptilia - a grade-based definition Archosuaria – Clade-based group

1. Scales 4-Chambered heart
2. Lack of feathers Parental Care
3. Lack of hair Vocal Communication
 Probainognathus +

Tritylodonts+

Sinocodon+

Morganucodonts+ Monotremes

Docodonts+

Early Cynodonts
Haramiyids+
Triconodonts +

Multituberculates +

Ruta et al. (2013).

Most stable definition:
Clade-based definitions of Mammalia

Metatherians

Eutherians
Rowe (1988).
Crown-group definition:
 Implications of Endothermy
A. Energy Requirements – Endotherm requires 10X energy as a similar sized
ectotherm.
Therefore, selection favored

Efficiency in food processing

•Dentition (specialized, precise)
•Evolution of masseter
•Formation of secondary palate

Cardiopulmonary efficiency
•Extrusion of nuclei from red blood cells
•Separation of oxygenated/deoxygenated blood
•Muscular diaphragm
•Thoracic ribs
•Respiratory turbinates
 Implications of Endothermy
B. Behavioral Implication – Because endotherms can generate own heat, they can be
active at cold temperatures.

Endothermy permitted nocturnality.

Selection favored:

i. Hair for insulation

ii. Development of olfactory and auditory capabilities

The evolution of endothermy generated the selective forces that favored most of the
traits we consider to be mammalian traits.
 Classic Idea.

Extinction of dinosaurs at the end of the Cretaceous permitted the radiation of

mammals, resulting in modern mammalian diversity.

Lots of current studies are testing this notion by estimating the timing of mammalian
radiation (e.g., O’leary et al., 2013 vs. Springer et al., 2013).
 TRENDS IN ORGANIZATION FROM "MAMMAL-
LIKE REPTILES" TO MAMMALS
 Enlargement of the temporal fossa

03/12/2023
 movement of jaw muscle origins from temporal region to the
braincase and the zygomatic arch (masseters)

@Dr. EJC 2017

 enabled "rotary" jaw action
 Gradual increase in size of dentary at expense of other jaw bones

 trend to dentary-squamosal jaw articulation

 reptiles and primitive synapsids have an articular-quadrate jaw

articulation
 evolution of the mammalian jaw from Steven Carr, Memorial University of
Newfoundland
 some cynodonts had both, i.e. double articulation
 quadrate-articular AND squamosal-dentary
 eventual loss of surangular then depression in squamosal
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(glenoid fossa)
 eventual transformation of articular and quadrate
 Evolution of middle ear
 stapes present in most tetrapods
 quadrate of skull becomes incus

03/12/2023
 articular of lower jaw becomes the malleus
 developmental evidence: malleus formed from proximal end

@Dr. EJC 2017

of Meckel's cartilage
 angular of lower jaw becomes the tympanic
 development of a bony secondary palate change from
homodont to heterodont dentition
 change from polyphyodont to diphyodont dentition
 change from single to double occipital condyles
 evolution of mammalian posture
 loss of cervical and lumbar ribs
 evolution of endothermy 33