Exam 1 on Friday, August 16!

Lectures 1 to 5
(through Adaptation & Functional Morphology)

What are fossils good for?

Macroevolution: we can examine patterns
and hypothesize mechanisms of
evolutionary change over long periods of time

Study guide and sample questions are posted

Around 30 questions
(multiple choices + short answers)
Bring UCD Scantron 2000, a pencil and an eraser

Can studies of population genetics alone

explain patterns that we see in the fossil
record?

Unlikely!

Dont be late!

Long-term evolutionary patterns that are

known from the fossil record, and are not
predictable from the living world alone:

Species longevity

Long-term evolutionary patterns that are

known from the fossil record, but are not
predictable from the living world alone:

Taxonomic turnover

Species can exist for long and

variable periods of time (from
1-25 million years, on average)

Higher taxa (genera, families,

orders, etc.) turn over (change) at
a high rate over geological time

Long-term evolutionary patterns that are

known from the fossil record, but are not
predictable from the living world alone:

Modes of preservation
1. Unaltered
2. Altered

Mass extinctions
Episodes of elevated rates of
extinction have occurred, more
than once. Are they periodic?

Permineralization
Recrystallization
Dissolution & Replacement
Carbonization

Most attributes of most organisms dont fossilize

Biological Characteristics
that make it more, or less, likely to be preserved
in the fossil record:

Fossil record is made up (largely)

of mineralized skeletons
Having a mineralized skeleton greatly improves an
organisms (slim) chances of being preserved in
the fossil record

Why do organisms make skeletons?

Anatomical
What are skeletons made of?

Ecological
How do organisms make skeletons?

Biogeographical

What are organisms made of?

Most skeletal materials have two primary components:

What can their skeletons tell us about their mode of life,

and the environment in which they live?

Uniformitariansim
The present is the key to the past

1. Organic
Proteins
Polysaccharides
2. Inorganic
Mostly crystalline (atoms arranged in a highly
ordered, repeating pattern)
Formed both intracellularly and extracellularly
Most enclose or are enclosed by an organic matrix
Composed of major elements, minor elements,

Can observe these sedimentary processes acting today, thus

can infer that the same processes operated in the past

trace elements, and isotopes of elements

Principle of Superposition

Principle of Original Horizontality

Principle of lateral continuity

Most attributes of most organisms dont fossilize

Geological Characteristics
that make an organism more, or less,
likely to be preserved and discovered in the fossil record:

Depositional environment
Geological history
Time
Discovery

Relative Age vs. Absolute Age

Ways of telling time from rocks and fossils:

Relative methods:

Relative
order of
events

Numerical
ages of
events

Pattern
Rocks

Fossils

Process

Superposition Uniformitarianism
Steno 1669

Hutton late 1700s

present is key to past

Faunal
succession

Natural
selection

Smith early 1800s

Darwin 1859

Absolute methods of telling time:

Rocks are made of minerals
Minerals are made of elements or compounds
Elements are made of atoms
Some elements are radioactive (radiogenic)
Radioactive elements decay
continuously, spontaneously, and
at a predictable rate (measured in a laboratory)
We can measure the ratio of parent to daughter
atoms to determine the age of a rock that
contains minerals containing radioactive
elements

Stable isotopes of oxygen

16O
18O

(~99%)
(~1%)

The ratio of 18O to 16O measured in natural waters or skeletons

temperature (and salinity) dependent
The ratio varies as the environmental temperature varies
16O

has less mass than 18O (because it has fewer

neutrons), so it is more mobile in the environment

DEATH

Biostratinomy: decay, disarticulation, deposition

Biological

Chemical

Physical

bacterial decay
borers
scavengers

dissolution
mummification

abrasion
borings
disarticulation
transport

BURIAL

Diagenesis: fossilization (sensu lato)

bioturbation

oxygen

isotopes can tell us about the temperature of

formation of skeletons of marine organisms; seasonal
growth periodicities; etc.

recrystallization
permineralization
carbonization

replacement

DISCOVERY

Organisms are rarely

preserved and found as
fossils
BUT they are sometimes
preserved excellently in
certain localities

Types of growth
All organisms with a mineralized skeleton
must enlarge the skeleton to accommodate the
growth of soft tissues

Addition
Change

Lagersttten (mother loads)

sites of exceptional preservation
Art Rogers

Variation in organisms

Quantify

size
and

Shape
=
Morphometrics

Growth of organisms commonly

involves allometry

Allometry
:growth of a part of an organism in relation to

Change in shape that accompanies

a change in size

the growth of the whole organism

Isometric growth: relative size of morphological
characters does not change with absolute size

Often results from competing

requirements of:
surface area (diffusion, heat loss, friction)
cross-sectional area (muscle force, tension)
volume (buoyancy, body mass)

Allometric growth: relative size of morphological

characters changes with absolute size following a
power function
Y = b Xa

Processes that can shift gene frequencies:

Processes that can shift gene frequencies:

Natural selection

Genetic drift chance fluctuation in gene frequency

variation exists among individuals

greater effect in small populations

the variation is heritable

Inbreeding reduces genetic variability within populations

differential survival occurs

Migration mixing prevents genetic divergence
Will increase those gene frequencies that are more fit (more

Mutation occur spontaneously

ultimate source of genetic variation

adaptive) in some environments

can be beneficial, neutral or lethal depending on

Most influential force in shifting gene frequencies

environment

in large populations

Biological Species Concept (BSC)

Ernst Mayr, 1942

Groups of actually or potentially interbreeding

natural populations,
which are reproductively isolated
from other such groups
Defined by interbreeding behavior
reflects underlying genetic cohesion
Species are still recognized morphologically, in practice

Rationale:
Similarity in morphology represents underlying genetic cohesion

Allopatric speciation
Geographic isolation

Parapatric speciation
Partial geographic isolation

Sympatric speciation
No geographic isolation

Species evolve as populations diverge via

Phylogenetic systematics

(Hennig, 1966)

cladogenesis or shift mode via anagenesis

and become reproductively isolated
(and usually morphologically distinct)

The methods of systematics provide a way to organize the

extraordinary diversity of life, and make sense of it in an
evolutionary framework

from one another

Systematics identifies evolutionary patterns
generated by the process of common descent
Systematics is the only means by which
macroevolutionary
patterns can be identified and studied

Phylogenetic patterns generated from branching processes

of cladogenesis may be represented in two different ways:

Characters
Phylogenetic hypotheses (cladograms or evolutionary trees)

1. Cladogram: maps the distribution of characters among

taxa

are generated from characters (features that are heritable)

Clades = groups that include all the descendants from a single

common ancestor are monophyletic groups imply a relative
pattern of ancestry (recent, distant)

Polarity = direction of character transformation in

Sister groups = two groups that share most recent common

ancestor
Paraphyletic groups = include some, but not all descendants from
a single more recent common ancestor

evolution
Primitive (plesiomorphic) = ancestral, more general
Derived (apomorphic) = more specific

Polyphyletic groups = include some, but not all descendants from

a single more distant common ancestor

Methods of phylogeny reconstruction

Characters

How are hypotheses of phylogenetic relationship constructed?

Phylogenetic hypotheses (cladograms or evolutionary trees)

are generated from characters (features that are heritable)

1. Choose the groups of organisms whose phylogeny you are

interested in

Homology = features that share common ancestry

2. Select the characters > homologues (not analogues)

are often, but not always, similar to one another

are informative about evolutionary relationships
Analogy (homoplasy) = features that are similar, but NOT
because of common ancestry
Convergence; parallelism; reversal

3. Select a point of reference > polarity determination (primitive;

derived)
Outgroups (sister group)
Stratigraphy (lowest); Ontogeny (youngest)
4. Criterion for evaluating alternatives > parsimony; maximum
likelihood
Given the data in hand, the simplest explanation to account for
those data is preferred over more complex explanations;
the statistically most likely explanation is preferred.

Morphological phylogenetic analysis

Advantages of molecular sequence data:
Advantages:
is tangible
relatively easy to observe
more tests available for homology hypotheses
can use it to relate the organism to the environment
(functional morphology, paleoecology, etc.)

Abundant characters
Only four character states possible
Relatively easy to analyze computationally (now)

Disadvantages of molecular sequence data:

Disadvantages:
defining characters and character states is difficult
relatively small number of characters can be specified
we dont yet know much about rates of morphological
evolution, and variation among those rates

Difficult to relate to the environment directly

Difficult to assess homology of nucleotide similarity
Assume clock-like rates of change
Not realistic in all cases
Because so few character states, homoplasy likely

Adaptation and Natural Selection

Why should we study adaptation?

What is adaptation?
Adaptation concerns the
fit of an organism to its environment;
the appropriateness of design of an organism for the
environment (physical, chemical, biological)
in which it lives
How does adaptation work?
(Most) adaptations arise through
the process of natural selection

Scaling Effects
Interpret allometry to be a result
(effect) of size-dependent
scaling in certain
physical/chemical parameters

1. To understand how organisms live today in their

natural environment
2. To reconstruct functional morphology of organisms
(living and extinct)
3. To evaluate theories about the long-term mechanisms
of evolution

Larger animals

Smaller

Heat loss
Heat generation

easier to keep constant body temperature

in low ambient temp.
easier to be overheated in high ambient
temp.

Gigantothermy

Metabolic Rate per 1kg of Body Mass

The instructions for

generating the structure
(molecular sequences
in the genome)

historical factor
PHYLOGENY

Thomas (1978)

What a structure is
made of (minerals, soft
tissues) and how it is
built (growth and
development)

ADAPTATION
functional factor

GROWTH
morphogenetic factor

How a structure is used (feeding,

moving, reproducing, etc.)

Structure

Function

Structure
Structure
Structure
Structure

Function
Function
Function
Function

Morphospace
Simplified depiction of complex
shape variations using selected
variables (measurements)