14.

1
Macroevolution: evolution occurring above the species level, including
the origination, diversification, and extinction of species over long
periods of evolutionary time
Scientists observe using fossil evidence can identify large
scale patterns in biodiversity, can reconstruct climates and
environments of ancient organisms
Can also study macroevolution by studying organisms
A. The Motors of Macroevolution: Speciation and Extinction
D1 + originations + immigrations extinctions - emigrations = D2;
D stands for diversity (total # of species in a particular clade)
Changes in diversity through time can be studied by looking at
the interplay btwn speciation and extinction
B. Rates of Origination and Extinction
Rates are calculated as changes in their numbers occurring over
some unit of time
o Ex. per million years, in geographic stages
Standing diversity: the # of species (or other taxonomic unit)
present in a particular area at a given time
o If standing diversity falls for too long and reaches 0, the
entire clade will become extinct
Alpha (): rate of origination; Omega (): rate of extinction
o If > for long time, standing diversity will ; if opposite,
then standing diversity will decrease
o *in most taxa, > but by only a little!!! Too much = will
be overrun with species of that particular taxon
turnover: the disappearance (extinction) of some species and
their replacement by others (origination) in studies of
macroevolution. The turnover rate is the # of species eliminated
and replaced per unit of time
o relatively low turnover rates (snails, clams) = their species
lased much longer and new species appeared less
frequently
o fast turnover rate: each species only lived briefly and new
species continued to arise
why diversity can decline
o an increase in extinctions
o if origination rates drop dinosaurs! Had high turnover
ratenearly 100% of dinosaur genera became extinct at
end of their particular formation and the typical origination
rate of dinsosaur genera per formation was also nearly
100%
o unusual that no new kinds of dinosaurs evolved to take
their place

extinction can happen 2 ways: can increase or can drop

drastically
o a sustained drop in origination rates or a strong increase in
extinction rates

Chapter 4: The Tree of Life

4.1 Tree Thinking
phylogeny: a visual representation of the evolutionary history of
poopulations, genes, or species represents branching pattern
of evolution over time
clade: an organism and all of its descendents
o smaller clades are nested in larger ones
tips: terminal ends of an evolutionary tree, representing species,
molecules, or populations being compared
branches: lineages evolving through timw btwn successive
speciation events
node: a point in a phylogeny where a lineage splits (a speciation
event)
internal nodes: nodes w/in a phylogeny representing ancestral
populations or species

4.2 Phylogeny and Taxonomy

monophyletic: a term used to describe a group of organisms that
form a clade
different conclusions can result from different lines of evidence,
Linnaeuss system was built on an understanding of structural
similarities that was relatively basic compared with the tools of
scientists today
4.3 Reconstructing Phylogenies
characters: heritable aspects of organisms that can be compared
across taxa

o shared character sttes that were ancestral to a group are

NOT informative for defining that group
ex. humans and cats have skulls but it doesnt help
us decide that they belong to the same clade b/c
repotiles and fish have skulls too
taxon: a group of organisms that a taxonomist judges to be a
taxonomic unit, such as a species or order
o taxa that diverged from each other recently would have
more character sttes in common than taxa that divereged
farther in the past b/c closely related taxa have a longer
shared history and have spent less time on their separate
evolutionary paths whereas taxa that split long time ago
have more time to evolve and are generally more
divergent from each other
synapomorphy: a shared derived character (i.e one that evolved
in the immediate common ancestor of a clade and was inherited
by all of its descendants)
o only these coan proide clear understanding of relationships
among organisms
o ex. production of milk evolved in the immediate common
ancestor of living mammals and is absent from the closest
relative of living mammals such as birds and iguanas
cladistics: phylogenetic methods that construct trees by grouping
taxa into nested hierarchies (clades) according to their shared
derived characters (synapmorphies)
o clade is monophyletic if it can be cut from the larger tree
w/ a single cut = if it includes an organism and all of its
descendants
homoplasy: character state similarity not due to shared descent
(e.g produced by convergent evolution or evolutionary reversal)
evolutionary reversal: the reversal of a derived characteristic
state to its ancestral state
o ex. snakes descend from 4 legged reptiles but
subsequently lost their limbs
convergent evolution: the independent origin of similar traits in
separate evolutionary lineages diff. lineages can
independently evolve the same trait
systematists identify synapmorphies to generate phylogenies.
Homoplasy can crete the mistaken impression that species are
closely related when theyre actually not

3.1 The Ancient Earth

19th century scientists debated how old the Earth is

3.2 A

Darwin concluded that the Earth formed through gradual erosion

over course of 300 million years and if it had taken hundreds of
millions of years for a relatively small geological formation to
reach its current state, Darwin surmised that the Earth must be
billions of years old
William Thomson (Lord Kelvin) criticized Darwin argued that the
world cant be that old; his argument is based on temperature
instead of formation of rocks
o Use current temperature of rocks to estimate how long
they had been cooling since hot rocks cool at a steady rate
o Estimated that Earth is only 20 million y/o but hes wrong!!
He assumed that the planet was a rigid sphere but the
panets interior is dynamic
Studies of radioactive isotopes show that the earth is 4.568
billion years old
Vast Museum
Darwin referred to earth as a vast museum b/c the fossil record is
far from complete
Most organisms dont fossilize because other organisms eat them
Largerstatte: a site with an abundant supply of unusually well
preserved fossils from the same period of time, often including
soft tissues
o In most cases, the fossilized animals were swept into
anoxic, lifeless pools, lagoons, or bays where microbes and
other scavengers couldnt destroy their bodies instead,
they were trapped in fine sediment which preserved their
tissues in stone
o Ex. Burgess Shale in British Columbia has yelied 65k+
speciments of mostly soft bodied animals representing at
least 93 species
o Important also because it acts like a snapshot of an entire
ecosystem has has long since vanished
Importance of fossils
o Determine how extinct organisms developed over time
(morphological changes such as growth)
o Record the decline of organisms, showing marks left on
them by illnesses/injuries
Biomarkers: molecular evidence of life in the fossil record; can
include fragments of DNA, molecules (ex. lipids), isotopic ratios
o Some re large molecules thar are the produce of a long
series of anzymatic reactions
o Some are so distinctive that geochemists can even identify
the group of species that produced them
o Example: Okenone in Australia

Today, the only known source of okenane is a

biological molecule called okenone and the only
organisms that produce okenone are purple sulfur
bacteria ancient Australian rocks contain high
levels of okenane reveals tht purple sulfur bacteria
were present in earth 1.64 billion years ago
Today, purple sulfur bacteria are rare and found in
extreme environments with low oxygen and high
levels of sulfur the abundance 1.64 bil y/a
supports hypothesis that the oceans at that time
were toxic to organisms

Isotopes
o Ratio of different oxygen isotopes in whale fossil teeth
indicates whether they consumed freshwater, seawater, or
water with an intermediate level of salinity
o Provide clue to organisms metabolism
Different plants have different ratios of carbon
isotopes depending on how they carry out
photosynthesis
o Ratio of carbon isotopes in animals reflect kinds of
animals/plants they eat
Technology allows scientists to gain new insights into the
behavior and appearance of extinct species from their fossils

3.3 Lifes Earliest Marks

Molten crust hardened lighter formations of rock rose to form
continents gases escaped from rocks to form atmosphere
water arrived on surface of planet, possibly escaping from
Earths rocks as vapor or delivered by comets/asteroids basins
btwn continents filled with water, forming oceans
The moon formed from earth colliding with debris
Burial of earths crust and collisions in its history have destroyed
almost all of the planets original surface
o Crust of earth broke into plates, hot rock rose up in some
cracks between plates and added to their margins while
opposite margins of the plates were drivn down under the
crust; as the rock sank, it become hotter until it melted
aaway
Before life began, the only source of carbon on earths surface
would be ceom lifeless sources (like volcanoes) but once life
emerged, it would have produced abundant amunts of organic
carbon which would be incoroporated into sedimentary rocks
o Rocks formed after origin of life record this shift earliest
sign of life most likely produced by photosynthetic bacteria
found in Greenland

o Research by Rosing and Frei

Case study by Abigail Allwood possibly discovered stromalites
in some of the oldest geological formations on earth
o Stromatolites: layered structures formed by the
mineralization of bacteria
Potential sighs of life date back as far as 3.7 billion years ago.
The oldest known fossils that are generally accepted are 1.45
biollion years old. The earliest signs of life are microbial, and
microbes still constitute most of the worlds biomass and genetic
diversity

3.4 The Rise of Life

living things can be divided into 3 domains!
Bacteria: one of two prokaryote domains of life; includes
organisms such as E.Coli and other familiar microbes
o Single celled organisms, may be shaped like rods,
filaments, spheres
o Exist in a bunch of niches as predators on other bacteria,
as photosynthesizers, as heterotrophs, chemoautotrophs
o Have membrane that contains peptidoglycan and a unique
set of 5 proteins that carry out RNA polymerization (this is
unique to bacteria only)
Archaea: one of the two prokaryote domains of life; resemble
bacteria, but are distinguished by a number of unique
biochemical features
o Single celled organisms, may be shaped like rods,
filaments, spheres
o Live in wide range of habitats, not capable of
photosynthesis
o Found in fossil record 3.5 billion years ago

Eukarya: a domain of life characterized by unique traits including

membrane-enclosed cell nuclei and mitochondria; includes
animals, plants, fungi, and protists (single-celled eukaryotes)
o Eukaryotes are roughly 100 times bigger than bacteria and
archaea and they have a nucleus
o Emerge in the fossil record 1.8 billion years ago