INTRO TO CLASS

Biodiversity = measure of diversity between living things

Measured on 3 scales:
-genetic = diversity within species (ex: between individuals); diversity at the fine
level
-ecosystem = broadest scale; a measure of the number of ecosystem types on a
continent
- species = most common; a measure of the number of different species in a defined
area

extinction
-fossil record shows that 99% of all spcies that have ever lived are extinct
-extinction rate varies for different organisms (ie mammals on average last 1M
years, herbivores last longer than carnivores)
-extinction is the ultimate fate
-there are timesin the fossil record where there are spikes in extinction, known as
mass extinctions (a large population of the earths species were wiped out)
-65M years ago was the last mass extinction where a large comet hit the earth. The
impact was vaporization of a large part of the earth. The material got ejected into
the atmosphere and came back down as ash. There were tsunamis, earthquakes, etc
as a result. The impact also created a dust cloud, reducing the amoint of light and
heat on the earth. This lack of sunlight prevented plants from growing, causing
herbivores to die, which caused carnivores to die. This resulted in animal
extinctions. This impact event wiped out the dinosaurs

fossils
-fossils form in sedimentary rock where bodies of organisms get trapped in mud
-age of fossils in relatively known based on order (most recent fossils on top while
older fossils are at the bottom)

biodiversity crisis of today

-current bird and mammal extinction rate is 100-1000X higher when compared to
organisms of similar kind from the past. This extinction rate is comparable to the
dinosaur extinction rate
-if this continues, this will be indistinguishable from the extinction event 65M years
ago

conservation biology
-main goal is to preserve biology in all its forms
-this field draws from other biology fields

American Burying Beetle: An endangered species (extinct in Canada)

- conservation biologist are trying to save this beetle from extinction
 - these beetles do important work and have an important role in their
ecosystem
- beetles are important decomposers
- beetles release nutrients that other organisms, like plants, can use

Passenger Pigeons
-extinct in Ontario
-extinct globally
- used to be common
-were an important food source for beetles so this was a factor for the beetles
-easy prey for hunting
-overhunting lead to extinction

Endophytes (prof studies this)

-microorgnaisms living entirely inside plant tissue without causing disease or
symptoms (ie fungi)
-most are undescribed
-represent a huge pool of most unexplored biodiversity
-some are mutualistic
-benefits for plants:
> chemical defenses made by the fungus that protect the host from herbivores
> increased resistance to drought and heat
- benefits for humans:
>taxol = an anti-cancer drug discovered in Pacific yew trees which is produced by an
endiphytic fungus that lives in the tree

LECTURE 1: PHYLOGENY 1

Phylogeny is the study of evolutionary relationships among organisms

How many organisms are related to you?

- All of the animals on the slide are related to us
- Everything that is alive on earth today is descended from the same ancestor!

The Universal tree of Life

- Shows relationships of all living things today
- The tree splits into 3 major branches (archae and bacteria are unicellular,
eukaryote which are multicellular)
- Today there are at lest 10M species on earth and all of that diversity came
from the ancestor that lived 3.5-4.5 billion years ago

Phylogeny
- Basically reconstructing the family tree of species
- Important for the use of decision making in conservation biology
- Several sources of information used to reconstruct phylogeny (listed on
slide)
 - Systematics is a branch of biology that studies resemblance based on 2
factors:
o Morphological
o Biochemical
- Molecular systematics uses DNA sequences to compare DNA base pair for
base pair to see how similar or different genes are to infer relatedness
o Ex: tulips, human, fungi are physically different but the human and
fungi have similar molecular factors

Silversword alliciance
- DNA studies have shown that very different looking plants are actually
closely related in terms of their genetics
- There are mutations in key developmental genes dictates how the plant looks
- When species can interbred and hybridize, this indicates that they share a
recent common ancestor because they have a lot of genetic similarity

Characters and Traits

- If two different species share a common trait, there are different reasons for
this
o Homology (similarity due to shared ancestor)
o Analogy (similarity due to shared function ie organ does same
function that were independently acquired for survival)
Shared trait not seen in the most recent shared ancestor
- All terrestrial vertebrates share the same bones but they are different in
shape and size and used for different functions. They have all been inherited
from a similar ancestor
- The whale and ichthyosaurs most recent common reptile did not have a
flattened tail so the tails must have evolved independtly after the two split off
from their common ancestor so this is an example homoplasy

Distinugishing homolog from analogy

- Important to reconstruct phylogeny
- Ex: chimp and human skull unlikely to hve independent origin because the
structures share features that are not essential to function

Evaluating molecular homologies

- 5/11 bases are the same not enough similarity to have a common ancestor
- differences can be explained just by two mutations
- computer programs can analyze and detect shifts of DNA to find sequences
that match even if they are offset by a few bases this distinguishes
coincidence from true homologies

Phylogenetic trees
- branch points near the top of the tree indicate a more recent divergence
- taxon = any classified group (species, family, etc) plural is taxa
o example drawn by prof
 letters represent diferent living taxa

Cladograms
- doesnt contain any information other than the order of splitting and how
recent their ancestors are
- doesnt tell us the amount of change that has taken place since splitting
- altnerative to shared ancestral character is a shared derived character
(happened after the group has split off from its ancestor; doesnt need to be
present in all members of the group but can only be found in some as long as
it is not found in another group)
- ex of cladogram fossil dinos and birds on slides
- lowest node on the tree indicates a very early evolution (meaning it occurred
in the distant past)

LECTURE 2: PHYLOGENY II

Recall: the diagram is basically showing the relationship between dinos and modern
birds

Character table example (on separate piece of paper)

- D has no shared derived characters so it is the outgroup
- Cant determine evolution order of 1 and 3 so you can list them how you want
- Important to have te outgroup because we need it to see about the derived
state

Why do we need an outgroup?

- using a monotreme in this example because its a mammal but not a placental
mammal however because it is still a mammal you can use it as an outgroup
- can also use a marsupial but the fossil record isnt clear if marsupials evolved
before placental mamalls
- basically with the montreme outgroup, we can see that hair is the loss of a
character and not the acquisition so the absence of hair is the shared derived
condition

Cladograms and Timing

- cladograms dont tell us how much tme has passed, just the order of which
divergence accured

Phylograms
- shows some more information than a cladogram; shws the amount of genetic
changed between species
- in the picture, we are looking at the changes in a homologous gene between
all the species

Ultrametric Trees
- tell us when the last common ancestor between two species lived
 - basically shows when each species diverged from a common ancestor

Methods for Building Trees

- maximum parsimony
o each tree is treated like a hypothesis and the one with the fewest
mutations is the correct answer
o chose characters that are homologous
o avoid homoplasies (characters that are the same due to similar
environmental pressure)
o the example with amniotes is an example of why not to use
homoplasies to make trees
4 chambered heart is homoplasy
since we used a homplasy, the cladogram isnt parsimounous
- Autamoprohies
o Doesnt provide any useful information in parsimony analysis
o In the example, the cladogram would look the same even without the
knowledge of the hair in the leopard. Hair is not a useful information

LECTURE 3: PHYLOGENY III; SPECIES CONCEPT

Recall: parsimony (trees requiring few mutation events are more likely to be correct
so therefore they would be parsimonious)

Methods for Building Trees (cont)

- distance method
o homologous DNA should accumulate mutations at more or less the
same rate (especially if the mutations arent affecting survival)
o neutral variation = variation that doesnt affect ability to survive or
reproduce (ie: muations in intergeneic DNA, introns, silent mutations,
mutations that subsititue amino acids without changing the function
of a polypeptide)
o when comparing homologous DNA, the genetic distance is the
different in the number of the base pairs
o genetic distance should be proportional to the time passed from
divergence
o this information can be used to construct a phylogeny
o allows for the use of autapomorphies
o some genetic changes are weighted differently because thy dont
occur with equal frequency (a rare event would be weighted more
heavily)
ex: transition subsititution (ie purine for purine) are more
common than transversion substitution (ie a purine for a
pyrimidine) therefore transversion are more heavily wegithed
o if a mutation occurs but another mutation reverses it, this still gets
factored in
- combining methods
 o both methods are compared

Molecular Clocks
- used to see the amount of time that has passed since divergence
- different genes evolve at different rates and these rate reflect te importance
of these genes for fitness and survival
o ex: RNA genes evolve slowly so mutation accumulate slowly (these
genes used to identify divergences from a long time ago)
o ex: mitochondrial DNA evolves fast (used to identify recent
divergenes)
- time example: simultaneous divergence occurs in the two lineages so 2
happen in one and 2 happen in the other (making a total of 4 seen in the
sequence) but since only 2 occur in each lineage, the time is 20000 years
- the clock doesnt always run smoothly there will be irregularity where
some changes are favoured and others are not

Phylogeny in Conservation
- conservation goal: protect as much biodiversity as we can
- when trying to save and protect species in hotspots, the species are more
closely related so the genetic diversity in these regions may be limited
- evolution can only happen if there is a pool of diversity for natural selection
to act upon so if there is little genetic diversity, there is little material to act
upon
- in picture with r1,r2,r3:
o W is the weight that each species is assigned based on uniqueness
(how closely related it is to others)
o T stands for the total weights added up
o R stands for region
o P stands for percent
o So the totals as percents show us how much diversity is in each region
(the higher percentage, the more priority to conserve because there is
more diversity)
o P2 evaluates how much weight is placed on regions with animals not
in the highest priority region (which is region 3)
To show where diversity is more that is different than region 3
In this example, R1 is second priority because it has the second
high genetic diversity compared to r3

Phlyogeny and Classification

- taxonomy = science of classigiying and naming living things
- you want monophyletic taxa

What is a Species
- common definition: biological species concept (a species is a group of
individuals that has the potential to breed together in the natural
environment)
 - gene flow = mixing of alleles between different populations of species
- gene flow doesnt occur between species ecause of isolation mechanisms that
act as barriers to prevent interbreeding

Prezygotic Isolation Mechanisms

- before fertilization
Temporal Isolation
- breed at different times
Habitat isolation
- different habitats
Behavioural Isolation
- species specific matings songs, rituals, chemical signals
Mechanical Isolation
- anatomical incompatibility
Gametic Isolation
- mating occurs successfully but no fertilization (sperm die before reaching
ova or cant fuse with ova)

LECTURE 4: WHAT IS A SPECIES? (CONTD)

Postzygotic Isolation Mechanisms

- act after fertilization
Reduced hybrid viability
- hybrids die before reaching maturity (assures mixing of alleles is stopped
because the hybrid doesnt survive)
Reduced hybrid fertility
- hybrids are sterile (they cant pass their alleles)
Hybrid breakdown
- the second generation of hybrids is frail/sterile (Stops the mixing at this
stage)

sometimes more than one mechanism is working to maintain integrity of the species
(prevents from mixing)

isolation via reduced hybrid fertility isnt always 100% (ex: lions and tigers). It only
turns down the gene flow but doesnt stop it completely

this shows the criteria of breeding isnt always straightforward. In theory, it works
but in the real world it can get tricky

biological species concept focuses mainly on biological isolation but sometimes we

dont have the information about breeding bhevaiour of a species

Alternative: Morpholigical species concept

- species defined by physical/ biochemical traits
- ex: number of surface teeth in flies
 - can be used to define any spcies (unlike the biological species concept)
- problem: subjective as to what characteristics to use

Alternative: Ecological species concept

- describes species in terms of role they play in nature or what ecological niche
they are part of

Alternative: phylogenetic species concept

- defines species based on their unique genetic history
- species need to be monophyletic groups
- detailed phylogenies unavailable
- depends on what genes you use and how many genes you use to make
different species

Application of Species Concepts

1. Marine Copepods
- used to be considered as a single species based on their physical appearance
- recently, Carol Lee looked at individuals from different locations and she did
a phylogentic analysis on them to see if they would form one single branch
tip
- she found there were many different branches
- she did mating tests to see if indiviudlas from different branches could
interbreed and she found that they couldnt
- so the phylogentic and biological concepts support that there are actually 8
different species
- these species are cryptic species (indistinguishable based on morphology)

2. Elephants
- there are two species (African and Asian) and are clearly distinguishable
physically
- African elephant was traditionally regarded as one species even though it
was apparent not all are the same (there were different ones)
- These two African elephants are more similar to each other but there are
some differences but they were just considered to be one species
- They live in different habitats so they dont interbreed in nature so they
should be classified as different species (this proposal is supported by
phylogenetic analysis)
- This means that there are two habitats to try to conserve, not just one
habitat, in order to preserve the most elephant diversity

Speciation
- how we go from one population into two new populations that cant
interbreed anymore
- allopatric: population divided by a geographical barrier. After they become
isolated, they start to slowly change genetically and they become so different
that even if they come back into contact they wont breed
 - sympatric: speciation without a geographical barrier

Allopatric
- barrier blocks gene flow so there is no exchange of alleles
- allele frequencies slowly diverge
- isolation mehcnisms evolve
- sometimes there isnt enough genetic difference to make another species

Geographic Isolation
- two types:
o dispersal (some individuals cross over some sort of a barrier and now
there are two separate poulations that dont exchange genes very
frequently, arising in two different species.)
o vicariance (a single population is split into two by a spontaneous
barrier and there is no movement from one location to another)
- dispersal ex: fruit flies (the ones on newer islands have diverged sometime in
the near past while the ones on older islands diverged earlier and have a
more distant ancestor)

LECTURE 5: SPECIATION (CONTD)

Allopatric Speciation
- a population becomes divided by a barrier that blocks gene flow
- the genetic makeup slowly diverges
- as they become genetically different, isolation mechanisms evolve
- ex: snapping shrimp that have diverged due to vicariance. Theres a land
bridge that fromed about 3M years ago and due to the similar speces on both
sides of the land bridge, it was thougt that species used to live all together. A
phylogeny was constructed and it was found that sister species lived on
opposite sides of the land bridge
o the land bridge formed between north and south America. This land
bridge filled in gradually and not all at once so the species diverged
slowly over time

Dispersalism
- many species have a bizzare distribution
- ex: some plants are found in different continenets that are widely spread out
- how did these plants be able to colonize these lands that are sperated that
oceanic barriers?
- Some scientists argue that dispersal can explain this
- Not everyone agrees with this view

Alfred Wegener
- argued continental movement describes the bizarre distribution instead of
dispersalism
- basially argued for continental drift
 - proposed that at one time all the continents were all joined together into one
continent, that he named Pangea
- this continent began breaking up 200M years ago
- evicence:
o jigsaw puzzle: the edges of the continents seem to fit together like a
puzzle
o fossil distributions: number of species that were found in distant parts
of the world that dont seem to have capapbility ot cross the barriers
that separate the regions so they mustve all been in one area at some
point (all lived together on Pangea when the continents were joined
together)
o matching rock formations: same geologically similar rocks in separate
areas (mountain ranges match up)
o ancient climates: glacial striations in S. Africa that show that the
continents mustve been closer together and Africa mustve been close
to Antarctica
- he couldnt figure out why continnents would move. He proposed that they
plow their way through the ocean floor but there is no evidence on the ocean
floor to prove this
- because he couldnt explain how they move, his theory was ridiculed
- in the 1960s, there were technological advances that prove his theory
- modern evidence:
o seafloor spreading: oceanographers started to map the ocean floor
and found features such as ridges of submarine volcanic mountians
and systems of deep trenches (ie Mariana Trench).
Plate tectonics has explained this
Volcanic mountain ranges are thought to be pieces of
the seafloor moving apart
Trenehces are when one part of the crust is going
underneath another piece
- Theory of plate tectonics: earths crust is divided into tectonic plates
o Movement of plates is very slow
o This movement changed continent distribution

Changing patterns of continents

- most important period was the last 400M years
- 400M years ago there were separate pieces and they jumbled together 260M
years ago
- 160M years, Pangea starting the break down into two continents
- 135M years ago, Gondwana (one of the continients) began to break up. All
the fragments started to move north and eventually collided together
- Wallaces Line is a transitional line that essentially seperates species and the
species on either side of this line are very different
- Note: Laurasia broke up much more recently than Gondwana so species
between NA and Europe are somewhat similar but there are still differences
Continental drift and vicariance
- continental drift is an important source for vicariance
- the ratities are similar but all distinctily similar
o this diversity has arisen by the breakup of Pangea
o the ancestor is thought to have lived in Gondwana
o as Gondwana split, allopatric species occurred on the isolated
fragments
o this lead to the diversity of the present species we have today

Geetic change in isolated populations

- mechanisms that lead to gnetic change:
o 1. Genetic drift: ex given is the forst which doesnt discrimante on
moth colour. So when 80% ar elimated, we dont know what will be
left over and whatever is left over is a different population that will go
on to pass their genes
o 2. Natural selection: probably the most important mechanisms
ex: apple maggot fly (original habitat was hawthorn fruit but
settlers in the area started planting apple trees and the flies
started going to the apples)
apples mature quicker so flies using apples were favoured by
natural selection so now the apple flies and the hawthorn flies
are separate species
o 3. Sexual selection: some indidvuals have better ability to get mates

Sympatric Speciation
- speciation without geographical isiolation
- how does it happen?
o Polyploidy
An error occurs in cell division when the sperm and egg meet.
If the new cell from the error is viable, it will meet with
another cell. If the combination results in an odd number of
chromosomes, the chromosomes dont pair up and it is sterile

Lecture 6: Speciation cont and DNA Barcoding

Polyploidy (speciation without barrier)

- few examples of this in animals (because there are only 2 sets of homologous
chromosomes)
- very common in plants because more sets of homologous chromosomes
o because plants can fertilize their own ova (self-fertilization)
- associated with an increase in size (very common in the plant kingdom)
o thi size difference occurs at the overall plant but also at the cellular
level
- a study has found that about 70% of plants have gone thorugh polyploidy at
some point in their history
o polyploidy has been important for the diversity that we see in plants
Disruptive Selection (speciation without barrier)
- ex: the bird
o two different beak thickness (aka two different morphs)
o intermediate sized beaks are rare because those beaks are inefficient
at handling on any type of seed
o over time, this divergence can lead to speciation if birds mate with
their own morph (assortative mating)
o combo of assortative mating and disruptive selection leads to
speciation

DNA Barcoding
- Canadian invention
- Basically a big international science project but is centered and started in
Canada

Traditional Methods
- based on morphology
- dichotomous keys to decide what a species was
- many taxonomists required if we want to identify all the species so this
method is cumbersome to use

DNA Barcoding
- identifiy species based on differences in the DNA
- differences are species specific
- there is also differences in individuals of the same species, closely related
species, species in the same family (there is variation at different levels)
- we need to find variation that distinguishes between species
- we need to find the ideal gene

Ideal Gene
- should be a gene that everything shares (ubiquitous)
- gene needs to vary between species but not vary within the species

Good Candidate:
- ITS region of ribosomal genes
- Every cell has ribosomes
- Synthesize proteins
- Ribosomes have 2 part structure (large and small subunit that fit together)
- Each subunit has its own componenets
- S is a measure of size (the larger the number, the bigger the object)
- ITS regions are cut out to make the final rRNA
- ITS regions mutate without harm to the organism because even if they
mutate, they are cut out so they dont affect the organism
- ITS regions are the same in a species but vary between species
Techniques in DNA Barcoding
1. PCR
- targets one region of the DNA using primers
- multiple copies made for a chemical analysis
- there are regions (18S, 5.8S, 28S)
- that dont change very fast (highly conserved) which are close to regions
that change rapidly so we can make primers that will bind to these regions
and copy the inbetween (variable/ITS) region
2. Chain termination sequencing
- normal nucleotides and fluorescently labeled nucleotides
- these fluorescent nucleaotides stop any further elongation (the enzyme
attaching nucleotides cant do so anymore)
- this results in a different length DNA in each product
3. Gel Electrophoresis
- seperates fragments according to size
- smallest fragments move the largest distance in the gel
- the original DNA strang can be sequenced based on this information

Barcode Gap
- essential to make gene useful for barcoding
- gap refers to the amount of difference of information when comparing
indivudla of same species vs indviduals of different species
- ideally, the amount of variation is small in individuals of the same species vs
the amount of variation btween species
- the ITS gap is not equally good for everything (good for individuals of same
species but not between species or its not a good measure for every species)
- easier to distinguish genera (Rather than species)

Other Barcoding Regions

- CO1 gene found in mitochondria
- Mitochondria has its own DNA, which CO1 is part of
- Found in organisms that do aerobic respiration
- Used in animal barcoding but not in plant barcoding

Cryptic Species
- DNA barcoding has discovered many new species
- Case study: butterfly
o Adults look alike so theyve been classified as one species
o The caterpillars come in different colour morphs
o CO1 DNA barcoding was used to see the variation
o Turns out there is different species because barcode gaps were found
o So this one species is actually 10 cryptic species
o Cryptic species = species that cant be identified based strictly on
appearance

Graph shows that dna barcoding finds species faster

LECTURE 7: DNA BARCODING (CONTD) AND SPECIES RICHNESS THROUGH
TIME

Barcoding Applications
- food quality assurance
o can confirm what species is being sold (ie when selling fish, you can
determine if it is actually what they say it is)
o screen for contaminants (ie if ground beef has other ingredients)
o pathogen screening (ie ecoli)
- environmental monitoring
o looking for presence of certain species that indicate contamination
(indicator species)
o detect invasive species (ie if theyre from another part of the world)
ex: silver carp (brought to NA to provide food for restaurants.
Lived in sperated fish poonds. A few years ago was a flood and
a river merged with the ponds and now these fish are
extending their ranges. Concerns about the fish going to Lake
Erie. DNA barcoding detected silver carp DNA but no live fish
have been found yet. These carp are very destructive because
they are like vaccumms they eat plankton which other fish
depend on because they are the basis of the food chain. They
are also really big. They have no stomachs because plankton is
easy to digest. Could have major impact on native fish. These
fish jump, so concern about injury to humans)
ex: bugs eating away at leaves
- wildlife protection
- bird/wildlife stikes
o ie when birds are involved in aircraft collisions

Critcism
- criticisms for different reasons by many biologists (shown in pie chart)
- barcoding doesnt provide as much info as morphology does (morphology is
rich)
- sp concept barcoding is too vague

Barcoding is Unjust
- it is expensive, not accessible to everyone
- this is problem but not a good reason to not use it at all
- trying to get costs down

Doesnt have species concept

- controversy about what is and isnt a speciesta
- however this is a problem throughout all of taxonomy and will be until there
is one clear definitation of a specie concept
Phentic
- iBOL phylograms dont take into account many differences between species
and their shared characters
- the data can be downloaded and reanalyzed to this is only a matter of how
the data is displayd at one point

Funding Issues
- barcoding takes money away from traditional taxonomy but in fact, since
barcoding, funding has gone up
- no evidence that barcoding is taking money away from other types of
taxonomists

Damaging to children (other category)

- not a valid criticisms

Traditional Taxonomist Attacks

- identification accuracy
o traditional taxomy methods have a high error rate (due to a variety of
reasons)
o error with barcoding seems to be lower and not higher than that of
traditional methods
- morphology isnt rich at the species level (some species cant be identified as
different using traditional methods)
- traditional methods use smaller specimens while barcoding uses more
specimen so the sample size is bigger, leading to more accurte results

Differences
- morphological gap is used in traditional methods
o this is similar to the barcode approach
o the barcode gap needs to be more flexible (ie how much of a gap is
required to distinguish between species)

Species Richness
- commonly used way of assessing biodiversity
- is a simple number
- overlooks a lot of diversity occurring at other levels (ie rarity of a species)
- richness varies on a number of scales (ie some continents have higher
richness, more richness closer to equator and less close to the poles)

The Geologic Recod

- a history of the earth based on fossils
- 3 major eons (largest time periods) (phanerzoic most important with
respect to animal life and the one we are living in right now)
- eons subdivided into eras, divided in periods, divided into epochs, divided
into ages
 - boundaires between blocks represent some time of extinction (ie Mesozoic
era the time of the dinosaurs boundaries of it are mass extinctions)
o large time blocks are more dramatic extinctions and smaller blocks
are smaller extinctions

- earth and other planets came out of a dust cloud

o dense pockets became planets
o low density pockets dissipated
o debris (comets and asteroid) hit planets. This is how it is thought that
earth got the water)

Stromatolites
- oldest known fossils
- composed of mainly bacteria so they are considered to be alive
- bacteria secretes mucous that traps sediment and bacteria move upwards
and continue trapping sediment in more mucous so its layered
- when the layers fossilize, stromatolites for
- life originated soon after liquid water was on earth

First living cells

- prokaryotic
o DNA clustered in cell (no nucleus)
o Very simple organisms
- Anerobic
o Didnt require oxygen

Origin of O2
- cyanobacteria can also do photosynthesis
- iron oxides in old rocks shows when oxygen was present in the environment
- cyanobacteria were probably the first photosynthetic organisms
- oxygen presence increased very slowly through time
- oxygen revolution = sudden increase in O2
- o2 can damage living things so o2 probably killed many microbes
- aerobic respiration is basically the reverse of photosynthesis
- aerobic resipiration gets more energy from glucose than glycolysis does

Eukaryotic Cells
- first eukaryotes can be dated in fossils
- some evidence that they existed before this fossil
- have cytoskeleton