Basic Cladistics
-
Upload
donna-mariel-calimpong -
Category
Documents
-
view
215 -
download
0
Transcript of Basic Cladistics
7/29/2019 Basic Cladistics
http://slidepdf.com/reader/full/basic-cladistics 1/91
7/29/2019 Basic Cladistics
http://slidepdf.com/reader/full/basic-cladistics 2/91
• The Phylogeny of an organism is traced back,
it connects through shared ancestors to lineages
of other organisms.
**phylogenetic tree
Cladistics or Phylogenetic Systematics
7/29/2019 Basic Cladistics
http://slidepdf.com/reader/full/basic-cladistics 3/91
• Given that closely related species share acommon ancestor and often resemble eachother, it might seem that the best way to uncover
the evolutionary relationships would be withoverall similarity.
• Q: In other words, out of a group of species, if two are most similar, can we reasonably
hypothesize that they are closest relatives?
YES or NO?
7/29/2019 Basic Cladistics
http://slidepdf.com/reader/full/basic-cladistics 4/91
• Overall similarity may be misleading becausethere are actually two reasons why organisms
have similar characteristics and only one of them is due to evolutionary relatedness.
• homologous feature (or homology)- Whentwo species have a similar characteristicbecause it was inherited by both from acommon ancestor
• Ex. eventoed foot of the deer, camels, cattle,
pigs, and hippopotamus is a homologoussimilarity because all inherited the featurefrom their common paleodont ancestor.
7/29/2019 Basic Cladistics
http://slidepdf.com/reader/full/basic-cladistics 5/91
7/29/2019 Basic Cladistics
http://slidepdf.com/reader/full/basic-cladistics 6/91
7/29/2019 Basic Cladistics
http://slidepdf.com/reader/full/basic-cladistics 7/91
• analogous feature (or homoplasy)- When twospecies have a similar characteristic because of convergent evolution
• Convergent evolution - when unrelated speciesadopt a similar way of life, their body parts maytake on similar functions and end up resemblingone another
• Ex. The paddle-like front limb and streamlinedbodies of the aquatic animals shown in the figurefor convergent evolution are examples of analogous features.
7/29/2019 Basic Cladistics
http://slidepdf.com/reader/full/basic-cladistics 8/91
• Only homologous similarity is evidence that
two species are evolutionarily related.• Q: If two animals share the highest number of
homologies, can we reasonably assume theyare closest relatives?
• YES or NO?
• a homology may be recently derived or anancient retained feature; only shared recent
homologies (called synapomorphies) areevidence that two organisms are closelyrelated.
7/29/2019 Basic Cladistics
http://slidepdf.com/reader/full/basic-cladistics 9/91
• Ex. The hand of the first vertebrates to live on
land had five digits (fingers).
• Many living terrestrial vertebrates (such as
humans, turtles, crocodiles and frogs) also
have five digits because they inherited them
from this common ancestor. This feature is
then homologous in all of these species.
7/29/2019 Basic Cladistics
http://slidepdf.com/reader/full/basic-cladistics 10/91
• In contrast, horses, zebras and donkeys have justa single digit with a hoof.
• Clearly, humans are more closely related to
horses, zebras and donkeys, even though theyhave a homology in common with turtles,crocodiles and frogs.
• The key point is that the five digit condition is the
primitive state for the number of digits. It wasmodified and reduced to just one digit in thecommon ancestor of horses, donkeys and zebras.
7/29/2019 Basic Cladistics
http://slidepdf.com/reader/full/basic-cladistics 11/91
• In an important work (first published inEnglish in 1966) by the German entomologistWilli Hennig, it was argued that only sharedderived characters could possibly give usinformation about phylogeny.
• The method that groups organisms that share
derived characters is called cladistics orphylogenetic systematics.
• Taxa that share many derived characters are
grouped more closely together than thosethat do not. The relationships are shown in abranching hierarchical tree called acladogram.
7/29/2019 Basic Cladistics
http://slidepdf.com/reader/full/basic-cladistics 12/91
• The cladogram is constructed such that the
number of changes from one character state
to the next is minimized. The principle behind
this is the rule of parsimony - any hypothesis
that requires fewer assumptions is a more
defensible hypothesis.
7/29/2019 Basic Cladistics
http://slidepdf.com/reader/full/basic-cladistics 13/91
7/29/2019 Basic Cladistics
http://slidepdf.com/reader/full/basic-cladistics 14/91
7/29/2019 Basic Cladistics
http://slidepdf.com/reader/full/basic-cladistics 15/91
0 1 2 30 to 1 = 1 step 1 to 0 = 1 step 1 to 2 = 2 steps
7/29/2019 Basic Cladistics
http://slidepdf.com/reader/full/basic-cladistics 16/91
7/29/2019 Basic Cladistics
http://slidepdf.com/reader/full/basic-cladistics 17/91
0 1 2 30 to 1 = 1 step 1 to 0 = 1 step 1 to 2 = 1 step
7/29/2019 Basic Cladistics
http://slidepdf.com/reader/full/basic-cladistics 18/91
0 1 2 30 to 1 = 1 step 1 to 0 = NA 1 to 2 = 1 step
7/29/2019 Basic Cladistics
http://slidepdf.com/reader/full/basic-cladistics 19/91
7/29/2019 Basic Cladistics
http://slidepdf.com/reader/full/basic-cladistics 20/91
7/29/2019 Basic Cladistics
http://slidepdf.com/reader/full/basic-cladistics 21/91
7/29/2019 Basic Cladistics
http://slidepdf.com/reader/full/basic-cladistics 22/91
Titanotheres
7/29/2019 Basic Cladistics
http://slidepdf.com/reader/full/basic-cladistics 23/91
7/29/2019 Basic Cladistics
http://slidepdf.com/reader/full/basic-cladistics 24/91
7/29/2019 Basic Cladistics
http://slidepdf.com/reader/full/basic-cladistics 25/91
0 1 2 30 to 1 = 1 step 1 to 0 = NA 1 to 2 = 1 step
7/29/2019 Basic Cladistics
http://slidepdf.com/reader/full/basic-cladistics 26/91
Multiple origins
1
2
3
7/29/2019 Basic Cladistics
http://slidepdf.com/reader/full/basic-cladistics 27/91
7/29/2019 Basic Cladistics
http://slidepdf.com/reader/full/basic-cladistics 28/91
7/29/2019 Basic Cladistics
http://slidepdf.com/reader/full/basic-cladistics 29/91
• If the character has only two states, then the task of distinguishing primitive and derived character states isfairly simple: The state which is in the outgroup isprimitive and the one found only in the ingroup is
derived.• It is common practice to designate the primitive states
as 0 (zero) and the derived states as 1 (one). If you aregoing to calculate trees by hand, this will certainlymake your calculations easier.
• On the other hand, if you are using a computerprogram to calculate a tree, it isn’t necessary todesignate the plesiomorphic state as 0 (zero):
7/29/2019 Basic Cladistics
http://slidepdf.com/reader/full/basic-cladistics 30/91
• The first step in basic cladistic analysis is todetermine which character states are primitiveand which are derived.
• The outgroup comparison method is the primaryone in use today.
• In outgroup comparison, if a taxon that is not a
member of the group of organisms beingclassified has a character state that is the same assome of the organisms in the group, then thatcharacter state can be considered to be
plesiomorphic.
• The outside taxon is called the outgroup and theorganisms being classified are the ingroup.
Asteroidea
7/29/2019 Basic Cladistics
http://slidepdf.com/reader/full/basic-cladistics 31/91
Asteroidea
Sea stars and starfishes
• Outgroup comparison
7/29/2019 Basic Cladistics
http://slidepdf.com/reader/full/basic-cladistics 32/91
7/29/2019 Basic Cladistics
http://slidepdf.com/reader/full/basic-cladistics 33/91
• In common cladistic usage, a monophyletic groupis a taxon (group of organisms) which forms aclade, meaning that it contains all thedescendants of the possibly hypothetical closestcommon ancestor of the members of the group.
• The term is synonymous with the uncommonterm holophyly.
• Monophyletic groups are typically characterizedby shared derived characteristics(synapomorphies).
7/29/2019 Basic Cladistics
http://slidepdf.com/reader/full/basic-cladistics 34/91
7/29/2019 Basic Cladistics
http://slidepdf.com/reader/full/basic-cladistics 35/91
• In current usage, a paraphyletic group consists of all of the descendants of a possibly hypotheticalclosest common ancestor minus one or more
monophyletic groups (most usually one).• A paraphyletic group is thus 'nearly'
monophyletic (consistent with the meaning of the prefix 'para', namely 'near' or 'alongside'.)
• A polyphyletic group is any group other than amonophyletic group or a paraphyletic group,which like a paraphyletic group contains onlysome of the descendants of their closest
common ancestor, but unlike a paraphyleticgroup is not characterized by the missingdescendants forming one (or more) monophyleticgroups.
7/29/2019 Basic Cladistics
http://slidepdf.com/reader/full/basic-cladistics 36/91
• A clade is a group of taxa consisting only of an
7/29/2019 Basic Cladistics
http://slidepdf.com/reader/full/basic-cladistics 37/91
• A clade is a group of taxa consisting only of an
ancestor taxon and all of its descendant taxa.
• In the diagram provided (a cladogram), it is
hypothesized that all vertebrates, including ray-finned fishes (Actinopterygii), had a common
ancestor all of whose descendants were vertebrates,
and so form a clade.
• Within the vertebrates, all tetrapods, including
amphibians, mammals, reptiles (as traditionally
defined) and birds are hypothesized to have had a
common ancestor all of whose descendants were
tetrapods, and so also form a clade.
• The tetrapod ancestor was a descendant of the
original vertebrate ancestor, but is not an ancestor of
any ray-finned fish living today.
7/29/2019 Basic Cladistics
http://slidepdf.com/reader/full/basic-cladistics 38/91
The relationship between clades can be
described in several ways:
1. A clade is basal to another clade if it containsthat other clade as a subset within it.
• In the example, the vertebrate clade is basal
to the tetrapod and ray-finned fish clades.Note:(Some authors have used "basal"
differently to mean a clade that is less species-
rich than a sister clade, with such a deficit
being taken as an indication of 'primitiveness'.
Others consider this usage to be incorrect.
7/29/2019 Basic Cladistics
http://slidepdf.com/reader/full/basic-cladistics 39/91
• A clade located within a clade is said to be
nested within that clade. In the diagram, the
tetrapod clade is nested within the vertebrate
clade.
• Two clades are sisters if they have an
immediate common ancestor. In the diagram,
crocodiles and birds are sister clades, as aremodern amphibians and amniotes.
7/29/2019 Basic Cladistics
http://slidepdf.com/reader/full/basic-cladistics 40/91
• Terminology for characters
The following terms are used to identify shared or distinct
characters among groups:
• Plesiomorphy ("close form") or ancestral state, alsosymplesiomorphy ("shared plesiomorphy", i.e. "shared close
form"), is a characteristic that is present at the base of a tree
(cladogram). Since a plesiomorphy that is inherited from the
common ancestor may appear anywhere in a tree, itspresence provides no evidence of relationships within the
tree. The traditional definition of reptiles (the blue group in
the diagram) includes being cold-blooded (i.e. not maintaining
a constant high body temperature), whereas birds are warm-
blooded. Since cold-bloodedness is a plesiomorphy, inherited
from the common ancestor of traditional reptiles and birds, it
should not be used to define a group in a system based on
cladistics.
7/29/2019 Basic Cladistics
http://slidepdf.com/reader/full/basic-cladistics 41/91
7/29/2019 Basic Cladistics
http://slidepdf.com/reader/full/basic-cladistics 42/91
• Apomorphy ("separate form") or derived state
is a characteristic believed to have evolved
within the tree. It can thus be used toseparate one group in the tree from the rest.
• Within the group which shares the
apomorphy it is a synapomorphy ("sharedapomorphy", i.e. "shared separate form"). For
example, within the vertebrates, all tetrapods
(and only tetrapods) have four limbs; thus,
having four limbs is a synapomorphy for
tetrapods. All the tetrapods can legitimately
be grouped together because they have four
limbs.
7/29/2019 Basic Cladistics
http://slidepdf.com/reader/full/basic-cladistics 43/91
• Homoplasy is a characteristic shared by
members of a tree but not present in their
common ancestor.• It arises by convergence or reversion. Both
mammals and birds are able to maintain a
high constant body temperature (i.e. they are'warm-blooded'). However, the ancestors of
each group did not share this character, so it
must have evolved independently. Mammals
and birds should not be grouped together on
the basis that they are warm-blooded.
7/29/2019 Basic Cladistics
http://slidepdf.com/reader/full/basic-cladistics 44/91
7/29/2019 Basic Cladistics
http://slidepdf.com/reader/full/basic-cladistics 45/91
CLADOGRAM
WHAT A CLADOGRAM ACTUALLY SAYS
7/29/2019 Basic Cladistics
http://slidepdf.com/reader/full/basic-cladistics 46/91
WHAT A CLADOGRAM ACTUALLY SAYS
ABOUT RELATIONSHIPS
• The trees that result from cladistic analysis are relativestatements of relationship and do not indicateancestors or descendants.
• For example in the tree above, Prorodon teres and
Prorodon marina are hypothesized to be sister taxaand to share a more recent common ancestor witheach other than with Coleps; but the prorodontids (P.teres+P. marina+Coleps) all share a more recent common ancestor with one another than with the
Placidae (Placus + Spathidiopsis).• The tree does not explicitly hypothesize ancestor-
descendant relationships. In other words, the treehypothesizes that Prorodon and Coleps are related, but not that Prorodon evolved from Coleps or that Coleps
evolved from Prorodon.
7/29/2019 Basic Cladistics
http://slidepdf.com/reader/full/basic-cladistics 47/91
TAXON
A B C D
Character
1 1 1 0 0
2 1 1 0 0
3 1 1 1 0
4 1 1 1 0
5 1 1 1 0
6 1 1 1 0
7 0 1 0 0
8 0 1 0 0
9 0 1 0 0
Applying cladisticsTAXON
A B C D
A 6 4
B
C
D
We now count the
derived similarities
between A and C.
7/29/2019 Basic Cladistics
http://slidepdf.com/reader/full/basic-cladistics 48/91
TAXON
A B C D
Character
1 1 1 0 0
2 1 1 0 0
3 1 1 1 0
4 1 1 1 0
5 1 1 1 0
6 1 1 1 0
7 0 1 0 0
8 0 1 0 0
9 0 1 0 0
Applying cladisticsTAXON
A B C D
A 6 4 0
B
C
D
And the similarities in
derived traits between
A and D.
7/29/2019 Basic Cladistics
http://slidepdf.com/reader/full/basic-cladistics 49/91
TAXONA B C D
Character
1 1 1 0 0
2 1 1 0 0
3 1 1 1 0
4 1 1 1 0
5 1 1 1 0
6 1 1 1 0
7 0 1 0 0
8 0 1 0 09 0 1 0 0
Applying cladisticsTAXON
A B C D
A 6 4 0
B 4 0
C 0
D
The completed table
shows all pairwise
similarities.
7/29/2019 Basic Cladistics
http://slidepdf.com/reader/full/basic-cladistics 50/91
TAXON
A B C D
A 6 4 0
B 4 0
C 0
D
Phenetics Vs. cladistics
TAXON
A B C D
A 6 7 3
B 4 0
C 5
D
Notice that the matrices produced
by phenetics (right) and cladistics
are quite different.
P C
7/29/2019 Basic Cladistics
http://slidepdf.com/reader/full/basic-cladistics 51/91
• The similarity
matrix is now used
to create the
cladogram.
• A is most similar to
B.
Applying cladistics
TAXON
A B C D
A 6 4 0
B 4 0
C 0
D
A B
7/29/2019 Basic Cladistics
http://slidepdf.com/reader/full/basic-cladistics 52/91
• The similarity
matrix is now used
to create the
cladogram.
• Next, (A and B) are
together most
similar to C.
Applying cladistics
TAXON
A B C D
A 6 4 0
B 4 0
C 0
D
A B C
7/29/2019 Basic Cladistics
http://slidepdf.com/reader/full/basic-cladistics 53/91
Phenetics Vs. cladistics
The phylogenies produced by
phenetics (right) and cladistics are
also quite different.
P C
A C B D A B C D
7/29/2019 Basic Cladistics
http://slidepdf.com/reader/full/basic-cladistics 54/91
A B C D
Now what?
• We can now plot the
changes that
occurred on the tree,getting a feeling for
the actual process.
7/29/2019 Basic Cladistics
http://slidepdf.com/reader/full/basic-cladistics 55/91
Reconstructing phylogenies (1)
• The number of possible trees quickly increases
as the number of taxa goes up.
A B
2 taxa
A B C C B A A C B
3 taxa
A B C D A B C D A C B D A C B D A D B C
4 taxa
+
7/29/2019 Basic Cladistics
http://slidepdf.com/reader/full/basic-cladistics 56/91
• Mathematically, the number of possibletrees is:
Reconstructing phylogenies (2)
(2n - 3)!
2n-2(n-2)!
• This quickly gets out of control.
• With many taxa and characters, computer
algorithms must be used.
Taxa trees
4 156 9458 135135
10 34459425
7/29/2019 Basic Cladistics
http://slidepdf.com/reader/full/basic-cladistics 57/91
Exhaustive
Branch-and-bound
Heuristic
7/29/2019 Basic Cladistics
http://slidepdf.com/reader/full/basic-cladistics 58/91
7/29/2019 Basic Cladistics
http://slidepdf.com/reader/full/basic-cladistics 59/91
Monophyly of Endomychidae
Consensus Tree Length:
89 CI: 57RI: 81
7/29/2019 Basic Cladistics
http://slidepdf.com/reader/full/basic-cladistics 60/91
Monophyly of Endomychidae
Nelsen Consensus Tree
Length: 89CI: 57
RI: 81
Monophyly Supported by:
Fronto-clypeal ridge present
7/29/2019 Basic Cladistics
http://slidepdf.com/reader/full/basic-cladistics 61/91
Monophyly of Endomychidae
Nelsen Consensus Tree
Length: 89CI: 57
RI: 81
Monophyly Supported by:
Fronto-clypeal ridge present
Head without antennal grooves
Tarsi 4-segmented, simple
h l f d h d
7/29/2019 Basic Cladistics
http://slidepdf.com/reader/full/basic-cladistics 62/91
Monophyly of Endomychidae
Nelsen Consensus Tree
Length: 89CI: 57
RI: 81
Monophyly Supported by:
Fronto-clypeal ridge present
Head without antennal grooves
Tarsi 4-segmented, simple
7/29/2019 Basic Cladistics
http://slidepdf.com/reader/full/basic-cladistics 63/91
7/29/2019 Basic Cladistics
http://slidepdf.com/reader/full/basic-cladistics 64/91
7/29/2019 Basic Cladistics
http://slidepdf.com/reader/full/basic-cladistics 65/91
7/29/2019 Basic Cladistics
http://slidepdf.com/reader/full/basic-cladistics 66/91
7/29/2019 Basic Cladistics
http://slidepdf.com/reader/full/basic-cladistics 67/91
7/29/2019 Basic Cladistics
http://slidepdf.com/reader/full/basic-cladistics 68/91
7/29/2019 Basic Cladistics
http://slidepdf.com/reader/full/basic-cladistics 69/91
7/29/2019 Basic Cladistics
http://slidepdf.com/reader/full/basic-cladistics 70/91
7/29/2019 Basic Cladistics
http://slidepdf.com/reader/full/basic-cladistics 71/91
7/29/2019 Basic Cladistics
http://slidepdf.com/reader/full/basic-cladistics 72/91
7/29/2019 Basic Cladistics
http://slidepdf.com/reader/full/basic-cladistics 73/91
7/29/2019 Basic Cladistics
http://slidepdf.com/reader/full/basic-cladistics 74/91
7/29/2019 Basic Cladistics
http://slidepdf.com/reader/full/basic-cladistics 75/91
7/29/2019 Basic Cladistics
http://slidepdf.com/reader/full/basic-cladistics 76/91
7/29/2019 Basic Cladistics
http://slidepdf.com/reader/full/basic-cladistics 77/91
7/29/2019 Basic Cladistics
http://slidepdf.com/reader/full/basic-cladistics 78/91
7/29/2019 Basic Cladistics
http://slidepdf.com/reader/full/basic-cladistics 79/91
7/29/2019 Basic Cladistics
http://slidepdf.com/reader/full/basic-cladistics 80/91
7/29/2019 Basic Cladistics
http://slidepdf.com/reader/full/basic-cladistics 81/91
7/29/2019 Basic Cladistics
http://slidepdf.com/reader/full/basic-cladistics 82/91
7/29/2019 Basic Cladistics
http://slidepdf.com/reader/full/basic-cladistics 83/91
7/29/2019 Basic Cladistics
http://slidepdf.com/reader/full/basic-cladistics 84/91
7/29/2019 Basic Cladistics
http://slidepdf.com/reader/full/basic-cladistics 85/91
7/29/2019 Basic Cladistics
http://slidepdf.com/reader/full/basic-cladistics 86/91
7/29/2019 Basic Cladistics
http://slidepdf.com/reader/full/basic-cladistics 87/91
7/29/2019 Basic Cladistics
http://slidepdf.com/reader/full/basic-cladistics 88/91
7/29/2019 Basic Cladistics
http://slidepdf.com/reader/full/basic-cladistics 89/91
7/29/2019 Basic Cladistics
http://slidepdf.com/reader/full/basic-cladistics 90/91
7/29/2019 Basic Cladistics
http://slidepdf.com/reader/full/basic-cladistics 91/91