Feel free to
discuss this and other assignments with other students in the class. You can
learn as much from them as you can from the TA and myself (sometimes).
Reconstructing phylogenetic relationships and assigning species to higher taxa
on the basis of those relationships is one of the primary goals of many
paleobiologists. This exercise is designed to give you practice in the use of
taxonomy and phylogenetic systematics ("cladistics").
4.1 Introduction
to Systematics
If it were not for
taxonomy and sytematics, our understanding of the relationships among living
things would be formless and chaotic. Taxonomic classification provides the
hangers on which we organize our knowledge of morphology and express hypotheses
of relationships. The classification system that we use today was established
by Carolus Linnaeus, a Swedish biologist, in the 18th century. It provides a
hierarchy of categories used to classify each living organism. From the largest
(most inclusive) to the smallest (least inclusive), these categories include:
|
Order
of Hierarchy
|
Mnemonic
(memorization device) |
|
Kingdom
|
Kings
|
|
Phylum |
Play |
|
Class |
Chess |
|
Order |
On |
|
Family |
Funny |
|
Genus |
Green |
|
species |
squares |
|
|
|
In our studies of
the actual organisms, we will focus mainly on the higher categories of phylum and
class. We will deal with lower levels, such as specific genera and species
principally when a specific one helps illustrate some important concepts
(otherwise the names just get too numerous!). In order to keep information
organized however, you will need to remember the order of this hierarchy. One
way to memorize the order of these categories is to use a mnemonic such as the
one above, or, if you prefer, make one up yourself.
4.1.1 Species
Concepts and Binomial Nomenclature
Linneaus
established in his taxonomy a system of binomial nomeclature, in which every
living organism has a binomial ("two-part name") consisting of its
genus name and species name, in that order. The best-known example of a
binomial is Homo sapiens, the genus and species to which modern humans
belong. The first letter of a genus name is always capitalized; the species
name is never capitalized, and it never appears without the generic name or its
initial (i.e. Homo sapiens or H. sapiens). Both names must have a
certain Latin form and, for this reason, are underlined or italicized. All
countries use both binomial nomenclature and the hierarchy of categories
established by Linneaus (sub-categories are also often added); this makes it
possible for systematists around the world to use a common system of names,
regardless of their native language. If this seems odd, stop and imagine, for
just a moment, what would happen if we all had to memorize a taxonomic name in
all its possible translated forms. Can you imagine the confusion and extra
names that might result? (It sometimes happens anyways)
For students,
learning classification is one of the first steps toward sorting out the many
new names and morphological characters that appear each lesson. At least at
first, students usually prefer that classification remains as unchanging as
possible. After all, who wants to see five alternate classifications, when one
is coomplicated enough? Systematists, however, are part of a dynamic science
for which an unchanging classification would represent a lack of progress in
understanding the organisms they study. Both new evidence, such as new fossil
discoveries, and new approaches constantly change our way of looking at
relationships.
One of the most
important changes in systematic thinking in recent years has been a shift to
the use of phylogenetic systematics or "cladistics", as some people
call it. According to phylogenetic systematics, whose philosophical father was
a German entomologist named Willi Hennig, classification should always reflect
evolutionary relationships of the organisms involved. Hennig's recommendation
may sound obvious, but in fact most classifications were not-and many still are
not-constructed strictly on the basis of evolutionary relationship.
4.1.2 Taxonomy
vs. Systematics
Systematics is not
taxonomy (and vice versa). Taxonomy is merely the classification and naming of
organisms. It is not necessarily dependent on systematics, but most people
agree that it should be. Grouping organisms according to the extent to which
they are related is the most logical method, but other (subjective and somewhat
less logical) methods have also been used.
Taxonomy is useful for identifying and discussing organisms. Systematics
is a scientific discipline in its own right.
4.1.3
Phylogenetic Systematics
Phylogenetics is a
method for determining the evolutionary relationships of organisms. The goal is
to try to understand which organisms are most closely related to each other.
This is accomplished by comparing ancestors and descendants.
4.1.3a How does
it work? Trees . . .
Evolutionary
relationships are represented by phylogenetic trees (a.k.a. phylogenies). In
this section you will become familiar with the terminology used to describe
phylogenetic trees and the relationships of the organisms in the phylogeny.
How this is
different from taxonomy? Taxonomy is
merely the naming and classification of organisms (Remember: Kingdom, Phylum,
Class, Order, Genus, Species).
Phylogenies, on the other hand, give paleobiologists valuable
information about the relationship between an ancestor and its
descendents. It provides a hypothesis
for the evolution of related organisms.

Figure 4.1 An example of a phylogenetic tree. This phylogeny indicates that a horse and whale share a common
ancestor (implied by the node B’). They are more closely related to each other
than either is to the trout. However, the
horse, whale and trout are distantly related because they share an ancestor at
node A’ (from Lucas,2000).
Tree terminology….
Indicate that there has been a change in character state Nodes= Branch Points Terminal Taxa TIME PRESENT PAST

POLYPHYLETIC
Groups on a phylogenetic
tree:
PARAPHYLETIC MONOPHYLETIC


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A monophyletic
group is often called a clade. It includes an ancestor and all of
its descendants .
A polyphyletic
group is one that is derived from many ancestors.
A paraphyletic
group is one that includes the ancestor, but not all of its descendants.
Sister groups share a common ancestor and
are each other's closest relatives.
(For example, D&E and B&C in the above figure)
4.2
Excercises
4.2.1
Questions
Use
the tree below to answer the following questions.
G F E D C B A

1. Circle those which are
monophyletic groups:
ABC
CDE EF
EFG ABCD
BCEF DEFG
2. Which pairs are more closely
related? (circle the right answer)
AB or BC
CD or EF EF or
DG AB or EF
3. AB is the sister group to?
_________
4. D is the sister group to?
_________
4.2.2 Characters
The
study of phylogenetics groups organisms into groups (clades, as you learned
earlier). A clade represents an ancestor and all of its descendants. Clades are
united by shared derived characters (synapomorphies). But what is a character?
A character is any
definite aspect of a particular organism. This means that any organism can have
an almost infinite number of characters. So which ones do we use? Put another
way, what makes a character appropriate for use in phylogenetic systematics?
At this time, the
best characters are discrete characters. Discrete characters have a
limited number of possible values. Discrete characters can either be binary
or multistate. Binary characters are those that only have two states,
like Present/Absent. For example, the ability to roll ones tongue is either
Present or Absent. Or, a beetle may be colored either yellow or blue.
Multistate characters on the other hand, have more than two states. For
example, eye color can either be blue, brown or green, that's three character
states.
Continuous
characters
such as size, are measurements on a continuous scale, and they are very hard to
use. For example, the length of a femur can be 10.1cm, 12 cm, 10.4cm, 11.2cm,
etc. The problem is, "Are these sizes "different enough" to
group into separate groups? Or "similar enough" to group into the
same group?" What is "different" and "similar" when it
comes to size? These terms are very subjective and therefore, they will not
help us to define any groups.
4.2.3 Coding
Characters
The easiest way to
do this is to create a code for the characters and their various character
states and to use that information to make a character matrix.
Try this out with
the example on the next page. If you get stuck, be sure to ask for help. Wait
for the rest of the class. We'll go over these characters together.

4.2.4 Character
Terminology
An apomorphy
is a derived character that is shared by all of the members of a clade, but is
not possessed by the immediate ancestors of the clade. If a character is shared
by more than one of the descendants, it is called a synapomorphy, or a
shared derived character. If a character is possessed by only one descendant,
it is termed an autapomorphy.
A plesiomorphy
is an ancestral character. It is a character that is possessed by an ancestor
and, possibly, any of its descendants. Basically, shared ancestral characters (synplesiomorphies)
are not useful in phylogenetic reconstruction. For example, it would be folly
to use "lungs" as a character for grouping all primates because lungs
arose much farther back in the tree of life as an adaptation to the
colonization of land. Thus, “lungs” is not a derived character at the level of
primates because they evolved in a common ancestor much, much earlier. The way
“lungs” can be used as a phylogenetic character is as a shared derived
character for lungfish (the first organism to possess lungs) plus all the other
vertebrates related to lungfish that have them (this includes primates). (Note: “derived” and “primitive” are
relative terms).
In addition, shared
derived characters that developed convergently are not useful in phylogenetic
reconstruction. Many animals have wings: birds, bats, pterosaurs, insects to
name a few. Wings however, are made in very different ways in birds, bats etc.
They are analogous (similar) but not homologous (having the same
genetic and developmental origins), and are therefore described as convergent.
These different animals have "converged" in their evolution of
similar features. The wings of bats are probably a shared derived feature for
bats, but wings in the general sense are developed convergently. Bats are
mammals and insects are arthropods, and there is abundant evidence that birds
are derived from dinosaurs.
The only characters
that help us in phylogenetics are synapomorphies. Synapomorphies define clades.
The others provide information about the taxa in question, but do little to
inform us as to their evolutionary relationships.
Cladistics is a
relative science. We are always conducting phylogenetic analyses on different
scales (like the difference between your family tree, the primate tree and the
tree of all life). It is, therefore, important to note that a synapomorphy at one
level on a phylogenetic tree, could be viewed as a plesiomorphy on another
scale.
Using the tree that
you created, and the characters that you mapped on, answer the following
questions...
1. What one synapomorphy
defines clade ED (Include the character and the change in character state)?
2. Of all the characters
analyzed, which one is an autapomorphy? What taxon has that character?
3. When looking at clade ED,
what is an example of a plesiomorphic, or ancestral character for this group?
(Remember that means that the character arose earlier than that clade, so its a
character that ED shares with other taxa.)
Each
group has a bag that contains fifteen different species from the Kingdom
Hardwarea, Phylum Fastneria.
Your groups’ job is to construct a hierarchical classification of the 15 species in the bag. In doing so, you must identify the defining features (characters) of each class, order, family, genus and species. You can give them names if you want, but that’s optional and not the purpose of this exercise.
In
devising hierarchical schemes, some people work best “from the bottom up”, by
grouping similar species in the same genus, similar genera (the plural of
genus) in the same family, and so on...
Other
people find it more natural to work “from the top down”, by finding the major
divisions among the objects, and then further subdividing those.
Some
ground rules: each class must contain
at least one order, each order at least one family, etc... This means, for example, that several
species could belong to the same genus, several genera to the same family, and
so on...
In
the space on the next page, sketch out your hierarchical scheme and list the
defining morphological features (=characters) of each class, order, family,
genus and species. You will then
compare your group’s classification scheme to the ones devised by the other lab
groups.
Things
to think about:
1.
Is
function a useful way to classify objects?
2.
When
the same characters appear in different groups, is this the result of ancestry
(i.e., the character occurred in the common ancestor) or convergence (i.e., the
character evolved independently in the two groups).
3.
Can
classification be a good guide to evolutionary relationships?