The Classification & Evolution of Caminalcules
Author(s): Robert P. Gendron
Source: The American Biology Teacher, 62(8):570-576.
Published By: National Association of Biology Teachers
DOI: http://dx.doi.org/10.1662/0002-7685(2000)062[0570:TCEOC]2.0.CO;2
URL: http://www.bioone.org/doi/full/10.1662/0002-7685%282000%29062%5B0570%3ATCEOC
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The Classification & Evolution of
Caminalcules
Robert P. Gendron
R
EADERS of this publication understand the with a detailed fossil record. With these fossils, stu-
importance, and difficulty, of teaching evolu- dents can construct a phylogenetic tree from the
tion in an introductory biology course. The bottom up in a way that they find conceptually
difficulty arises, in part, because evolutionary pro- meaningful.
cesses are slow and generally cannot be observed, The laboratory activity consists of three related
even over the course of an entire year. One way to exercises:
circumvent this problem is by using simulations
1. Classifying living Caminalcules into taxonomic
(Stebbins & Brockenbrough 1975; Tashiro 1984; Allen
categories (genera, families, etc.)
et al. 1987; Thelen 1988; Hammersmith & Mertens
2. Using the classification to develop a tentative
1990; Welch 1993; Nolan & Ostrovsky 1996; Dickinson
phylogenetic tree
1998; Lach & Loverude 1998). Another approach
3. Constructing a phylogenetic tree based on the
is to teach evolutionary concepts by constructing
fossil record.
phylogenetic trees (Vogel & Ewel 1972; McComas &
´ One of the main goals of the lab is to illustrate
Alters 1994; Bilardello & Valdes 1998). I have devel-
oped a lab in the latter category for our General the intimate connection between the classification of
Biology program. The idea for this lab came from living species and their evolutionary relationships.
an exercise written by Vogel and Ewel (1972) in
which students developed a classification of fasteners Classification of the Living Caminalcules
(nails, screws and bolts). The lab described here,
however, uses ‘‘organisms’’ with a fossil record, the Students begin by arranging the 14 living species
Caminalcules. into a hierarchical classification (Figure 2). First, they
Caminalcules (Figure 1) are imaginary organisms combine species into genera using the criteria that
invented by the late Joseph H. Camin (Sokal 1983). members of a genus should resemble each other
According to Sokal (1983), Camin created his organ- more closely than they resemble members of other
isms by starting with a primitive ancestor and gradu- genera. (With Caminalcules, as with many real organ-
ally modifying the forms according to accepted rules isms, physical resemblance is usually a good indica-
of evolutionary change. Camin’s intent was to tion of common ancestry.) Using the same criteria,
develop a known phylogeny (something that is gener- genera are combined into families, families into
ally unobtainable for real organisms) that could be orders, and so on. Depending on whether the students
used to critically evaluate different taxonomic tech- are taxonomic ‘‘splitters’’ or ‘‘lumpers,’’ their classifi-
niques such as phenetic and cladistic analysis. cation scheme might stop at Order or go all the way
For the purpose of teaching evolution to college up to Phylum. If class time is short, the students
and high school students, the Caminalcules offer can construct their preliminary classification at home.
several important advantages (McComas & Alters Another way to speed things up is by having the
1994). First, because Caminalcules are artificial organ- entire class work on it together, with the instructor
isms, students have no preconceived ideas about how acting as moderator and facilitator.
they should be classified or how they are related. This exercise teaches several important concepts,
This means that students have to concentrate on beginning with the idea of hierarchical classification
principles rather than prior knowledge when con- itself. Teachers of more advanced classes may wish
structing a phylogenetic tree or classification. Second, to discuss theories of classification (Vogt 1995; Ridley
unlike everyday objects such as fasteners, the Cami- 1996). The concept of convergent evolution is also
nalcules have a ‘‘real’’ evolutionary history, complete introduced, as described below.
Once the students have completed their classifica-
tion, I lead a class discussion with the aid of an
Robert P. Gendron is Professor of Biology at Indiana University overhead projector and transparent images of the 14
of Pennsylvania, Indiana, PA 15705; e-mail: rgendron@
grove.iup.edu. living Caminalcules. I begin with Caminalcule 2 and
ask what other species belong in the same genus.
570 THE AMERICAN BIOLOGY TEACHER, VOLUME 62, NO. 8, OCTOBER 2000
Figure 1. Fourteen living and 57 fossil Caminalcules. A number is used to identify each species in lieu of a name.
CLASSIFICATION OF CAMINALCULES 571
Figure 1. (Continued).
Most students want to put 2, 3, 4, 12 and 22 together dents consider characters such as color pattern, body
(Figure 1). If so, I ask them to split the five species shape, presence of elbows and head ornamentation,
into smaller genera. The most common mistake at they quickly decide that 3 and 4 belong in one group
this point is to put 3 and 12 in a genus by themselves (genus), and 2, 12 and 22 belong in another. Some
because they both have claws. This provides an would put 22 into its own genus, which is acceptable.
opportunity to point out that the classification should Having classified the five species to everyone’s satis-
be based on all available characters. When the stu- faction, we then discuss convergent evolution, using
572 THE AMERICAN BIOLOGY TEACHER, VOLUME 62, NO. 8, OCTOBER 2000
Figure 2. Example of a hierarchical classification of the living Caminalcules. If one looks at the true phylogenetic tree in Figure
2 it is apparent that this is not the best classification scheme. Specifically, Genera 3 and 4 are more closely related to Genus
5 than they are to Genus 2. Without first looking at the fossil evidence, however, students are much more likely to come up
with something like this.
3 and 12 as an example. (The two Caminalcule example, would suggest the tree in Figure 3. One
cyclopes, Species 1 and 16, are also convergent.) I of the key concepts here is that of the most recent
point out that convergent evolution is said to occur common ancestor. Students learn that when they put
when a similar trait evolves independently in two two species, say 19 and 20, in the same genus this
separate lineages. For example, fish and whales both implies that these species share a common ancestor
have the same shape, but based on their skeletal not shared by other genera (the phylogenetic principle
structure, endothermy, lactation, etc., whales clearly of classification). When there are three or more species
belong with the mammals. The characteristics they in a genus, students must decide which two of those
share with fish evolved independently as an adapta- species are most closely related (Figure 4).
tion to an aquatic environment. Another way to The same procedure is applied through the higher
identify convergent evolution, though not available classification levels. For example, if two genera resem-
to the students until later in the lab, is to determine ble each other more closely than they do other genera,
if the character in question was absent in the most this is presumably because they share a unique com-
recent common ancestor. This is equivalent to saying mon ancestor. Thus, students learn that even in the
that the characters are analogous rather than absence of a fossil record it is possible to develop
homologous. a tentative phylogenetic tree that corresponds to the
I encourage the students to be taxonomic ‘‘splitters’’ classification scheme. It is not necessary that the
rather than ‘‘lumpers’’ for heuristic reasons; it makes phylogeny exactly match Camin’s true one, and it
it easier to introduce the concept of convergent evolu- is unlikely that it will, given the intuitive approach
tion as well as to emphasize the need to examine used here. The students can discover any errors for
all available characters very carefully. In contrast, themselves after completing the final exercise.
Sokal (1983) lumps species 2, 3, 4, 12 and 22 into
one genus.
A Phylogenetic Tree Based on the Fossil
Record
A Phylogenetic Tree Based Only on
For this exercise each group of two to four students
Living Species needs a large sheet of paper, scissors for cutting out
In the second exercise students use their classifica- Caminalcules, a meter stick for drawing lines on the
tion of the living Caminalcules to construct a phyloge- paper, and glue for attaching the Caminalcules. For
netic tree. The classification shown in Figure 2, for paper we use end rolls which the local newspaper
CLASSIFICATION OF CAMINALCULES 573
Figure 3. This phylogenetic tree is based on the classification of living species shown in Figure 2. The members of each genus
share a common ancestor not shared by other genera. The same is true for each of the four families and two orders.
species (58 and 74) represented by 18-million-year-
old fossils. The students continue the tree from there.
To make the exercise more manageable, I use a subset
of the original Caminalcules. I pruned some of the
branches from Camin’s original tree, leaving 14 living
and 47 extinct species (Figure 5). For the complete
set, see Sokal (1983). Note that some species are
represented by both living and fossil specimens.
Students enjoy this exercise and generally do a
good job of piecing together Caminalcule evolution.
There are, however, several pitfalls that may catch
even the most careful among them. These pitfalls
provide an excellent opportunity to discuss concepts
such as gaps in the fossil record and evolutionary
Figure 4. When a genus is made up of three (or more) species stasis (Figure 5). For example, when they get to
students must decide which two of the species share a com- Species 67 (12 million years old), most students
mon ancestor not shared by the other. This diagram indicates
assume it must have branched off from Species 30
that Species 2 and 12 are more closely related to each other
than either is to 22. We hypothesize that 2 and 12 have a (13 million years). This would require an unlikely
common ancestor (y) that is not shared by 22. scenario in which an evolutionary trend towards
heavy crushing claws is suddenly reversed to give
rise to what look like forked tentacles. Once this is
gives away or sells very cheaply. We cut the paper pointed out, students will deduce correctly that 67
into sheets about 28 inches on a side. branched off further down and that there is a gap
Each fossil Caminalcule (Figure 1) is identified by in the fossil record.
its species number and its age (in millions of years) Once their tree is complete the students compare
in parentheses. Make sure the students do not cut it to Camin’s phylogeny and reconcile any discrepanc-
off these numbers. Since the oldest fossil (Species ies. Then I ask them to identify, either in lab or as
73) is 19 million years old, students draw 20 horizon- an assignment:
tal lines on the sheet of paper and label them from 1. The most recent common ancestor of any two
0 (present time) at the top to 19 at the bottom. I species
usually show the class how to begin the phylogenetic 2. Additional examples of convergent evolution
tree by placing Species 73 in the middle of the 19 3. Examples of vestigial structures (e.g. the inner
million-year line. This species gave rise to two new toe of Species 66 at 3 million years)
574 THE AMERICAN BIOLOGY TEACHER, VOLUME 62, NO. 8, OCTOBER 2000
Figure 5. The Caminalcule evolutionary tree. Adapted from Sokal (1983). Some of the branches in the original tree have
been removed.
CLASSIFICATION OF CAMINALCULES 575
4. Examples of evolutionary stasis (e.g. Species 14 instructor monitor each group’s progress in order to
and 13 remain unchanged for 13 million years) identify problems as they pop up. This provides an
5. An example of rapid adaptive radiation (e.g. opportunity for the instructor to discuss with each
the many lineages that arise from Species 43 at group some of the concepts mentioned above.
7 million years). Additional information about the Caminalcules and
a copy of my laboratory exercise are available at
These observations can easily be applied to real world
http://www.iup.edu/;rgendron. Modern theories of clas-
examples and lead to interesting discussion questions
sification and phylogenetic analysis are covered in
such as:
more detail in evolution texts such as Ridley (1996).
1. How do vestigial structures provide clues about
a species’ evolutionary history? Acknowledgments
2. What ecological conditions might result in the
rapid diversification of some lineages (e.g. the The Caminalcules were developed by the late
mammals at the beginning of the Cenozoic) or Joseph Camin and published by Robert Sokal. I am
the long-term stasis of others (e.g. horseshoe grateful to Dr. Sokal and the editors of Systematic
crabs and other ‘‘living fossils’’)? Biology for permission to reproduce the Caminalcules
3. Some caminalcule species became extinct. What here and on my Web site. I would like to thank the
factors might increase or decrease the probability many faculty and students who have contributed to
of extinction in the real world? the development of this lab over the years. Thanks
also to B. Forbes, S. Forbes, J. McElhinney, T. Peard,
Teachers should also use the tree to emphasize the S. Vogel and an anonymous reviewer for critiquing
important principle that evolutionary change occurs the manuscript.
through the modification of pre-existing structures.
We have used this lab in our nonmajors’ General
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