A Simple Model Illustrating Problems of Phylogeny and Classification John M. Burns Systematic Zoology, Vol. 17, No. 2. (Jun.

, 1968), pp. 170-173.

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A SIMPLE MODEL ILLUSTRATING PROBLEMS OF PHYLOGENY AND CLASSIFICATION

Abstract
A model consisting of 37 pieces of hardware, representing as many animal species in a single phylum, has been used successfully at several levels of undergraduate and post-graduate experience to teach problems of beta taxonomy and the arbitrariness, subjectivity, and limitations of higher classifications. Students find this model challenging, stimulating, and thoroughly engaging. The model is simple, durable, inexpensive, and easy to manip-ulate; it fits comfortably in one laboratory penod; it involves a minimum of characters; and it presupposes no familiarity with the morphology and accompanying jargon of any particular group of organisms.

lecture or discussion of relevant Although my chief systematic interests lie at and around the species level, this paper reports a laboratory exercise con-cocted in January 1963 to illuminate major problems commonly encountered in the synthetic taxonomic process of delimiting higher categories. Students are often at best but dimly aware of problems at this level and, in particular, of the arbitrariness, subjectivity, and limitations of higher clas-sifications. Classifications themselves are undeniably useful; unfortunately their utili-zation on the part of many students is virtually blind. 1 designed the following exercise for a single laboratory period of the introductory biology course at Wesleyan University, primarily to provide critica1 insight into various aspects of beta taxonomy but, secondarily, to offer some relief and mind-expansion in a largely cellular-molecular undergraduate curriculum. The exercise has been well received in five years of use in introductory biology. Moreover, it has pro-voked interest and enthusiasm at other levels of biological sophistication: from undergraduates in science courses for non-science majors at Wesleyan; from graduate students in a course in speciation at Mountain Lake Biological Station of the University of Virginia; and from college and university teachers in a summer institute for teachers of biology at Williams College. 1 usually preface the exercise with an exploratory

them in an orderly hierarchic scheme that more or less features of taxonomic theory and append a moderatereflects these relationships. reading assignment to assure an adequateAssume that al1 of these background. Despite a few dif-ferences of opinion animals belong to a single on specific points, 1 still favor the terse account of phylum and limit your taxonomic prin-ciples on pages 1 to 24 in Simpson classification of them to the (Bull. Amer. Mus. Nat. Hkt . , 85, 1945: 1-350). following taxonomic The exercise is in two parts, of which the first is categories: class, order, by far the more important. family, genus, species. (Avoid finer subdivisions PART 1. You will receive an envelope (markedsuch as superorder or "Classification") containing 37 assorted pieces ofsubfamily. ) small hardware. ( D o not lose any of these pieces or Some species may be mix the contents of any two envelopes.) Each objectconsidered more primitive than others and perhaps represents a different species of animal. [See Fig. l.] directly ancestral to others. Study these "organisms" carefully, compar-ing each (Hence some of the species with every other species, to detect the"specimens" with which you similarities and differences among them. Use al1are working available "taxonomic characters" to work out the relationships between the organisms and to arrange

A SIMPLE MODEL ILLUSTRATING

PROBLEMS

171

FIG.l.-A

typical first look at the 37 "species" of this model.

have, and so forth. may be "fossils" rather tlian recently col-lected and preserved specimens of living species.) You will have to make considerable use of linear measurements as charac-ters. Assume that appreciable intraspecific variation is non-existent. After contemplating the entire problem before you, prepare a formal presentation of your interpretations in the following way: 1. Make a phylogenetic diagram of al1 the species. 2. Make a classification embracing al1 of the species. In writing out this classifi-cation, give the characteristics of each taxonomic group. 3. Steps 1 and 2 are intricately related. Justify both your phylogenetic arrange-ment and your classification by explain-ing, for example, why you have judged some forms to be more primitive and others more advanced, why you have seen fit to make the groupings that you

Students are often astounded when they first 4. Discuss the following questions briefly: 1. How is phylogenetic relationship pour out their hardware (Fig. 1 ) . Some laugh. A few may in-ferred from morphology? stare blankly at the parts for 3. What difficulties arise in the process of minutes on end. Such individtranslating a phylogenetic diagram into a u a l ~may need comments or classification? questions to get started. But, c. 1s information lost in this process? in general, the best proce-dure d. What arbitrariness, if any, is inheris (1) to leave the students ent in the practice of modern evolualone, letting each pore over tionary taxonomy and classification? his hardware and ponder his 5. Construct a dichotomous key (which can be artificial rather own phylogenetic arrangement than natural) allowing ready and rapid identification of and the tentative groupings he each species. would make

condition) but also drives home the wavs in in shifting from phylogeny to classification; which information is lost in going from an and then ( 2 ) to encourage the students to evolutionary diagram to a hierarchic compare notes. They are usually surprised at classification. Students grapple directly with how much their interpretations differ and such conflicting force-S as splitting versus frequently get into heated but healthy lumping and, more important, ver-tical versus horizontal classification and are often severely argurnents. lik distressed when they recog-nize that in some They may disagree strongly-much e instances they cannot avoid rather arbitrarily taxonomistswha placing related species in different major professional about t is groups. They see that even if two students are primitive and what advanced (and what agreed on a phylogeny, they may yet produce these terms mean ,) ,, about what direction difSYSTEMATIC ZOOLOGY sequence takes (e.g., small an apparent to large, or the reverse), about what ferent but valid classifications consistent with could that phylogeny. Some students want to discard conceivably give rise to what-in short, about the screw eye. Some of those who perceive most of the relationships they are that the screw eye will not readily fit in their trying to determine. They may ask-at phylogenetic scheme are brought face to face times. in real anguishu some with the concept of convergence: they find it Are characters easiest (but still not altogether satisfying) to more important than others? (e.g., Are difpostulate in-dependent origin of threads in the ferences in color or size, say, as trivial as they screw eye and the screws. Most students find often appear to be?); 1s it true with Ijving the exercise challenging and imaginative. organisms, as it seems to be here, that what are good characters in one group are not They usually emerge from it more appreciative of the difficulties of practicing necessarily very helpful in another? taxonomy, somewhat disillusioned with and Some students are disturbed to learn that bothered by classification, aware of many of there is no one correct solution, that the the limitations of our system, and prepared to exercis i a n ~ t h i n black e s gbut and white, take classifications, in future, with a grain ( at and that (given the information in the least ) of salt. "svecies" an Over and above this, the model has a various d the rules of the number of purely practica1 considerations to model) many interpretations are accept-able. Most realize that, despite this, there arerecommend it. In contrast to sets of real organisms, it is simple, uniform, compact, numerous arrangements and groupings that are ~ l a i n l vindefensible. Almost al1 durable, cheap (even for a large class), quick to assemble, and easy to handle; and lost come to discern a series of parallels between pieces are immediately replaceable from a the model and the biological situation. ready supply of spare parts. It re-quires no The model not only emphasizes the dif-knowledge of morphology and anatomical ficulties and differences of opinion amongterminology of any particular animal group. workers in establishing a phylogeny with And the number of charac-ters involved is incom~lete data at hand lthe standard
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limited, which simplifies and shortens theA SIMPLE MODEL ILLUSTRATING exercise without in any way vitiating itsPROBLEMS instructive purpose. The essence of the exercise is Part 1, of which section 5 (dealing with formation of a key) is tangential and therefore expend-able. Part 11 is also inessential, but less so in that it provides a partial antidote to one of the devices of Part 1.
PART 11.

In Part 1 you were asked to assume that appreciable intraspecific variation is nonexistent. This-like many other features of this exercise-is a huge oversimplification. un - ward along the Take one of the envelopes marked "Samordinate. ples." It contains samples of 21 "adult males" 1 3 2 5 Place the resulting 4 of each of two "species" collected at the same LENGTH, histograms one above time and place. (Individuals of both species are FIG.2.-Variation in "body" the other. [See Fig. 2.1 represented here by wires; color of the wire of each of two "species" of what basis, other On wire. serves to distinguish N.= 21 and % = 3% cm. than color, can you distinguish the two the two species.)species? If you had Assume that eachonly one or two sample pelfectly specimens of each reflects the populationspecies, could you te11 from which it wasthem apart using this drawn and furthercharacter? Explain. If assume that eachthe environment in these two population waswhich species live should exposed throughout its undergo relatively rapid change, which develo~ment tospecies might have the bet-ter chance of identical surviving? Why? environmental conditions so that individual variation in Depalliment of the sample stems fromBiology, Wesleyan Uni-uersity, genetic variation in Middletoutn, population. Measure each Connecticut 06457. in to the both

nearest meter, record vour measurements, calculate the mean length for each sample, compare the two means thus obtained, and then plot the frequency distribution for each sample in the form of a bar graph, with length increasing to the right along the abscissa and number of individuals increasing
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