Plant Taxonomy: The Systematic Evaluation of Comparative Data

SECTION 1

The Meaning of Classification

Taxonomy is dynamic, beautiful, frustrating, and challenging all at the same time (fig. 1.1). It is demanding philosophically and technically, yet it offers intellectual rewards to the able scholar and scientist. It can be manifested in works of incredible detail as well as in logical and philosophical conceptualizations about the general order of things. It has strong implications for interpreting the reality of the world as we can ever hope to know it.

Because taxonomy has deep historical roots, the past is never escaped. This places an increasing burden upon practitioners to understand old and new material. The past must be dealt with for older results, and every new discovery must be digested and incorporated. As Constance aptly put it, My ideal taxonomist, therefore, must be very versatile indeed, and should preferably be something of a two-headed [i.e., two-faced] Janus, so that one set of eyes can look back upon and draw from the experience of the past, and the other pair can be focused upon deriving as much of value as possible from developments on the present scene (1951:230).

Taxonomy is a synthetic science, drawing upon data from such diverse fields as morphology, anatomy, cytology, genetics, cytogenetics, chemistry, and molecular biology. It has no data of its own. Every new technical development in these other areas of science offers promise for improved portrayal of relationships of organisms. This is a demanding aspect of taxonomy for a practicing worker, because it is virtually impossible to understand completely all of these different data-gathering methods, yet highly desirable to be able to master as many as possible. Furthermore, the accumulation of data and their interpretation never cease. Not only do new techniques of data-gathering provide more information that must be brought to bear on understanding relationships, but also these new interpretations reveal new taxonomic groups that must be understood and utilized. These are some of the reasons why taxonomy (and its parent discipline, systematics) has rightly been called an unending synthesis (Constance 1964), an unachieved synthesis (Merxmüller 1972), or even more poetically, the stone of Sisyphus (Heywood 1974).

FIGURE 1.1    An example of the challenges facing the plant taxonomist is shown dramatically by this bizarre landscape, which could represent an obscure area of the earth or perhaps even another planet, with completely new and different plant forms. If this scene were on earth, we would have considerable biological information on plants in general, e.g., modes of reproduction, structures, and functions, and a good background of ideas on how to proceed with classification of these groups based upon historical classificatory records. If on another planet, however, attempting a predictive classification of these forms would be unbelievably difficult, with nothing known about modes of reproduction, structures and their functions, mechanisms of evolution, or even what is an individual or population. This same type of overwhelming challenge was faced by plant taxonomists on this planet approximately 500 years ago. (From Lionni 1977, frontispiece)

CHAPTER 1

A Few Definitions

Classification, Taxonomy, and Systematics

Taxonomy has had various meanings over the past 150 years, and particular confusion with systematics has prevailed. Systematics no doubt was used very early as a casual self-evident term (Mason 1950:194) to refer to the ordering of organisms into rudimentary classifications. This activity has occurred ever since people have lived on earth (Raven, Berlin, and Breedlove 1971). The early documented use of the term systematics (as systematic botany) can be traced at least as far back as Linnaeus (1737a, 1751, 1754), and it has persisted to the present day although in modified form. Linnaeus (1737a:3) stated that we reject all the names assigned to plants by anyone, unless they have been either invented by the Systematists or confirmed by them. In 1751, he used the term (as botanico-systematici, p. 17) to refer to workers who carefully distinguish the powers of drugs (in plants) according to natural classes. He made the definition of a Systematic Botanist even more clear in his preface to the fifth edition of the Genera Plantarum:

The use of some Botanic System I need not recommend even to beginners, since without system there can be no certainty in Botany. Let two enquirers, one a Systematic, and the other an Empiric, enter a garden fill’d with exotic and unknown plants, and at the same time furnish’d with the best Botanic Library; the former will easily reduce the plants by studying the letters [i.e., features of diagnostic value] inscribed on the fructification, to their Class, Order, and Genus; after which there remains but to distinguish a few species. The latter will be necessitated to turn over all the books, to read all the descriptions, to inspect all the figures with infinite labor; nor unless by accident can be certain of his plant" (1754:xiii, 1787:lxxvi).

Books using the term systematic botany appeared thereafter (e.g., Smith’s An Introduction to Physiological and Systematical Botany 1809 and Nuttall’s An Introduction to Systematic and Physiological Botany 1827). Mason, although admitting the difficulty of establishing the place of its first use, ventured the opinion that systematics might possibly have even preceded it [i.e., the use of taxonomy] (1950:194) and gave Lindley (1830b) as the earliest reference.

A biologist interested in relationships during this early period mostly studied morphological features and accordingly grouped organisms into units. This ordering of organisms into groups based on similarities and/or differences was (and still is) called classification. This is a very old term going back to Theophrastus in the third century B.C. (see 1916 translation). The Swiss botanist, Candolle (1813), in the herbarium at Geneva, coined taxonomy (as taxonomie)¹ to refer to the theory of plant classification. It later became more generally used for the methods and principles of classification of any group of organisms and is still used basically in this way (e.g., Simpson 1961). From this point to the publication of the theory of evolution by means of natural selection by Darwin (1859), the two words, taxonomy and systematics, were regarded as synonyms, although the latter was used much more frequently. During this time, classifications were believed to reflect the plan of natural order created specially by God, and man was simply rediscovering the Divine Plan. Biologists engaged in these activities of classification were called interchangeably either taxonomists or systematists. Since Darwin’s time, systematists have not only continued their interest in classification, but also have attempted to understand evolutionary relationships among the groups so ordered. Furthermore, some systematists have become interested in the process of evolution itself, that is, in the mechanisms that produce the diversity. Consequently, a systematist today may study many different aspects of evolutionary biology that are far removed from the morphological investigations of a century ago. For a useful overview of themes and progress in plant systematics during the past half-century, see Stevens (2000a).

The basic methodology of modern systematics is outlined in table 1.1. Data are gathered from organisms and their interactions with the environment and used to answer questions about classification, phylogeny, and the process of evolution. Specific examples of systematic studies might be analyzing the patterns of adaptive radiation within a particular group of species, comparing DNA sequences for reconstruction of phylogeny, or investigating patterns of intra- and interpopulational genetic variation. A similar and equally legitimate viewpoint was presented by Blackwelder and Boyden (1952), who indicated three steps: (1) recording of data; (2) analysis of the data for making classifications; and (3) synthesis of these generalizations for insights on phylogeny and evolution. Wilson aptly pointed out:

most systematists by choice are not problem solvers in their method of working. It might be said that the perfect experimental biologist selects a problem first and then seeks the organism ideally suited to its solution. The systematist does the reverse. He selects the organism first (for reasons that are highly individual) and only secondarily chances upon phenomena of general significance. The chief value of his discoveries is that they are typically of the kind that would not be made otherwise. If the systematist has an ideal program, it runs refreshingly counter to the conventional wisdom: select the organism first, as a kind of totem animal (or plant) if you wish, then actively seek the problem for the solution of which the organism is ideally suited (1968:1113).

TABLE 1.1 Outline of Methodology of Systematics

I. Accumulation of Comparative Data

A. From the Organism

1. Structures

2. Processes (interactions among structures)

B. From the Organism-Environment Interactions

1. Distributionsa

2. Ecology

II. Use of Comparative Data to Answer Specific Questions

A. Classification (most predictive system of classification at all levels)

1. Method and result of grouping of individuals

2. Level in the taxonomic hierarchy at which the groups should be ranked

B. Process of Evolution

1. Nature and origin of individual variation

2. Organization of genetic variation within populations

3. Differentiation of populations

4. Nature of reproductive isolation and modes of speciation

5. Hybridization

C. Phylogeny (divergence and/or development of all groups)

1. Mode

2. Time

3. Place

a Floristics, or the documentation of what plants grow in particular regions, is deliberately not listed in this table as a separate question, nor does it find a specific place in the areas of systematics in figure 1.2. Determining where particular plants grow is a very legitimate and valuable activity within systematics, but it is essentially data-gathering of distributions of plant groups that have already been classified. Some floristic projects, however, especially of poorly known regions (e.g., Rzedowski and McVaugh 1966; McVaugh 1972a, b) involve considerable amounts of classification as well as original historical scholarship. To this extent, they become more revisionary, and less floristic, in character (for these and other distinctions, see Stuessy 1975). Many innovations in floristic work are presently occurring, especially using Web-based technologies (see symposium introduced by Kress and Krupnick 2006).

This is a very important perspective. Were all biologists interested only in seeking answers to basic biological processes, then all would work with model organisms such as Drosophila or Arabidopsis, both easily manipulated in the laboratory. This is certainly one approach to biological science, but if we do not continue to investigate other organisms, we will miss out on other interesting and useful processes (Greene 2005). Where would molecular biology (and molecular systematics) be without polymerases that work at high temperatures, which were found in the course of systematic studies on bacteria of thermal pools (Saiki et al. 1988). Even model-organism researchers themselves recognize the importance of adding other representatives of different life forms (Fields and Johnston 2005).

Because of the diverse nature of these studies that were spawned by evolutionary theory, a collective term was needed to designate these different activities and the people so engaged, and another term also was needed for describing the more traditional activities of classification. As a result, the term systematics (or systematic biology) has come to have a meaning different from and broader than taxonomy. The definition used by most people and preferred here is that of Simpson: the scientific study of the kinds and diversity of organisms and of any and all relationships among them (1961:7). A slightly simpler way of defining systematics is the study of the diversity of organisms (Mayr 1969c:2; cf. Wilson 1985, for his similar definition the study of biological diversity). Diversity is such an important concept in systematics that a useful and delightful perspective on this was provided by Constance:

Much as I respect the giant strides that have been made in clarifying basic principles and processes of wide applicability, I have chosen to celebrate diversity. It is well enough to know that all music can be reduced to a relatively few notes and a minimum of ways of evoking and receiving them in the human ear. This does not suggest to the music lover that symphonies, sonatas, and operas are redundant because their parts and processes can be analyzed. All literature, after all, is merely spun out of words. Human beings are a lot alike, but it does not necessarily follow that there is no point in knowing more than one of them (1971:22).

Myers provided a slightly more general definition of systematics: the study of the nature and origin of the natural populations of living organisms, both present and past (1952:106). The term biodiversity is now often used (e.g., Fosberg 1986; Wilson 1988; Minnis and Elisens 2000) to refer to the total biotic diversity on earth.

Some biologists have equated taxonomy with systematics (Lawrence 1951; Crowson 1970;² Radford et al. 1974; Ross 1974; Jones and Luchsinger l986; Stace 1980; Minelli 1993; Singh 2004);³ Wilson echoed this viewpoint and talked about a pure systematist, who can be defined as a biologist who works on such a large number of species that he has only enough time to consider classification and phylogeny. If he narrows his focus, his unique knowledge provides him with a good chance to make discoveries in genetics, ecology, behavior, and physiology, as well as in taxonomy. But then we come to know him as a geneticist, or an ecologist, or a behaviorist, or a physiologist (1968:1113). Swingle (1946) made a distinction between systematics and taxonomy with the latter regarded as dealing with phylogenetic classification, and the former being broader to include taxonomy and nomenclature. Mason (1950; and followed by Porter 1967) took a different approach and regarded systematics (specifically systematic botany) as the data-gathering aspect (e.g., as Systematic Anatomy, Systematic Cytology, Systematic Genetics) and taxonomy as the interpretive phase in constructing classifications and in revealing evolutionary relationships. Wheeler offered a cladistic definition of systematics as the production of cladograms that link taxa through their observed variation (2005:71), a much too limited definition in my view. Smith defined systematics as that branch of science which investigates the philosophy that underlies a classification (1969:5), which would be closer to the definition of taxonomy used in this book. Schuh followed a similar definition describing systematics as the science of biological classification (2000:3). Wägele talked about biological systematics as the science of the systematization of organisms and the description of their genetic and phenotypical diversity (= biodiversity) (2005:10). For general agreement with the definitions used here, see Danser (1950), Davis and Heywood (1963), Blackwelder (1967a), Sylvester-Bradley (1968), Mayr (1969c), Darlington (1971), Stuessy (1978b), Woodland (2000), and Judd et al. (2002). For additional discussion of these (and related) concepts, see Small (1989). The term synthetic biology has emerged recently (e.g., Benner and Sismour 2005), but this deals with creating artificial life and using different units of the natural world to form new functioning systems. There also exists the term syntaxonomy, but this refers to classification of plant communities (e.g., Fremstad 1996). The relationships among terms as used in this book are shown in fig. 1.2.

Because most fields have a set of principles, and because taxonomy is the main focus of this book, I herewith list the six points that to me seem significant for our field (from Stuessy 2006):

1. All known life originated on earth during the past 3.5 billion years.

2. Due to evolutionary processes (e.g., speciation, hybridization, extinction), life-forms show natural patterns of relationship to one another.

3. Using selected features of organisms (characters and states), we determine patterns of relationship that we assume reflect these evolutionary processes.

4. Humans need hierarchical systems of information storage and retrieval to live and survive, including dealing with the living world.

5. The assessed patterns of organismal relationship are used to construct hierarchical classifications of coordinate and subordinate groups that are information-rich and have high predictive efficacy; these are the taxonomic hypotheses that change with new information and new modes of analysis.

6. Names are assigned to the classified groups to facilitate communication about them.

FIGURE 1.2    Diagram of conceptual and procedural relationships among and within areas of systematics. The tighter connection between the bottom two areas emphasizes their closeness as aspects of evolutionary processes, short-term on one hand, and long-term on the other. (From Stuessy 1979b:622)

Every worker would have a slightly different set of principles, without a doubt, but to me these points seem fundamental.

Nomenclature

Two other terms, nomenclature and identification, are sometimes confused with classification and systematics. After groups of organisms have been classified, names must be given to these groups so that communication about particular units will be facilitated and so that continued progress in classification can be made (Hitchcock 1916). The naming of groups of organisms and the rules governing the application of these names together are called nomenclature. Nomenclature cannot, in fact, be wholly separated from the classification that it serves…. New taxonomic advances will always pose new nomenclatural problems, whatever form of taxonomy is employed, just as new social forms require new linguistic expression. New nomenclatural procedures will enable new possibilities in taxonomy to be formulated and examined. The process is fruitful, inevitable and in many respects essential to progress (Whitehead 1972:216). Many philosophical and methodological topics on nomenclature could profitably be discussed, such as using code numbers (numericlature; Little 1964; Hull 1966) or descriptive codes (sunegs, genus spelled backwards; Amadon 1966) instead of descriptive binominals, possible roles for uninominals (Michener 1963, 1964; Lanham 1965; Steyskal 1965), elimination of Latinization of names (Yochelson 1966), the numerous features of the established International Code of Botanical Nomenclature (McNeill et al. 2006), comparisons with the zoological, bacteriological, and viral codes (see Web sites respectively: www.iczn.org, www.ncbi.nlm.nih.gov, www.ictvonline.org), the proposed biological code for all organisms (BioCode; Greuter et al. 1998), and the proposed phylogenetic code (PhyloCode; Cantino and de Queiroz 2005). But these topics fall largely outside the scope of this book (for some comments, especially on the PhyloCode, see Chapters 8 and 10). The reader is referred to discussions in Lawrence (1951), Jeffrey (1977a), and Simpson (2006) for salient points of the Botanical Code; case studies are provided by St. John (1958), but without answers!

Identification

Identification, on the other hand, involves referring an individual specimen to a previously classified and named group (Jardine 1969a). For example, if one walks outside and picks a small branch with leaves from a tree, takes the specimen back to the laboratory and attempts to find a name for it, what is being sought is an identification of the specimen—not a classification. Many years ago, some taxonomist did classify the species represented by the individual tree. It is possible, of course, that the species of tree has never been classified before (unlikely in temperate countries, but entirely plausible in poorly known areas of the tropics), in which case, this would need to be done and an appropriate new name provided. Many innovations in identification have been suggested and discussed (e.g., Dallwitz 1974; Pankhurst 1974, 1975, 1988a; Sneath and Chater 1978; Mascherpa and Boquet 1981), especially information content of keys, multiple access keys (polyclaves, Hansen and Rahn 1969, Simpson and Janos 1974; Duke and Terrell 1974; Westfall, Glen, and Panagos 1986; Robert et al. 1994; these are also called tabular keys, Newell 1970, 1972, 1976; Götz 2001), use of artificial neural networks (Clark 2003), and use of computers for automated identification and key construction (Watson and Milne 1972; Malesian Key Group 2004; MacLeod 2007), including electronic field guides (Agarwal et al. 2006). The Internet is now becoming a location for placing these interactive keys (Wilson and Partridge 1986; Farr 2006). Details of these topics fall outside the scope of this book; see Pankhurst (1978) for a good introduction.

These two activities, identification and nomenclature, are directly associated with classification (fig. 1.2). For purposes of communication, groups of organisms without labels are not at all useful to humans, and similarly, without identification, the names given originally to the classified groups cannot be applied to newly collected individuals. These associated activities, therefore, are not only directly related to, but are also most important for, classification. Despite their importance, however, they must never be regarded as overshadowing the more significant effort of studying the organisms themselves and developing classifications to reflect relationships. Although Myers overstated the case, it is basically true that If systematics deals primarily with the nature of populations, such appurtenances as nomenclature are seen in proper perspective, as mere adjuncts to systematics, necessary in speaking of populations but of minor importance (1952:107). Labels are meaningless and unnecessary unless they refer to individuals or groups with information content, such as units within a classification.

Biosystematics

Biosystematics was introduced by Camp and Gilly (as biosystematy; 1943a:327; see also Camp 1951) as an attempt (1) to delimit the natural biotic units and (2) to apply to these units a system of nomenclature adequate to the task of conveying precise information regarding their defined limits, relationships, variability, and dynamic structure. Determining these natural units often involved a program of artificial crossing studies among populations. In this fashion, the isolating mechanisms and genetic compatibilities were revealed that led to discovery of the natural units. As a result of this definition, any studies involving breeding programs between taxa have been regarded as involving biosystematics, even though the data sought were for determining evolutionary relatedness of taxa rather than for discovering natural units. Böcher remarked that "In my opinion the place of biosystematics is closer to cytogenetics and ecology than to taxonomy in a narrower sense. Our main interest is not classification but evolution. This, of course, does not mean that we should never deal with classification. But frequently it will be better to leave problems of taxonomic rank and nomenclature to taxonomists sensu strictu, or to cooperate with them. The main goal of a biosystematist is to try to unravel the evolution of a group of taxa, what the evolutionary forces were and how they worked together in each particular case" (1970:3–4) (these sentiments also were echoed by Merxmüller 1970 and Stebbins 1970b). With the proliferation of other laboratory studies (including DNA) in systematics during the past several decades, however, sometimes biosystematics has been used to refer to any kind of experimental systematic study, i.e., involving any type of data-gathering except traditional morphology and anatomy. Further, because of the very broad definition of systematics now in use by most workers (and used in this book), the need for the term biosystematics has surely lessened and, in the minds of some, nearly disappeared (Johnson 1970). It is still used occasionally, however (e.g., Catling 2001; Lowrey 2002; Nieto Feliner et al. 2005), and in one instance as a synonym of systematics (Lecointre and Le Guyader 2006).

Experimental Taxonomy

Experimental taxonomy (or experimental systematics) is another term most often used for laboratory-based studies other than (or in addition to) crossing or breeding data (Hagen 1983, 1984). These are not experiments in the strict scientific sense, but rather the gathering and analyzing of different kinds of comparative data that ordinarily are generated in the laboratory (e.g., cytology, phytochemistry, DNA sequences, computer manipulations) and used to generate and test hypotheses (Gilmartin 1986; Donoghue 1987; La Duke 1987). The term experimental taxonomy did derive from actual experimental investigations of the nature of plant species by reciprocal transplants of clones into different environments (Gregor 1930, 1938; Hall, Keck, and Hiesey 1931; Hall 1932; Gregor, Davey, and Lang 1936; Clausen, Keck, and Hiesey 1939, 1941a), which effectively discriminated the genetic vs. environmental components of morphological variation. This was also called genecology by Turesson (1923) and others (e.g., Constance 1957). Although these valuable kinds of studies were continued (e.g., Müntzing 1969), experimental taxonomy has come to have a broader (and less precise) usage. Müntzing emphasized the importance of genetics and cytology in experimental taxonomy and commented that what experimental taxonomy can do is to establish the nature and occurrence of such intra- and interspecific differentiation that cannot be clarified merely by morphological, ecological and plant-geographical methods (1969:791). Rollins commented: The type of experimentation differs, depending upon the objectives. The most frequent are undoubtedly those associated with genetics and cytology in which the reproductive process or the level of polyploidy is under investigation. The most effective experimental approaches in taxonomy have combined work in the herbarium and field with studies in the greenhouse and experimental plot (1957:192). Some workers (e.g., Stace 1980) have regarded experimental taxonomy and biosystematics as synonyms.

New Systematics

The term new systematics was coined by Hubbs (1934) and popularized by Huxley to stress that Fundamentally, the problem of systematics, regarded as a branch of general biology, is that of detecting evolution at work (1940:2). That is, the focus is on understanding the mechanisms by which diversity is produced rather than solely on classification. This was a useful distinction at that time and helped strengthen the existing area of experimental taxonomy and aided spawning of biosystematics. The influence of Huxley’s book (1940) was substantial, and this is reflected in Mayr’s emphasis on populations and his explanation of the term (in contrast to the old systematics): "The new systematics may be characterized as follows: The importance of the species as such is reduced, since most of the actual work is done with subdivisions of the species, such as subspecies and populations. The population or rather an adequate sample of it, the ‘series’ of the museum worker, has become the basic taxonomic unit. The purely morphological species definition has been replaced by a biological one, which takes ecological, geographical, genetic, and other factors into consideration. The choosing of the correct name for the analyzed taxonomic unit no longer occupies the central position of all systematic work and is less often subject to argument between fellow workers. The material available for generic revisions frequently amounts to many hundreds or even thousands of specimens, a number sufficient to permit a detailed study of the extent of individual variation" (1942:7). Despite its utility at the time, however, new systematics is obviously a dated term and is rarely used today. Every age has had its own new systematics. As far as I am concerned, there always has been some new systematics and there always will be (Mayr 1964:13). As Sylvester-Bradley appropriately put it the ‘new systematics’ of today is something very different to the ‘new systematics’ of thirty years ago…. Perhaps the time has come to consider the publication of a ‘Newer Systematics’ (1968:176–177). There is now the truly new systematics (Schram 2004), which emphasizes the handling of large databases. Similar is comprehensive systematics (Stuessy and Elisens 2001), which utilizes different types of data and the computer to help reveal relationships.

The label new taxonomy was used by Cain (1959a) to refer to anticipated advances in making taxonomic comparisons by more quantitative means. This hope did not materialize in the way Cain envisioned, but the development of what eventually was called numerical taxonomy has yielded many useful results (see Chapter 7). Cain also introduced the term cryptic taxonomy (1959b) or cryptotaxonomy (1962) to refer to taxonomy in which the exact features used for comparisons have not been made explicit (i.e., most of the traditional intuitive approaches). The new taxonomy was meant to remedy this.

Comparative Biology

Comparative biology, also sometimes called the comparative method (Fisher and Owens 2004), is a term similar to systematics and regarded as synonymous by some (e.g., Nelson 1970). I view it here as broader, embracing any study that compares particular features of organisms. For comparative biology to be equivalent to systematics involves the asking of questions only about classification and/or evolution (table 1.1). But other very different questions also are sometimes asked that utilize comparative data for answers, such as in genetics, physiology, or descriptive and developmental anatomy. The focus of these studies is simply descriptive of form and/or function and not interpretive in the context of evolutionary relationships. It is probably true that the most meaningful questions answered with comparative data are, in fact, systematic ones, which may be one of the reasons why some workers (e.g., Mayr 1969c) regarded comparative biology as falling completely within systematics. Another and more broad approach was taken by Nelson and Platnick (1981), who regarded comparative biology as the science of diversity (1981:5), which includes the primary areas of systematics and biogeography and also the secondary areas of embryology and paleontology.

¹Some workers, e.g., Richter (1938), believed that taxonomy, if properly derived from its Greek roots, should be spelled taxionomy (or even taxinomy), but these suggestions for change were unfounded and unnecessary (Mayr 1966; Pasteur 1976) and have never been adopted.

²Although Crowson concluded that The words classification, systematics and taxonomy are now commonly treated as synonyms, an example of the confusion and carelessness in the use of words which is prevalent in so much modern writing (1970:18), he offered the view that Originally and properly, classification would have denoted the activity of placing things in classificatory groups, whereas systematics would be the body of general theory underlying this activity (p. 19).

³Griffiths (1974a) suggested that because systematics and taxonomy have been regarded by most authors as synonyms, the term metasystematics should be used for the more inclusive concept. To my knowledge, no one else has yet adopted this terminology. It would now be even more confusing due to use of the term metadata in systematics, i.e., comparative data associated with taxa, such as DNA sequences or chromosome numbers.

CHAPTER 2

The Relevance of Systematics

Because of the close relationship of taxonomy to systematics, a few comments on the relevance of the latter are appropriate. It would be surprising if systematics, which deals directly with organic diversity, did not relate to every aspect of human endeavor. The impact of systematics upon society is substantial and most easily reflected in satisfying our intellectual curiosity about the world in which we live, formulating principles and methods of classification applicable to many human needs and activities, helping preserve the world’s organic diversity for aesthetic and economic reasons, and more directly in developing economic potentials.

Importance of Systematics in Society

There is no general concern more important for an individual during his or her lifetime than coming to grips in some manner with how life came to be. Every person deals with this question in a different way; it is a highly individualized business with different answers approximating the number of questioners. But no matter how one addresses the problem, nor how one offers solutions, four general concerns nearly always come to mind: How did life originate?

How has life changed through time? What life now exists on earth? What were the origins and evolution of humans? Answers to these basic questions about the human condition all come from systematic biology. As Smith stressed, It is because the systematist has been able to remove himself from the fragment of earth time that is man-dominated, in order to view this planet and the prior phases of atomic evolution from an effectively external philosophical viewpoint, that he is particularly fitted to appreciate reality (1969:7–8). Insights from other disciplines are obviously involved, too, such as anthropology, biochemistry, geology, and genetics, but systematics is central to these issues. Systematics is involved with studies on the origin of life (e.g., Eigen and Winkler-Oswatitsch 1983; Hartman, Lawless, and Morrison 1987; Brack 1998; Schopf 2002; Thomas et al. 2006), with investigations of the evolution of life once it was created (e.g., Chaloner and Sheerin 1981; Willis and McElwain 2002), with studies of the biota on earth including the proper storage and documentation of all collected materials (e.g., Cohen and Cressey 1969; Banks 1979; Krupnick and Kress 2005), and with all kinds of morphological, anatomical, cytological, and biochemical investigations on the appearance and spread of the human species (e.g., Brown et al. 1982; Blumenberg and Lloyd 1983; Sibley and Ahlquist 1984; Rak 1985; Smouse and Li 1987; Carroll 2003; White et al. 2003; Macaulay et al. 2005; Mellars 2006). Systematics is truly fundamental for satisfying our intellectual curiosity about the nature of the world in which we live.

These considerations have become even more pertinent with the recent detailed exploration of our neighboring planet Mars. All sorts of amazing discoveries have occurred, but much attention has been placed on a ca. 4 billion-year-old meteorite given the accession number ALH84001 that was ejected from the surface of Mars ca. 15 million years ago and fell to earth in Antarctica ca. 11,000 years ago. McKay et al. (1996) reported small bacteria-like structures and organic signatures inside this meteorite. This was followed by reports of chains of magnetic crystals in this same sample (Friedmann et al. 2001; Thomas-Keprta et al. 2001) that resemble modern magnetite-chain-forming bacteria. Not all workers are convinced, however, that these chains can be explained only by biotic origins (Barber and Scott 2002). In addition, Weiss et al. (2000) estimated that the internal temperature of a meteorite of this nature, particularly as it was penetrating the earth’s atmosphere, would not have been above 40°C, easily within survival range of modern species of bacteria. Water ice has now definitively been documented on Mars (Bibring et al. 2004), as well as indications of possible past oceans (Head et al. 1999) and fluvial (Malin and Edgett 2003) and glacial (Head et al. 2005) geomorphic signatures. Whether this all means that present life on Earth originated from Mars, or evolved independently in parallel, or led to frequent early cross-dispersals (these would be cases of really long-distance dispersal!), remains to be seen. To emphasize once again, these are all questions that fall clearly within the purview of systematic biology.

No one in past decades can have failed to realize that the environment is of special concern for the continued survival of our species and for the maintenance of a desirable quality of life in our society. Our present environmental crisis has included recent debates over global warming and climate change (Thomas et al. 2004; Araújo and Rahbek 2006), but so far, there has been much more hand wringing than solutions proposed and accomplished. At the very least, it is obvious that as the human population increases and absorbs energy resources for its own maintenance, fewer total useful resources will be available on earth for future human needs and fewer will be available for most other species as well (Birdsall, Kelley, and Sinding 2001). In short, severe pressure is being brought to bear on all species in a most serious competition for survival (Prance and Elias 1977; Jenkins 2003; Hawksworth and Bull 2006). Tropical forests, which hold more undescribed species than anywhere else, are especially under pressure from deforestation and cultivation in developing countries of the world (Croat 1972; Gómez-Pompa, Vazquez-Yanes, and Guevara 1972; Raven 1976; Adams, Dong, and Shelton 1980; Myers 1980; Campbell and Hammond 1989; Prance 2001; Brook, Sodhi, and Ng 2003; Curran et al. 2004). As these habitats disappear, so do the species that are unique to them. As species disappear, so does their potential to aid our future needs, and we are left with one less weapon in our dwindling arsenal for survival. Further, most human beings are tremendously susceptible to the aesthetics of their environment to the extent that as diversity decreases and the world of our individual experience becomes more monotonous, its mentally therapeutic value declines proportionately with as yet poorly understood impact on the human condition. This relates to biophilia, our natural bond with other species (Wilson 1984; Kellert and Wilson 1993).

The systematist’s roles in the face of this loss of diversity are obvious: speak out loudly and clearly on conservation issues in which an informed opinion will be helpful (Mosquin 1971) and work for preservation of natural areas (it is far more economical in the long run to preserve a habitat that houses a rare and desirable species [in situ conservation] than to attempt to maintain it artificially apart from that habitat [ex situ conservation]); collect and inventory vigorously in those areas of immediate danger of destruction (Turner 1971); and help set aside germplasm resources to be saved for future breeding studies (Hawkes 1978), especially wild relatives of crops already of great economic value (e.g., wheat, rice, corn, soybeans, sunflowers). We also need long-term conservation perspectives (Willis and Birks 2006), which are particularly challenging due to regularly changing democratically elected governments in the developed world, and better prioritization of global conservation efforts (Brooks et al. 2006; Wilson et al. 2006). Elimination, or at least amelioration, of poverty also relates to successful initiatives for biodiversity conservation initiatives (Adams et al. 2004), as does reduction of corruption (R. J. Smith et al. 2003; Laurance 2004). Hedberg stressed: In a world with rapidly increasing human population pressures and accelerating exploitation it is imperative to utilize biological resources sagaciously on a sustained yield basis, and to this end we must have an adequate knowledge of its flora (1978:7). Many species yet to be described will have enormous value for food and medicine, and these are often encountered serendipitously in the course of general floristic work or in field work primarily devoted to other purposes (Iltis 1982). Systematics is essential for helping ensure our continued survival on this planet (Forey et al. 1994; Vane-Wright 1996; Leadlay and Jury 2006).

There is very good recent news with regard to systematists and conservation in that for the first time in history, the systematic biology community has come together to support a single top priority: to provide a complete biotic inventory of the planet, the well-known Systematics Agenda 2000 (Anonymous 1994). However remarkable this written consensus is, still lacking is translation of community consensus into funded reality, such as achieved by astronomers or high-energy physicists with their pieces of equipment with multimillion- or billion-dollar price tags. Positive steps toward implementation can be seen with the ALL Species Foundation project (Smith and Klopper 2002; Boom 2005), which aims to inventory all life forms within approximately 25 years. There is also now the Web-based Encyclopedia of Life, which is user-contributed and so may have a better chance to succeed.

Despite centuries of systematic work, we still know perhaps only 80 percent of the seed plants, 5 percent of the fungi, and an even smaller percent of the microbial world. We are still quite clearly on a new voyage of discovery of our own planet (Donoghue and Alverson 2000; Prance 2001; Brooks and McLennan, 2002). There is even considerable disagreement on the number of seed plants that inhabit the earth, i.e., the dominant vegetation, with estimates ranging from a low of 223,300 (Scotland and Wortley 2003) or ca. 260,000 (Thorne 2002) to more than 420,000 (Govaerts 2001, 2003; Bramwell 2002).

Clearly much more collecting is needed (Prance 2001, 2005), as is more monographic work (Stuessy 1993; Helgason et al. 1996; Hopkins et al. 1998; Kirschner and Kaplan 2002). Considerable attention has been given to the taxonomic impediment (Environment Australia 1998), i.e., the lack of trained personnel to get the job accomplished. While the lack of human resources is certainly an issue, employment of parataxonomists, those with some training, may provide a solution (Basset et al. 2000). Likewise, U.S.A. governmental funds have also been channelled successfully toward large training programs in monographic systematics (the PEET program; Rodman and Cody 2003). Certainly greater use of the Internet in making information on existing biodiversity more readily available is also to be encouraged (Bisby 2000; Godfray 2002; Wheeler 2004).

As a result of the need to inventory the planet, especially considering the current high rate of loss of biodiversity, some workers (Blaxter 2003; Tautz et al. 2003; Pons et al. 2006) have suggested completing a more rapid DNA inventory rather than a relatively time-consuming, normal, taxonomic approach of defining and describing new species based largely on morphological features. As one example, Fuhrman and Campbell (1998) found DNA sequences from deep-sea samples that were 30 percent different from any known organism. As another example, Venter et al. (2004) filtered several hundred liters of sea water from the Sargasso Sea, not known for its microbial diversity, and used whole-genome shotgun sequencing to reveal the existence of microorganisms. The results yielded 148 previously unknown phylotypes. Others have echoed the need for similar microbial assessments (DeLong and Pace 2001; DeLong et al. 2006; Mering et al. 2007; Not et al. 2007). Positive suggestions for inventorying fungi in the soil have also been expressed (Gewin 2006). Although such efforts tell us next to nothing about the organisms themselves, i.e., about their morphology, reproduction, life processes, and ecology, they can show levels of genetic diversity within a particular ecosystem. Some overly enthusiastic workers, however, have even advocated establishment of classifications based primarily on DNA sequences (Tautz et al. 2003; Blaxter 2004), but not surprisingly, not everyone agrees (Seberg et al. 2003). Moritz (2002) and Ennos, French, and Hollingsworth (2005) have properly stressed the importance of thinking not just about conserving taxa (or structures) but also about evolutionary processes, especially in dynamically changing taxa that provide us with difficult taxonomic boundaries.

Systematics can also help in developing further the economic resources that we already have. Biological control of agricultural pests, especially insects, has been used for decades with frequent success. For such endeavors to work well, systematists must be involved with proper identification of the organisms plus supplying data on their ecology and reproductive habits (Clausen 1942; Sabrosky 1955; Rosen 1986) to avoid unanticipated, unwanted, and economically ruinous results. Proper identification of plant materials is also important in customs work, as well as forensic applications (Coyle 2004). The proper use of land resources such as the building of new dams, new canals, and strip mines, is another area in which systematists play an indispensible role by advising on the possible ecological impact on organisms living in the region (Hedberg and Hedberg 1972). Further, the knowledge and techniques gained by systematists through study of relationships of wild species can often be used to improve our existing, cultivated, food crops by similar methods of cytogenetics, molecular biology, and artificial selection. Gathering of data on indigenous uses of plants (e.g., Duke and Vasquez 1994; Austin 2004) can also yield valuable economic results, in consort with acceptable profit sharing.

Systematics is also important in an even broader, but less obvious, philosophical way. This perspective was summarized well by Mayr:

The study of diversity has perhaps made its most important contribution to the development of new human conceptualizations, to a new approach in philosophy. More than anything else, it is the study of diversity which has undermined essentialism, the most insidious of all philosophies. By emphasizing that each individual is uniquely different from every other one, the students of diversity have focused attention on the importance of the individual, and have developed so-called population thinking, a type of thinking that is of the utmost importance for the interaction of human sub-groups, human societies, and human races. By showing that each species is uniquely different from every other species and thus irreplacable, the student of evolution has taught us a reverence for every single product of evolution, one of the most important components of conservation thinking. By stressing the importance of the individual, by developing and applying population thinking, by giving us a reverence for the diversity of nature, systematic and evolutionary biology have supplied a dimension to human conceptualization which had been largely ignored, if not denied, by the physical sciences. And yet it is a component which is crucial for the well being of human society and for any planning of the future of mankind. (1974a:8–9)

Contributions of Systematics to Biology

The importance of systematics not only extends generally to society, but also more specifically to other areas of biology (Mayr 1968a, b). One of the most pertinent contributions is the role that taxonomy plays as the data processing system for biology or in a less eloquent phrasing as the digestive system of biology (Heywood 1973a:139, 143). The number of data points that are being collected from organisms every day in biological research laboratories throughout the world is overwhelming. Literally millions of pieces of data are being gathered in the course of studies ranging from the genetic to the anatomical. Most of these data are tabulated and reduced in some fashion to answer specific questions posed by the biologist. In addition to helping answer these questions, however, these data can be used to assess relationships among organisms. Thus, the isolated pieces of information can be used profitably in a more general way. Although this generality is true, the data so collected are rarely complete by themselves for making systematic inferences; much additional study is almost always needed to develop truly comparative data for all the organisms under study by the systematist. In fact, even the data generated and used in traditional floristic and revisionary work is usually deficient in some respects (Watson 1971; Heywood 1973a; Stuessy 1981; Pullan et al. 2005). This data processing or digestion also allows for future specific questions to be asked that are more sophisticated and meaningful than past questions. The organization of data is enhanced by its attachment to organisms that are arranged hierarchically in a system of classification. We are engaged in the construction of a framework on which to hang or arrange the total available biological information, the data about life, whether on the molecular or the organismal or the population levels (Fosberg 1972:632). The informational content of a hierarchical arrangement of data is greatly superior to that of an arrangement in which data are all coordinate to each other.

A further step in making information about organisms more conveniently available for biological needs is through DNA barcoding (Hebert et al. 2003; Blaxter 2004; Hebert and Gregory 2005; Savolainen et al. 2005; Schindel and Miller 2005; Cowan et al. 2006). This initiative strives to assign a specific DNA sequence for all life forms. Clearly, the same sequence cannot possibly serve to distinguish all species in all groups, but the idea is to use one sequence within a particular group (e.g., a different one for Bacteria, Asteraceae, or Bryophyta) or even possibly combining different short sequences (Rubinoff, Cameron, and Will 2006). If successful and if widely used, DNA barcodes would have a marked positive impact on allowing more rapid and precise identification of organisms for ecological and many other studies. It would also facilitate prospecting for new biodiversity. Not all agree with these presumed benefits, however (Wheeler 2005; Brower 2006; Cameron, Rubinoff, and Will 2006; Hickerson, Meyer, and Moritz 2006; Meier et al. 2006), emphasizing that there is no substitute for a full understanding of organismic diversity (Ebach and Holdrege 2005; Will, Mishler, and Wheeler 2005) plus stressing the proven value of morphology for identification and classification (Will and Rubinoff 2004).

Systematics also helps us understand the processes of evolution, which is information used by many other areas of biology. The microprocesses of evolution, including individual variability, population variation, reproductive isolation, and modes of speciation, are all revealed through systematic studies. Some workers might prefer to call these kinds of investigations evolutionary biology or even more specifically, e.g., reproductive biology, population genetics, or speciation biology, but they all clearly fall within systematics as broadly defined in this book. The populational data are used by areas such as genetics, developmental biology, and even more distant subdisciplines such as game theory. Broader-scale evolutionary phenomena, sometimes called macroevolution (Stanley 1979; Vrba and Eldredge 2005), are also revealed through systematic studies, for example, trends in the specialization of seeds and seedlings and many other reproductive features of flowering plants (Stebbins 1967, 1970a, 1971a, 1974). These broader insights are likewise useful for other areas of biology (e.g., anatomy, morphology, and developmental genetics in the example of seeds and seedlings given above). Macroevolution has many different meanings and is controversial. Some workers believe that phenomena occur over long periods of time that are distinct from populational processes (Stanley 1979), and others (including myself) believe that they are accumulations of effects developed at the population and species levels (Stebbins 1975; Bock 1979; Charlesworth, Lande, and Slatkin 1982; Coyne and Orr 2004) plus extinctions (Nitecki 1984; Novacek and Wheeler 1992; Steadman 2006). Some workers further claim that the tempo of evolution is punctuated (i.e., with periods of stasis followed by periods of rapid divergence; Eldredge and Gould 1972; Gould and Eldredge 1977; Eldredge 1985; Gould 2002) as opposed to a more gradualistic view, and some even question the validity of the current modern evolutionary synthesis (Gould 1980). It seems clear that no new synthesis is needed (the present one is satisfactory to include all known micro- and macroevolutionary trends; Stebbins and Ayala 1981; Grant 1983a) and that both punctuated and graduated patterns must have occurred during evolution (Rieppel 1983b; Levinton 1988), more one way in some groups or more another way in others. See Zeeman (1992) for an effort at using catastrophe theory to resolve this conflict. The more interesting question is what kinds of groups show one type of pattern over another. See, for example, studies on divergence in minnows (Notropis) and sunfish (Lepomis) that suggest change due to gradual as well as punctuated phenomena (Douglas and Avise 1982). See also Malmgren, Berggren, and Lohmann (1983, 1984) for examples of punctuated gradualism in foraminifera.

Systematic studies also help reveal patterns of evolution that are useful and stimulating to other areas of biology. Patterns resulting from evolutionary processes occur at all levels of organization from the local population to ordinal and class lineages hundreds of millions of years old. The ancestor-descendent and associated patterns of relationship over long periods of evolutionary time, called phylogeny, and their reconstruction are of special interest to systematics. Phylogeny is important because it has much to do with the construction of classifications. Different opinions prevail, but most workers (myself included) believe that a classification of a group of organisms should in some measure reflect (or at least not be inconsistent with) its phylogeny. These phylogenetic classifications, therefore, contain the information that can be retrieved and used by other areas of biology. These hierarchies also stimulate ideas on such subjects as the origin of life, the origin and evolution of major groups of organisms (including the human species), and the development of ecological zones through geological time.

The reconstruction of phylogeny received a huge boost with availability of DNA data in the late 1980s and early 1990s. These new data offered much more precision in producing branching diagrams of organisms and in their statistical tests. In a broad context, these efforts are directed toward assembling the Tree of Life (Soltis and Soltis 2001; Cracraft and Donoghue 2004; Ciccarelli et al. 2006; Hodkinson and Parnell 2007). Tremendous progress has been made in obtaining genetic information for thousands of organisms and complete genome sequences for many hundreds, and this has led to all sorts of insights, especially for new classifications for specific groups (e.g., angiosperms; APG 1998; APG II 2003) and for all of life (Cavalier-Smith 1998). New textbooks in systematic botany are also now oriented in this way (Judd et al. 2002; Spichiger et al. 2004; Simpson 2006). Within lower forms of life (the Monera and Protista), however, there have also been surprises with lateral gene transfer now appearing commonplace (Doolittle 1999). This has led to a new concept of a Ring of Life (Martin and Embley 2004; Rivera and Lake 2004; McInerney and Wilkinson 2005) that becomes more bifurcating only toward the tips in the more derived Eucarya. However this sorts out in the future, there is no ignoring the importance that molecular-based phylogenies have had upon all aspects of biology (Harvey et al. 1996).

The relationships of some other areas of biology to systematics are so strong that they are virtually dependent upon data generated in systematic studies. Ecology, biogeography, and paleontology depend in this way upon proper classifications of organisms and accurate knowledge of their distributions. Wilson (1971), speaking in the role of an ecologist, stressed the need for continued support for taxonomic investigations so that the data can be used effectively in ecological work. It is fairly obvious that to understand interactions among organisms requires first knowing what they are and also where they are. Thorne (1972) stressed the absolute reliance of phytogeography upon sound evolutionary classifications, so that meaningful hypotheses on dispersal and vicariant events in particular groups can be formulated. Studies on relationships of organisms, including distributional patterns, have even given rise to new ideas on major events on earth. The impact that the documentation of similar fossil biotas in Africa and South America had upon the development of ideas about continental drift was substantial (Schopf 1970; Hughes 1972).

The original conceptualization developed from within systematics (e.g., populational perspectives) contributes significantly to a broadening of biology and to a better balance within biological science as a whole (Mayr 1969c:9). This balance is further aided by systematics reminding other biologists that the diversity of life can be (and should be) studied profitably at all levels of hierarchical organization rather than by focusing all attention through myopically reductionistic eyes only on chemical and physical attributes. Of course, it is quite possible that we could fully account for the properties of each whole if we could know the precise characteristics of all the parts and know in addition all existing relationships between them. Then we could reduce the characteristics of the whole to the sum of the characteristics of the parts in interaction. But this involves integrating the data not merely for three bodies, but for three thousand, three million, three billion, or more, depending on the whole we are considering (Laszlo 1972:8). Needless to say, this type of detailed understanding will be a long time in coming (if ever). In the meantime, a broad view of biology with an emphasis on organisms is entirely appropriate for continued advances to be made.

CHAPTER 3

The Importance and Universality of Classification

Of the numerous important contributions that systematics makes to society and biology, none is more significant than that provided by classification (and its theoretical and methodological umbrella, taxonomy). Classification is a pervasive human quality like the predisposition to sin, it accompanies us into the world at birth and stays with us to the end (Hopwood 1959:230). Although it cannot be denied that the construction of classifications provides intellectual satisfaction for those who make them (J. A. Moore, in Warburton 1967), and, in my opinion, this by itself is justification enough, many more positive features of classification also exist. Heywood suggested that the societal value of taxonomists and their classificatory efforts and products would be negligible: what effect would a strike of taxonomists have? The immediate effects would be few! A handful more people would die each day as the narcotics bureaus and emergency hospital services were unable to identify plant material; plant introductions might be halted. The papers on some biochemical-taxonomic topics might become even more bizarre without the advice of taxonomists! But it would be a long time before there were any serious consequences (1973a:143). But Isely (1972) showed convincingly, with an imaginative example of the hypothetical disappearance of all taxonomists and their works, that the long-term consequences of the cessation of taxonomic efforts would be disastrous for society. This would be especially true in the loss of the information content of classifications and identification services derived from them. The term classification has been used in two general senses: (1) the process of classification (i.e., the manner in which grouping and ranking of items is accomplished) and (2) the results of the process, viz., the resultant hierarchy of classes. The importance and universality of classification derive from both these aspects.

Process of Classification

The significance of the process of classification can be viewed within our innate mental activity and attempts at description and formalization of this activity. The human species has a compulsion for order. This need probably derives from a desire to deal with the environment in a predictable way. Knight described it well:

The world into which we are born is a booming buzzing confusion, and we only slowly learn to sort out things of like kind. Instinctively in babyhood, and later more self-consciously, we group things together and attach general terms to them; so that instead of a chaos of endless particular things without apparent order, we come to perceive a world with a finite number of classes of things. We thus begin to feel at home, even though the classes may need to be revised (sometimes painfully) and certainly seem to cut across each other so that everything belongs in more than one. We distinguish parts of ourselves from other things, and we then separate things that are accessible from things that are not, like the Moon, for which there is little point in crying. Some classes have sharp lines, and others have fuzzy ones; the divisions between colors, for example, seem hard to learn and are mastered some time after children have got size relations straight, and differ between cultures. Great scientists are Peter Pans, still anxious to classify and explain at an age when most people are concerned with money, power, and sex. (1981:16–17)

Because of our greater intellect, we alone of all species are smart enough to worry about ourselves and our existence on earth (hence the development of such disciplines as science, philosophy, and religion; for perspectives on how monkeys see the world, see Cheney and Seyfarth 1990:305-308), and the process of classifying everything in our environment (and the utilization of the resultant products) can be interpreted as an attempt to lower this risk of uncertainty of living. To be confused about what is different and what is not, is to be confused about everything (Bohm 1980:16). In effect, we are trying to describe and interpret reality (at least as far as we can ever come to know it). We classify everything in our environment including animate and inanimate objects such as furniture, cars, houses, clothes, diseases, and the elements (consider the periodic table or particles in the Standard Model; Seife 2003). We even rely formally on methods of classification in some nonbiological disciplines (e.g., linguistic analysis, Hoenigswald 1960, Jordan and Swartz 1976, Platnick and Cameron 1977, Halle, Bresnan, and Miller 1978; anthropology, Lomax and Berkowitz 1972; management theory, Goronzy 1969, Higgins and Safayeni 1984; psychology, Kee and Helfend 1977, Campbell, Muncer, and Bibel 1985, Chiribog and Krystal 1985; medical tests, Pepe 2003; and astronomy, Gehrels et al. 2006). Evidence for the pervasiveness and importance of classification for humans comes from studies of aboriginal peoples of the world who classify objects and ideas in their environment in much the same basic fashion as do professional western taxonomists (Berlin, Breedlove, and Raven 1966, 1974; Raven, Berlin, and Breedlove 1971; Berlin 1973, 1992; Atran 1990). Recent studies have shown that indigenous peoples in Amazonia use geometric concepts such as points, lines, parallelism, right angles, distance, angles, and other shape relationships without any schooling (Dehaene et al. 2006). Obviously many differences exist in the details of resultant classifications produced by aboriginal vs. progressive western cultures, but the process of classification is the same. Sometimes even the hierarchy of classes is remarkably similar, except for a tendency in aboriginal societies for many more subdivisions and variations for organisms and things that are economically and/or culturally very important (over-differentiation; Berlin 1973).

Because of the importance of the process of classification in our innate mental activity, we have attempted to describe this process more clearly and even to formalize our way of thinking about it. The early attempts go back to Plato and essentialism (Hull 1965) and to Aristotle and logical division (Atran 1990). With Plato’s essentialism, one could view attempts at classification as simply the search for the eidos, or essence of things, which would reveal their true natures and allow for proper communication about them as well as correct alignments together in classification. The search for essences and, in a broader sense, the attempt to order the world through the construction of classifications may be related to the nature of language. Bohm suggested that this may be due to the subject-verb-object structure of sentences: "This is a pervasive structure, leading in the whole of life to a function of thought tending to divide things into separate entities, such entities being conceived of as essentially fixed and static in their nature. When this view is carried to its limit, one arrives at the prevailing scientific world view, in which everything is regarded as ultimately