Systematics of Biological Systematics (Or, Taxonomy of Taxonomy) Author(s): Ernest Small Reviewed work(s): Source: Taxon, Vol.

38, No. 3 (Aug., 1989), pp. 335-356 Published by: International Association for Plant Taxonomy (IAPT) Stable URL: http://www.jstor.org/stable/1222265 . Accessed: 28/05/2012 19:58
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TAXON 38(3):335-356. AUGUST 1989

SYSTEMATICS OF BIOLOGICAL SYSTEMATICS (OR, TAXONOMY OF TAXONOMY) Ernest Small

Summary whichclarifyorganization amongorganisms by analysisof relationships, Taxonomyandsystematics, clarification of theseparameters. andnaming,ironicallythemselvesrequire Thefollowing classification, of researcharea;relationshipwith other biological sciences--whether are concerns:circumscription or not; subdisciplinary and nomenclatural or not, and whetherhierarchical classification; overlapping analysis of the disciplinarytitles: taxonomy, systematics, biosystematics,experimentaltaxonomy, genecology,and populationgenetics.Taxonomyand systematics,and theirinternalsubdivisions,may be conceived variouslyin their relationshipsto each other and to other sciences. All such systematizations sufferfrom the same problem as does biological classification:complex, overlapping,and indeed evolving relationshipsare distortedfor the purposeof simplification.Systematicdisciplinary titles are problematical,includingG. G. Simpson's widely employed definitionsof systematicsand as equivalent,mutuallyexclusive, contaxonomy. Systematicsand taxonomy have been interpreted tainingeach other (most desirablywith taxonomy as a subfieldof systematicsconcernedwith formal and overlapping. There should be encouragement for the trendof usingbiosystematics classification), as a synonym of biologicalsystematics,ratherthan in any of the numerousways the word has been employed. So oft in theologic wars, The disputants, I ween, Rail on in utter ignorance of what each other mean, And prate about an Elephant Not one of them has seen! John Godfrey Saxe (1865), The blind men and the elephant Introduction The six blind men in the above extract from the well-known Hindu fable inspected different parts of an elephant, and arrived at different conceptions of it. Each was partly right, but substantially misinterpreted the elephant for two reasons: failure to appreciate the limitations of their own viewpoints, and disregard of their companions' observations. To avoid the distorted conclusions of the blind men, scientists must comprehend the restrictions of their perspectives and the relative perspectives of the sciences. In response to the explosion of knowledge and the search for more, biology has become extensively subdivided into areas of specialization. Disciplines and sub-disciplines can be related in the same ways that formal taxonomic groups are related (that is, they can have relative rank, circumscription, and position). As others have pointed out (e.g., Blackwelder, 1967, p. 348), it is ironic that specialists in biological classification have failed to provide a clear systematization and clear nomenclature for their own field and its components. It has been argued, with appreciable justification, that "vague" terminology may be advantageous in science (Rosenberg, 1975): disciplinary titles are often mental straightjack1

Biosystematics Research Centre, Agriculture Canada, Research Branch, Central Experimental Farm, 335

Ottawa,Ont. KIA OC6,Canada.

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ets, limiting investigators to the theoretical foundation, if not indeed models, methodology and accumulated information, of what is conceived to be the circumscription of that title. It is interesting to contemplate what Charles Darwin's accomplishments would have been if he had remained disciplinarily restricted to taxonomic malacology. In attempting to understand the perspective and goals of a science - especially by defining it - there is danger of placing that science in a straightjacket. Nevertheless, as the taxonomist is well qualified to appreciate, a comprehensible terminology for scientific fields is desirable simply for clear communication. As will be reviewed, however, names of fields of classificatory science have been coined with limited, if any appreciation of overlapping, if not identical labels; have competed and evolved in response to what has been considered new and fashionable; and have mutated over the years, sometimes to the point of extreme ambiguity. In short, what is needed is a systematic examination of biological systematics, and a nomenclatural analysis and consequent synonymization of terms denoting systematically-oriented disciplines. Many of the semantic difficulties treated here could have been avoided if the well-known and easily comprehended term "classification" had been the basis for a title such as "bioclassification"; as Williams (1967) wrote: "the human race has always classified, for the use of any common noun implies a classification" (cf. various meanings of classification in, for example, Jardine (1969)). This paper turns the methodology of biological systematics upon itself. Specialized fields within systematics in its broadest sense are treated analogously to populations, and are considered as posing at least some similar phenomena of evolution (mutability, combinability, adaptive "fitness" to scientific and political needs) and problems of classification (specification of circumscriptional criteria, whether hierarchical or overlapping). I hope that the analogy is provocatively heuristic, but it has its limitations. In particular, there are no codified conventions for clarification of nomenclature for sciences and subsciences as there are for groups of organisms. Also, unlike groups of organisms, concepts of fields of science may be inherently subjective, and are often rather vague and flexible. It will be noted that I have reversed the normal order of taxonomic examination, by beginning with established names of scientific fields, and then searching for discontinuities between the designated concepts, rather than vice-versa. It would be simpler, if of academic interest only, to circumscribe scientific fields anew, and subsequently seek out the most applicable names. There is ample literature on what taxonomists and systematists do, and how and why they do it, which of course provide the bases for circumscribing these fields. It is particularly important to specify purpose (Gilmour and Turrill, 1941). McNeill (1979) concluded: "at least above the species level, the purpose of classification in biology is to provide a simplified expression of the interrelationships among organisms". The more general systematic objective is simply to clarify relationships among organisms. "Relationship" may be phenotypic or genotypic (or both), a grade or a clade (or both), combinatorial (simple hybridization, higher-level gene transfer), or just temporal. Whether or not it is acknowledged, systematists find it difficult or impossible to conceive of relationships except in the context of evolutionary theory (Anderson, 1974). Another principal consideration is human psychology, and here there are several constraints, perhaps the most important of which is the visual Gestalt. As Heywood (1973) noted, "because of man's visual sense, the biological classifications he makes are primarily morphological. He needs to see what he is doing ... there is a morphological bias built into the system before any of the taxonomic processes can begin". (Compare Stafleu's (1969) discussion of "non-visual systematics".) Another factor is that humans find it difficult to think except in terms ofa limited number of discrete, obvious groups (McNeill, 1979). Still another consideration is the inevitability, indeed desirability of artistic expression in systematizations. Cowan (1955) remarked "the 'best' classifications are those made by artists with the keenest appreciation of what is both useful and intellectually satisfying".

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Biological Science and OtherSciences Relationshipof Classificatory

Any field of research implies restriction to a discipline or subdiscipline, with its associated methodology and accumulated data. In the classical scheme of Tschulok (1910; see analysis in Major (1958)), eight quite separate "viewpoints" were applied to biology, namely composition, structure, physiology, genetics, ecology, history, chorology and taxonomy. The resulting sciences were alleged to be independent. However, when one considers the relationships between major fields, such as biology, geology, and chemistry, it becomes apparent that there can be considerable overlap. Biogenetic pathways may be as important to biology as to chemistry, the study of fossils as important to geology as to biology. Moreover, sciences evolve; for example, the delimitation of chemistry is becoming increasingly problematical (Garfield, 1986; Maddox, 1985). Clearly, there is so much overlap between fields of science as presently conceived that their circumscription (horizontal categorization) is often difficult. Without belabouring the point, it is clear that sciences and subsciences are subjective, arbitrary, human constructs, susceptible of different kinds of separation. A second problem is that of hierarchical recognition (vertical categorization), and in this regard the organization of university departments is illuminating. Gardiner (1988) remarked that an "identity crisis has developed in the biological departments of our universities." Should "genetics" be a subdivision of the department of "physiology"? Or vice-versa? (Note one resolution in Table 1.) The widely used Dewey Decimal Classification is the oldest library classification system, and is illustrative of the problem under consideration. As remarked by the editor of the most recent edition (Custer, 1979a): "classification may be applied not only to tangible objects and beings, but also to processes, to actions, to relationships, to mental concepts, in fact to any kind of subject or group of subjects the members of which show likenesses as well as differences". A portion of the system important to biology is shown in Table 1 (Custer, 1979b). Assignment of any given book to its position (analogous to identification of organisms) is done in key-like fashion by considering the set of alternatives at any decimal level. The system is hierarchical, and in theory the classes are mutually exclusive. In fact, depending on the weight allotted to an alternative, a book will be assigned to different classes by different librarians. This is of course different from identifying organisms on the basis of their characteristics, where the discriminatory value of an attribute should not arbitrarily vary with the subjective opinion of the observer. The Dewey system is an example of hierarchical organization of mutually exclusive groups, and for the purpose of systematizing biological science, is both advantageous and disadvantageous. It is good in being a stable, widely used, and at least somewhat reasonable system (perhaps like the Englerian system on which many herbaria are organized). It is bad in having been erected subjectively and somewhat arbitrarily, in lacking the flexibility of permitting the incorporation of better organization as it is conceived, and indeed in providing a rather rigid (rather typological) way of consideration. (Only minor modifications of the Dewey system have been made since its inception in 1873; the Library of Congress Classification system is more flexible.) A "logical grouping or classification of the zoological sciences and in particular of the systematics fields" was attempted by Blackwelder and Boyden (1952), whose hierarchical result is given in Table 2 (the indented tabulation was inferred by me from their text). However, these authors were not satisfied with the separation of their groups, and concluded that insofar as systematics is concerned, the many contributing sciences are completely interdependent, producing one comprehensive class. Some ways of interpreting the relationships of systematics and some principal related fields are shown in Figs. 1-4. The hierarchical arrangement of Fig. I is simplistic, but emphasizes well the highly varied data base of systematics. Figure 2, in which systematics is viewed as a central or core discipline receiving data from many other sciences, is probably the most attractive of the four figures to systematists themselves. Figure 3 interprets sys-

338 Table 1. Extractof Dewey Decimal classificationfor biologicalsubjects. 560 Paleontology Paleozoology 561 Paleobotany 562-569 Specificanimals and groupsof animals 570 Life sciences 571 (Unassigned) 573 Physicalanthropology 574 Biology 575 Organicevolution and genetics 576 Microbes 577 Generalnatureof life 578 Microscopyin biology 579 Collectionand preservationof biologicalspecimens 580 Botanicalsciences 581 Botany

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581.1 Physiology of plants (581.11 = genetics; 581.19 = biophysics and biochemistry)

581.2 581.3 581.4 581.5 581.6 581.7 581.8 581.9

Pathologyof plants Developmentand maturationof plants Anatomyand morphologyof plants Ecologyof plants Economicbotany (Not assigned) Tissue, cellular,molecularbotany treatmentof plants Geographical

tematics as one of many overlapping sciences of equal status, and probably is nearer to the viewpoint of most biologists than the preceding model. Alternatively, this could be illustrated by a completely joined network. Figure 4 illustrates three axes of a hyperspace in which each interacting discipline contributes a dimension (see Sattler (1987) for an analogous concept from the point of view of morphogenesis). Theoretically, one could conceptualize a systematic subdiscipline for every approach to biology: morphological (-anatomical), evolutionary (perhaps including paleobiological), genetical, chemical (-molecular), and mathematical systematics seem reasonable in terms of principal current approaches. Ecological (see Duke's (1977) discussion of "ecosystematics"), economic, geographical, physiological, physical, geological (i.e., stratigraphicfossil), psychological (i.e., ethological), and pathological (see Savile's (1979) use of susceptibility to fungi as an indicator of relationship) systematics are other activities of biological systematics involving separately recognized fields, although the hybrid disciplines are not usually conceived of as meriting subdisciplinary recognition. Although it may seem farfetched, there are even possibilities of hybrid fields between biological systematics and areas of inquiry usually conceived of as not belonging to the realm of science. Thus theology could be joined with biological systematics (as it indeed once was) to clarify the natural organization of living things by God(s); and art intermingles with biological systematics in producing aesthetically appealing systematizations. Biological classification clearly is most related to the companion sciences evolution and ecology, which are also synthetic, holistic, genetically-based, and organismic-populational. In practice, these fields are so complementary that one cannot avoid characterizing them as overlapping. Kruckeberg (1969b) noted "it is plant taxonomists, not taxa, that make distinctions between the formal disciplines of ecology, evolution, and systematics". Turrill (1952) observed that "taxonomists neither claim absolute national sovereignty over any ... field of research nor recognize such a claim by other biologists". Constance (1953) held that "for its basic data ecology-again like taxonomy-is almost wholly dependent upon other disciplines, and the attempt by some to maintain that ecology is quasi-independent

1989 AUGUST and Boyden's(1952) classificationof the zoological sciences. Table 2. Blackwelder A. Dealingwith one or a very few kinds Dealingwith the species Homo sapiens Medicine Psychology Physicalanthropology Dealing with domestic and a few other animals Veterinarymedicine Apiculture Animal breeding medical aspect Subjectswith a predominantly Physiologicalembryology Histology Cytology B. Dealing with kinds as such Deal principallywith determiningthe differencesand similaritiesbetweenkinds Comparativeanatomy Comparativephysiology Anatomicalembryology Comparativehistology Cytology Zoogeography Geologicaldistribution Assemble the data into orderand make them availablefor use Taxonomy Classification of broad significancefrom the classifieddata Draw conclusionsor generalizations Phylogeny Evolution Investigatethe causes of the observedfacts and thus contributeto their recognition Morphology Stratigraphy Genetics C. Dealing with kinds in relationto their environment Ecology(=bionomics = naturalhistory)

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of these disciplines and a sub-science in its own right because it has its own objectives, methodology, terminology, and practitioners has the aspect of a losing cause". In theory, at least, the three goals of clarifying genetic change over time, clarifying genetic organization among organisms, and clarifying adaptive environmental relationships, may respectively distinguish evolution, taxonomy, and autecology. Munroe (1964) wrote "systematics is an approach to biology rather than a department of it". Similarly Major (1958) concluded that sciences like systematics fundamentally amount to a point of view of plants which overlaps the viewpoints of other sciences. Considered in these perspectives, systematics may be characterized as an approach to or viewpoint of biology stressing genetic organization from the level of the individual upward. Similarly, evolution is an approach stressing (populational) change, and autecology is an approach stressing adaptation (cf. Fig. 4). Egocentricity of "Biological Systematics"; Systematics sensu latissimo Chauvinism is common in science, and some semantic distinctions obscure the essential nature of biological classification. The position has been taken that classificatory science is more fundamental or elementary than all other biological sciences, because all others proceed from a taxonomic base (Simpson, 196 1). This is of course true, but other sciences

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\LOG\ CAL SC/

SEVOLUTION
GENETICS ECOLOGY MORPHOLOGY BIOGEOGRAPHY EVOLUTION BIOCHEMISTRY
GENETICSBIO

ECOLOGY
GEOGRAPHY

MORPHOLOGY BIOCHEMISTRY SYSTEMATICS ECOLOGY

ECOLOGY
EVOLUTION BIOGEOGRAPHY EVOLUTION

GENETICS

BIOCHEMISTRY

MORPHOLOGY

SYSTEMATICS

Figs. 1-4. Models of relationshipsof some biological sciences. (Scienceschosen are exemplary, 2. Systematicsas a core not exhaustiveof the possibilities.)1. Systematicsas the apex of a hierarchy; 3. Systematicsas one of many overlapping scienceto which all otherscontribute; sciences;4. Systematics as one of many scientificdimensionsor perspectives. reflect basic human value perceptions comparable in scope to the "what is it" of taxonomy ("what good is it" for economics, "what does it do" for physiology, etc.), and all sciences have their indispensable values. Another overstated argument for the uniqueness of systematics is that it is the most inclusive of all biological sciences, synthesizing data from all other disciplines while generating none of its own (Simpson, 1961). The same claims can be made for ecology and evolution, and in any event many biological data are not of interest to systematics. Furthermore, systematists do collect their own data as evidenced by specialization in fields that have proven especially useful in providing data, such as morphology and chemistry. Simpson (1961, p. 5) stated that taxonomy has "almost a superscientific place among the sciences". However, as pointed out by Riedl (1977, p. 366), we should "dismiss the pretentions that master fields exist in the natural sciences". Simpson (1961, p. 6) noted the argument that, in a general sense, all science involves ordering, so that accordingly systematics and science are coextensive. Hennig (1950) stressed this idea, which seems to recur independently; Ayala (1968) stated "the goal of science is the systematic organization of knowledge about the universe". Indeed, science may be considered to be a "system". As pointed out by Rowe (1961), "since at least the time of Compte, the idea of arranging and classifying the sciences in a hierarchy according to level of inclusiveness or level of integration of subject matter has been common currency" (cf. Guttman, 1976; MacMahon et al., 1978). Table 3 orders sciences sequentially by the comprehensivenss of the units studied, and notes systematic products of each (cf. the highly imaginative schemes in Table 1.2, Bertalanffy (1968) and Table 1 (Griffiths 1974)). The term biosystematics, discussed in detail later, is rather presumptive (as is "biological

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Table 3. An analysisof disciplineson the basis of units studied and systematizations achieved. Unit studied Energy,subatomicparticles Atoms, molecules Tissues, organs Singleorganisms Groupsof geneticallyrelatedorganisms Groups of habitat-related organisms related Groupsof geographically organismsand organism-related regions Ecosystem Field of study Physics Chemistry Histology,anatomy Morphology(s. str.) Taxonomy Phytosociology Zoogeography, phytogeography Synecology Exampleof systematization Classification of elementaryparticles Periodictable of elements Tissue, organclassifications Growthform systems Taxonomic systems Phytosociologicalsystems provinces,phytogeoZoogeographic graphicregions Ecosystematicsystems

species"). As is clear from Table 3, systematization of biochemicals, biological organs, and biotas could legitimately be called "biological systematics", although in common with most current usage, the phrase is used in this paper for the systematics of groups of whole organisms, associated on the criterion of genetic relationship. "Organismic systematics" is more accurate than biological systematics, but could tIeoretically refer to classification of organisms on grounds other than the purely genetic, and for example could include the classification of biotas. In General system theory, von Bertalanffy (1968) presented an expansive "systematic" perspective of science. He sought natural "systems" (defined as complexes of interacting elements) in diverse spheres of knowledge ranging from physics through the social sciences, and further looked for descriptive laws bearing generally on such organization. For example, one might seek parallelisms in organization of a cell, a solar system, and a city. Bertalanffy was opposed to the reductionist approach of seeking explanation of organization simply on the basis of the most elementary (physical) constituents. Griffiths (1974) wrote in regard to Bertalanffy's approach, "I believe that his work is important for understanding physical systems, but am not convinced of the validity of generalizing the system concept in this way". It may be that von Bertalanffy has overstated the parallelism of organization addressed by different sciences. Griffiths (1974) pointed out difficulties with the delimitation of populations as systems sensu Bertalanffy (and hence of so-called "biosystematics"); and similarly difficulties with the delimitation of ecosystems (and hence of so-called "ecosystematics"). It is intriguing, if premature, to speculate that there is a common systematic structure and nature to all of reality that transcends elementary physical constitution. Frequency of Use of the Principal Titles: Systematics, Taxonomy, Biosystematics As remarked by Mason (1950) in his examination of these three terms, "it is usage and the history of usage that ultimately moulds the meanings of our words and terms". To clarify the comparative frequency of usage, several surveys were conducted, as described below and tabulated in Table 4. 1. The frequency of use of the terms was compiled, manually from Biological Abstracts from 1943 (when "biosystematy" was coined) to 1968, and by computer from 1968 to and including 1986 employing the Biosis service (Biosis, 1987). In Table 3, the title taxonomy in fact is based on taxa and *taxon*, where * represents any addition; thus cytotaxonomy, chemotaxonomy, taxonomic, taxonomical and taxonomist are incorporated in the total of 42,871. As a matter of interest, this included 1034 cytotaxonom* and 2189 chemotaxonom*. The term systematics is based on systematic* (systematic and systematics occurred in about equal frequency), and similarly biosystematics was based on biosystematic*. Of

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Table 4. Frequencyof use of the majorheadingsof biologicalclassificationscience. Biosystematics 1. 2. 3. 4. Use in ca. 12,000,000 titles of publications, 1943-1986 Use in 1746 titles of the BiosystematicsResearchCentre, 1973-1986 Use in ca. 17,500 names of journals(basedon Biosis, 1987) Use in names of ca. 2600 institutesand departmentswith systematiccollections(after Holmgrenet al., 1981; Arnettet al., 1986) 283 16 0 1 Systematics 10,064 16 22 11 Taxonomy 42,871 76 11 33

the total of about 12,000,000 titles surveyed, about 0.44% contained one of the three terms. The ratio of use of biosystematics: systematics: taxonomy was about 1:36:152. 2. The frequency of use in 1746 titles of all publications of my own institute, the Biosystematics Research Centre ("Institute" until 1986) was surveyed from its inception in 1973 up to and including 1986. The ratio of use of biosystematics : systematics : taxonomy is about 1:1:5. It seems that being in a "biosystematics institute" does stimulate one to use the term in titles more frequently than used by systematists in general. 3. The frequency of use in ca. 17,500 names of journals and special publications, based on Biosis (1987), was compiled. There were no uses of biosystematics, 11 of taxonomy, and 22 of systematics. It appears that systematics is considered a more appropriate term for journals than taxonomy, likely because of the prestige and greater generality attached to the former term. 4. The frequency of use in the names of a total of ca. 2600 institutes and departments (subdepartments in a few cases) was compiled from Holmgren et al. (1981; listing herbaria of the world), and from Arnett et al. (1986; listing locations of collections of insects and spiders). The only "biosystematics" institute was the one I represent (the Biosystematics and Beneficial Insects Institute of Beltsville, Maryland came into being in 1986, and was disbanded in 1988). There were 11 taxonomic institutes and 33 systematic institutes, once again systematics apparently having a more desirable connotation than taxonomy, although the latter is by far the most frequently used term in publication titles. Are Systematics and Taxonomy Distinct? This question has often been superficially addressed, frequently in otherwise authoritative texts. In the following I have limited the discussion to what I consider to be the pivotal considerations and key illustrative literature. The various ways of interpreting the relationships of taxonomy and systematics are shown in Figs. 5-9. Taxonomy was coined by A. P. de Candolle (1813, p. 19), who referred to "de la Taxonomie (2), ou de la Theorie des classifications appliquee au regne veg6tal"; and in footnote (2): "Mot form6 de ra4Ls ordre, et votos loi, regle". Mason (1950) concluded that in the context of de Candolle's book, classical taxonomy was concerned with relationships (primarily morphological), classification, hierarchical categorization, nomenclature, and description. Mason was unable to find a precise derivation for systematic, but noted that its use in biology as referring to a hierarchical arrangement of organisms may have predated the word taxonomy; however, he stated that taxonomy was older than systematic botany. By contrast, Camp (1951) suggested that systematics is a much older term, and Ornduff (1969) suggested that the two terms originated at about the same time. Noting that the literature he surveyed often used taxonomic and systematic synonymously, but that the latter term was often used to indicate comparative biological study,

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= SYSTEMATICS

SYSTEMATICS

5 TAXONOMY

6 SYSTEMATICS

SYSTEMATICS

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7 TAXONOMY SYSTEMATICS

9
6. MuFigs. 5-9. Conceptionsof relationshipbetweentaxonomy and systematics.5. Equivalent; tually exclusive; 7. Systematics as a component of taxonomy; 8. Taxonomy as a component of systematics;9. Overlapping. Mason (1950) advocated retention of taxonomy as the term of choice, and offered a relatively narrow definition of systematic botany centred on approach: "we may define Systematic Botany as the comparative study of any related (systematic) group of plants utilizing the research techniques of any of the divisions of botany". Thus classification, nomencla-

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ture, and description (i.e., what is usually conceived of as a "system") are left to taxonomy. Effectively, Mason's distinction separates as means (systematics s. str.) from what is usually considered the end of classificatory science (i.e., the system). In a similar vein, Davis and Heywood (1963, p. 452) wrote "much of the debate, in fact, centres around whether it is the aim of taxonomy to study products or processes". Of interest, Mason's implicit definition of systematics (a word not used in his 1950 article), which does not reflect common usage, may be construed as excluding taxonomy, and vice-versa (see Fig. 6). Simpson (1961), who also noted that taxonomy and systematics were commonly used synonymously (as he did in his earlier writings), proposed a distinction by which systematics is broader than taxonomy, including it (see Fig. 8; he also included classification, and nomenclature, which as noted in the following he excluded from taxonomy). Simpson stated "systematics is the scientific study of the kinds and diversity of organisms and of any and all relationships among them". This definition has been parroted in numerous texts, usually without appreciation, in my opinion, of what he went on to state: "in this definition the word 'relationship' is to be understood not in any particular, narrow sense (for instance, in the sense of phylogenetic connections), but in a fully general way, including specifically all associations of contiguity.. .". "Relationship" in the context of classificatory science is normally assumed exclusively to designate genetic ("phylogenetic", "evolutionary", "patristic", etc.) affinity; Simpson made clear, however, that he included biogeographical and ecological relationships. Since these are not heritable, they do not bear directly on what taxonomists-systematists have conceived of as the core of their domain. Homoplastic relationship (whether due to convergence or parallel evolution), chronistic relationship, spatial relationship, ecological relationship, and still other kinds of relationship may serve to clarify heritable relationships, and therefore may be of "systematic" relevance, but their inclusion in a definition of systematics is challengeable. Most systematists would probably accept the breadth of Simpson's definition insofar as it includes biogeography; but I think many would not accept Simpson's (1961, p. 9) inclusion of synecology as a branch of systematics, since the study of community relationships of organisms addresses a conceptually different kind of organization than systematics addresses. Simpson's comprehensive definition of systematics leaves the impression that any kind of relationship is appropriately studied under the heading of systematics. Similarly, the adjective "systematic" has been so frequently applied as in "systematic anatomy" that it might seem that any kind of comparative study is appropriately termed systematic (note Mason, 1950). However, Simpson (1961, p. 9) held that comparative autecology is essential for some aspects of systematics, but he nevertheless specifically excluded autecology from systematics. The position is taken here that the ultimate aim of systematists is the clarification strictly of genetic aspects of relationships, and thus it might seem that one is proposing to exclude some who like to be called systematists from the systematics club, and to restrict systematists from conducting the activities of other specialists. However, this review leads to the conclusion that historical attempts at circumscription of sciences and subsciences have hindered rather than promoted communication. While it is particularly appropriate that systematists inquire on what, fundamentally, distinguishes systematics (self-analysis is sometimes useful), it would be regrettable if this resulted in constraints on either research or outlook. It is not easy to find boundaries between the sciences, any more than circumscribing a city from its suburbs, satellites, and sister cities. Often individuals working in a city, and making valuable contributions to it, are not technically residents of that city, but of another. So long as the boundaries are permeable, they are not unduly restrictive, and may serve to unify efforts towards a common principal goal. Solbrig (1970, p. 4), an advocate of Simpson's definition of systematics, recognized four main types of relationships between plants, based on: descent (phylogeny), similarity (phenetics), space (geography), and trophic relations; he stated that systematists deal with all four, although emphasizing phylogenetic and phenetic relationships. In my view it is mis-

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leading to assign the latter two kinds of relationship to systematics, although admittedly such may bear indirectly on interpreting genetic affinity. Benson (1943, p. 99) expressed the idea well: "the real goal of systematic botany is of the organization--arrangement made up according to their myriads of living plants into a readily understandable system genetic or 'blood' relationships". For taxonomy, Simpson echoed a distinction of Gregg (1954) who in effect distinguished theory and practice, respectively as "methodological taxonomy" and "taxonomy proper". In Simpson's words: "taxonomy is the theoretical study of classification, including its bases, principles, procedures, and rules". Simpson noted the etymological justification for a theory-concerned definition of taxonomy (note later comments on theory vs. practice). Mayr (1982, p. 146) made a similar distinction, dividing taxonomy into microtaxonomy, dealing with methods and principles of recognizing and delimiting kinds (species), and macrotaxonomy, by which kinds are arranged as a classification. Simpson's resolution of the meanings of the terms systematics and taxonomy has been widely popularized, and has become a sacred academic cow. A reviewer of this paper wrote "I have one major question to ask, and I am sure that other readers who have worried about this subject will too. That is, what is wrong with Simpson's classical definitions of systematics and taxonomy? I have always regarded them as the best statements of the differences between the terms, and a large number of later authors have simply modified them. One of them deals with relationships--any and all relationships--and the other with classification." Above, the viewpoint is taken that including all relationships in the definition of systematics is not desirable because it includes a breadth of non-heritable variation that dilutes the focus of classificatory science. Further, Simpson himself arbitrarily excluded the study of some relationships. For example, it is instructive to note that he substantially excluded what has become the latest systematic bandwagon, molecular biology, because "it does not necessarily or (at present) even usually focus attention on the kinds or diversity of organisms, and therefore as a special biological science it is not a part of systematics" (1961, p. 7). In any event, as the above reviewer perceived, it is desirable to allocate classification under the rubric taxonomy. The problem, as pointed out in the preceding, is that Simpson did not fully do this; he assigned only theoretical aspects of classification to taxonomy, and in segregating the practical aspects of classification to systematics, he removed from taxonomy what most taxonomists understand to be included. The attempt to distinguish taxonomy and systematics on the basis of theory vs. practice has been widespread. However, as should be clear from the following, different writers have had contradictory ideas of the way the distinction should be be made, and indeed of just what is theory and what is practice. Camp (1951) specifically distinguished taxonomy as theory, systematics as practice: "we should keep clearly in mind that as originally defined by A. P. de Candolle (1813), the term 'taxonomy' must be retained for the study of the laws and principles underlying a system of classification. Systematics ... refers only to the classification of objects within a nomenclatural system" (note Fig. 6). Heslop-Harrison (1953, p. 128) defined taxonomy as "the study of the principles, practice and results of the classification of organisms", and systematics as "the practice of describing, naming and classifying living things" (i.e., systematics is a practical component of taxonomy; Fig. 7). Solbrig (1966, p. 116) stated "the relationship of taxonomy to systematics is somewhat like that of theoretical physics to the whole of physics" (i.e., taxonomy is a theoretical component of systematics; Fig. 8). Similarly, Heywood (1973), who accepted Simpson's definition of systematics ("the scientific study of the diversity and differentiation of organisms and the relationships (of any kind) that exist between them"), stated for taxonomy: "although this term has been widely misapplied in the past, there has been a tendency in recent years to restrict it to its original meaning, namely that part of systematics which deals with the study of classification, its bases, principles, procedures and rules. It therefore covers ... all such techniques as are involved in the mechanics of actually preparing

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classifications and keys of any kind from the raw data". (This inclusion in taxonomy of practical aspects of classification is clearly inconsistent with Simpson's narrow circumscription of taxonomy.) Blackwelder (1967) used taxonomy to refer to "the day-to-day practice of dealing with the kinds of organisms", and systematics to "the study of the kinds and diversity of organisms, their distinction, classification, and evolution" (Fig. 9). In most texts treating classificatory science (perhaps more evidently in North America than elsewhere), systematics is equated with taxonomy (e.g., Ross, 1974; Jones and Luchsinger, 1979; Radford et al., 1986; Fig. 5). In practice when a distinction is made, systematics is usually considered to be more inclusive than taxonomy. This is illustrated by Solbrig's statement (1966, p. 114), "systematic studies often but not always result in classifications". It has been said that the systematist is more concerned with processes leading to pattern, whereas the taxonomist is more concerned with the pattern per se, but the distinction is weak. Ornduff (1969) noted that "many of us would rather be called systematists than taxonomists, since in some circles the latter carries with it connotation of anachronism". "Biosystematics": Ambiguous, Pretentious, Superfluous, But Useful The publication of a collection of papers as The New Systematics (Huxley, 1940) precipitated problems concerning how biological classificatory science should be conceived and labeled. Although the general tone of the book was highly respectful of traditional taxonomy, and not suggestive of a need for new fields for taxonomy, nevertheless this was the result. As examples, consider Heslop-Harrison's (1953) contrast of "experimental" and "classical" taxonomy (a point of view criticized by Davis and Heywood (1963, p. 456)), and Mayr's (1942) contrast of "The Old Systematics" and "The New Systematics" (criticized by Blackwelder (1967, p. 342)). The climate of the time was marked by excitement that the "new", "experimental" techniques of chromosomal, breeding, and populational analysis were rejuvenating what many conceived of as the rather unscientific, exceedingly dull morphologically-based field of biological classification commonly known as taxonomy. New buzzwords became popular, although their import was not clear; in Constance's (1951) words: "I am none too sure in my own mind just what is meant by the terms Newer Systematics, Experimental Taxonomy, or Biosystematics". Blackwelder (1967, p. 4) appropriately criticized the situation: "During the past several decades some discussions of taxonomy have used terms that may be either derogatory in tone, ostentatious in implication, or pedantic in application. Among these may be cited ... The New Systematics ... biosystematics". Stafleu (1969) observed that "the new systematics" was "a somewhat overly optimistic term, since nothing ages as rapidly as the new". Camp and Gilly (1943) proposed the field ofbiosystematy: "As here defined, Biosystematy seeks (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". At the time, they entertained the thoughts, naive in retrospect, that "natural units" based on mode of reproduction existed below the species level, and a nomenclature for different kinds of breeding situations would be useful. None of an orgy of words they coined for their concepts was taken up except biosystematy, which soon mutated to biosystematics (the earliest use of the word I have encountered is Epling (1943)). Vickery (1984) pointed out that "biotaxonomy" would have been preferable in view of the stated nomenclatural goal. The term biosystematics never became widely applied in its original (albeit vague) sense. Cytogenetics was a dominant force in plant taxonomy for three decades following the coining of biosystematy (Moore, 1984), and the word became associated with botanical cytogenetic studies. Constance (1964) observed that "the cytogenetic period of taxonomy has been an extremely fruitful one, but it, too, has tended to become smug and authoritarian, and it is probably time for a change in emphasis". With the growing realization that fields other than cytogenetics were coming to the forefront of systematic research, biosystematics as a term mutated to become intolerably ambiguous, as noted in the following.

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Jones (1984) wrote: "Both Camp and BiScher have protested at the perversion of the original definition of "Biosystematy" (Camp, 1961; BScher, 1970) and I find myself very much in agreement with them"; and with regard to the derivative biosystematics, "it might seem that there is today some universal agreement as to its meaning, but I doubt it". In Biosystematic literature, Solbrig and Gadella (1970) tellingly comment "Compilers in many countries have worked hard at compiling this index. However, their ... definition of Biosystematics ... varied". In Plant Taxonomy and biosystematics, Stace (1980) quickly revealed (p. v) the lack of conviction of some who use the term biosystematics: "despite its title, the subject matter of this book is taxonomy". Monroe (1977), in a summary of a symposium on biosystematics, stated: "Some seemed to think of biosystematics as being an approach to systematics based on the characteristics of the living organism: experimental, behavioral, or genetic systematics would fall within this concept. Others used it for what was formerly called 'the new systematics,' that is, the whole array of indirect, theoretical, experimental, and modem methods as opposed to 'conventional' systematics". In the authoritative Principles ofangiosperm taxonomy, Davis and Heywood (1963) stated "much of what is included under the general title of biosystematics is simply part of the modern approach to taxonomy and represents usually an extended state of knowledge of taxa. The term biosystematics itself is in fact largely superfluous and serves mainly to describe one of the stages through which taxonomy passes". Johnson (1 970b) decided "perhaps the word 'biosystematics', as applied to some ill-defined discipline, should be allowed to fade away". Raven (1976) concluded that "it is likely that the term 'biosystematics' has been used in so many different ways ... that it has outlived its usefulness". More recently, Raven (1986) wrote "not surprisingly, the term biosystematics is largely being abandoned at present, in favor of 'plant population biology' and various other more specific alternatives". A growing number of authors has arbitrarily chosen to define biosystematics in a personal way, or to make biosystematics essentially coextensive with taxonomy and/or systematics. Stafleu (1969) observed the use of biosystematics as "evolutionary systematics of the living organisms in the widest possible sense". Stebbins' (1970) view was that "Biosystematics, in contrast to classical taxonomy, concerns itself primarily with the processes of evolution, and only secondarily with its end products. Perhaps this is not the original definition of the term, but I do not wish to be concerned with priorities and semantics". Solbrig (1970, p. 5), after stating that "biosystematics is usually interpreted ... as being essentially synonymous with systematics, although with an emphasis on the study of genetical and cytological phenomena", offered the following personal definition: "biosystematics can now be defined more precisely as the application of genetics (and cytogenetics), statistics, and chemistry to the solution of systematic questions in order to provide explanations about the diversity of organisms within the frame of the theory of evolution". Munroe (1977) wrote "I took systematics to be the general science of classification, and biosystematics to be that part of it which fell within the sphere of biology". Ross (1974) stated "I am using biosystematics here as the investigational field of systematics based on any scientific information that can be brought to bear on the problems of the evolution of species, whether they concern speciation or phylogeny". Stace (1980, p. 6) wrote: "it is unfortunate that the term biosystematics has been widened by some taxonomists to cover virtually any taxonomic activity not pursued in the herbarium or almost any newly acquired technique". Savile (1985) defined biosystematics in almost the same words (Appendix 1) that Simpson (1961) had used to define systematics. "Genecology": Systematics + Ecology + Evolution Turesson (1923) stressed ecology in coining the term genecology: "It seems appropriate for several reasons to denote this study of species-ecology by the term genecology (from the Greek 'genos', race, and 'ecology') as distinct from the ecology of the individual organism, for which study the old term autecology seems to be the adequate expression" For Turesson, genecology was the study of plant infraspecific variation in relation to

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environment, and included analysis of mechanisms producing infraspecific adaption. Heslop-Harrison (1964) interpreted genecology in the original Turessonian sense as a "synthetic discipline", merging ideas and methods from taxonomy, genetics, ecology, and physiology. Bennett (1964) stressed the evolutionary aspect of genecology in her redefinition of it: "the study of the genetic mechanisms which operate within organisms, and between organisms and their environment, at the level both of the individual and of the population viewed as a process, which at the population level produce those changes known collectively as micro-evolution." She recorded her view that the terms taxonomy, experimental morphology, new morphology, new systematics, genonomy, biosystematics, and neotaxonomy are not equatable with genecology, and the terms experimental evolution and micro-evolution are more restricted in scope than genecology. (Epling's (1943) ill-defined term genonomy, which was proposed to designate studies that then also fell under the terms "the new systematics", population genetics, and biosystematics, never received acceptance.) Lawrence (1951, p. 169) stated about genecology, "this term has not been accepted in America where it is confused euphonically with the medical term gynecology". A current dictionary definition of genecology (Gove, 1981) has a taxonomic aspect: "a branch of ecology concerned primarily with the species and its genetically variant subdivisions, with their position in nature and, with the controlling genetic and ecological factors" (italics added). However, Turesson (1922) held the view that the study of genetic and ecological variation of natural populations was independent of taxonomy. Langlet (197 1, p. 659) implied that while genecology could help taxonomy, unless the two were maintained as distinct fields, the term genecology ought to be abandoned. Nevertheless, as pointed out by Constance (1953), "genecology slipped away from the realm of plant ecology to that of plant taxonomy". Heslop-Harrison (1964, p. 237) did not quite concede this transfer: "an unfortunate trend during the post-war period has been apparent in recurrent attempts to assimilate genecology into taxonomy". So-called experimental classifications began with Turesson in 1922, and were based on transplantation, breeding, and analysis of environmental adaptations (for details see, for example, Baker (1952); Davis and Heywood (1963); Stace (1980)). It has been clear for at least two decades that genecological classification is a sterile exercise from the point of view of classification (e.g., Constance, 1957; Merxmiiller, 1963), whether based on arbitrary degrees of capacity to interbreed, adaptive differentiation, or a combination of these. Mayr (1982, p. 277) and Heslop-Harrison (1964, p. 237) noted the common interpretation of biosystematics as genecology. Stace (1980, p. 6) wrote: "biosystematics may ... be considered as the taxonomic application of the discipline known as genecology-the study of the genotypic and phenotypic variation of species in relation to the environments in which they occur". However, Heslop-Harrison (1964, p. 238) insisted that "biosystematics (or experimental taxonomy) should ... be preserved as something distinct from genecology ... ecological data and observations on genetic systems are a sine qua non of genecology, although by no means an essential part of taxonomy. Conversely, the nomenclatural and bibliographic studies which are an obligatory part of any taxonomic study are not necessarily significant for a genecological investigation of a species". Kruckeberg (1969a) stated "genecology constitutes, to date, the greatest point of contact between ecology and plant systematics", and "we may look upon genecology as a special branch of microevolutionary study". Clearly genecology is a mixture of systematics, ecology, and evolution, and insofar as it addresses "racial systematics" (cf. the following discussion of population genetics), is of critical importance to systematics. "Population Genetics": Systematics + Genetics Mayr (1982, p. 553) noted that population genetics, which is concerned with gene frequencies in (within, among) populations, is an ambiguous term, on the one hand reflecting a tradition of research on mathematically-based, theoretical, statistical populations, and

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on the other hand research on organisms. In the latter sense, the origin of population geneticshas been attributedto Chetverikovin 1926 (Adams, 1968), althoughMayr(1982, p. 553) acknowledgesthe contributionsof others. Mayr (1982, pp. 559-560) interpreted Chetverikov'stype of population genetics as essentially a combination of systematics(as it appliedto geographic races)andgenetics.Camp(196 1)noted that some equatepopulation geneticswith experimentalsystematics,and he himself equatedmuch of it to his own term, biosystematics.Ford (1964) usurped the field for ecology as ecological genetics. Langlet (1971, p. 659) observed that such usage essentially equates population genetics and genecology. Genes are hereditarydeterminants,and by definitionthe subjectof the field of genetics. Change in gene frequenciesclearly belongs to the field of evolution, and so one would expect a close relationshipwith systematics.However, if one confines the subjectof population genetics to the genotype, then the relationshipof it and systematicssubtly hinges on the basis of genealogical on cause and effect,since even those who advocateclassification in so on the fact do almost the basis of entirely relationships phenotype(afterdiscounting environmentalmodifications).The visible associateof populationalgenetics,populational phenetics,is squarelywithin the realm of Mayr'sterm (1982, p. 247) "populationsystematics".In practice,populationgeneticsas a term is appliedlike genecology,to intra-specific studies in which morphologicaldifferentiationis usually too limited for formal classifithe residue cation.And, like genecology,it is generallynot conceivedas preciselydesignating after taxonomy is deductedfrom systematics. "ExperimentalTaxonomy":Deceptiveand Unnecessary of biosystematicsis that it is "experimental A common interpretation taxonomy"(e.g., Heslop-Harrison, 1954; Boivin, 1960; Stace, 1980). As noted earlier, some have also assignedgenecologyto synonymy. Constance(1953) traced this phrase back to 1919 (cf. Langlet(1971, p. 658) who tracedthe term to 1920, and Heslop-Harrison(1964, p. 237) who tracedit to 1934). Not only is the meaningof the phraseunclear,but also it has never been obvious in exactly what respects experimentaltaxonomy is experimental.It seems that the presenceof laboratory,expensive and complicated apparatus,and novelty, concomitantsof much genuineexperimentation,have often been sufficientto label an activity as experimental(e.g., Munroe, 1960). Constance(195 1) stated"I fail to see how taxonomy, itself, can ever be regardedas experimental,unless one assumes that the success or failure of compatibilityor biochemical tests, for example, are per se proofs that two individuals do or do not belong to the the same taxonomic category".Presumablyit is this rationale that validates the so-called experimentalclassificationsmentioned in the preceding. The concernwith the "experimental" aspectof taxonomywas obviously partof the 20th centuryattempt to transformtaxonomy into a "science",since experimentationis widely (and incorrectly)considered to be a sine qua non scientific procedure.Lewis (1957) and Mayr (1982, pp. 30-32, 521, 855-856) noted that controlledlaboratoryexperimentation is just not that differentfrom intelligent observation and interpretationof uncontrolled naturalphenomena, especially when done from the comparativeperspective of the taxonomist. Indeed, even the so-called experimentaltaxonomist relies overwhelminglyon observationalevidence, usually only crudely controlled manipulation of a very limited portion of the genotype and phenotype being possible. Taxonomic and systematic interpretation are primarily of observational data, and the use of the admittedly att:ractive phrase "experimentaltaxonomy" is usually superfluous. Taxonomy, Taxa By comparisonwith de Candolle's(1813) coiningof the wordtaxonomy,the 20th century term taxon is a latecomer. According to Mayr (1982, p. 870), the term taxon was first

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proposed by Meyer-Abich (1926). Independently, Lam (reported in Sprague et al., 1948) proposed "to indicate a taxonomic group of any rank with the term taxon (plural taxa)" (cf. Lam, 1957; Morton, 1957; Rickett, 1958), and fortunately the word is now almost universally accepted among taxonomists in this one sense. (Unfortunately, the supplement to Webster's third international dictionary (Mish et al., 1986, p. 189), presents the following erroneous definition: "Taxon abbr taxonomic; taxonomy"). I believe that most modern taxonomists share Moss's (1983) viewpoint, "a taxonomist's basic concern is taxa, and how to group and explain them". In contrasting taxonomy and systematics, it is etymologically desirable to establish some consistency between the accepted formal unit of taxonomy, the taxon, and the science of taxonomy, and this is achievable by stressing, as in the following, the essentially formal classificatory nature of taxonomy. Beyond Taxonomy: Unconventional Systematization Formal, codified botanical nomenclature reflects the classical (Aristotelean) viewpoint of classification that things not merely should be, but are hierarchically ordered into absolutely discrete groups on the basis of attributes (Cain, 1959). This has proven to apply well to organisms, but there is a recurrent difficulty. Membership often proves problematical when groups are polythetically defined (i.e., "fully polythetically": group membership not being determined by invariable possession of any one character, but by preponderant possession of characters of a set; see discussion and references in Small (1989)). That is, some characters of a group suggest its membership in one covering group, whereas other characters suggest its membership in another covering group, and there is no really satisfactory way of making the choice. Formal taxonomy cleanly cuts the Gordian Knot of incongruent natural variation with a Procrustean solution: everything is forced into a uniquely named pigeon hole in a hierarchically stratified pigeon coop. There is a gentleman's agreement (to say nothing of the permissiveness of the Code of Botanical Nomenclature) that individual taxonomists of at least minimal competence are entitled to their own formal schemes, but there is a constant search for the Holy Grail of classical taxonomy, the one true system, with its place for everything, and everything in its place. When nature provides variation patterns that are just too complex to be accommodated by conventional taxonomy, so-called special-purpose, informal classifications are permissible (for discussion, see, for example, McNeill (1976)). Michener (1963; cf. Cowan, 1955, 1970) noted the possibility of systematic recognition of "overlapping taxa" as contrasted with conventional, mutually exclusive taxa. DuPraw (1964, 1965a, 1965b) observed that mathematically representing characters by a reduced number of multidimensional axes is quasi-taxonomic, and he used the oxymoronic phrase "non-Linnean taxonomy" (as noted by, for example, Johnson (1970a), while an ordination is an "arrangement", it is not a "classification"; hence it is not a taxonomy). In effect, such simplifying mathematical representations are obviously "systematizations", imposing and/ or revealing order. They are particularly useful for depicting non-hierarchically structured and/or overlapping variation, as is commonly found intra-specifically and in reticulate relationships, and in this respect can be in conflict with the nomenclatural assumptions of hierarchy and discreteness. Baum (1974) attempted to introduce multidimensional axis definition into formal nomenclatural taxonomy (without disturbing the time-honored principle of discreteness of taxa), but the unenthusiastic reception of this proposal (Stafleu and Voss, 1975) suggests suspicion that any form of numericlatural systematization could compromise formal taxonomy. Turrill (1935, 1938) conceived a gradient of taxonomic activity and associated classifications, that he suggested could be labelled sequentially by Greek letters. Alpha classifications are the best that can be done with available information, and one proceeds as taxonomic knowledge increases to the ideal, omega classifications. A parallel scheme is presented in Mayr, Linsley and Usinger (1953, p. 19) who noted a trend of designating

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and namingof speciesas alphataxonomy,placingthe speciesin a hierarchy characterization as beta taxonomy, and relatedevolutionarystudies as gamma taxonomy. It is an oversimplificationto imply, as do these Greek-labelledstages of taxonomic clarification,that all taxonomic problemsare solvable by acquiringenough information.As is well recognized, there is a substantialamount of weakly or confusingly structuredvariation that simply does not lend itself to "classification"in the accepted sense of the word. Indeed, Mayr (1982, p. 145) stated that gamma taxonomy is not strictlytaxonomy. It is at the point that conventionalformaltaxonomyproves insufficientto "systematize" heritablevariationthat a need has been recognizedto designatethe additionalinterestsof specialistsstudyingthe organizationof that variation. This need is met by the practiceof designatingthe more comprehensive field (that also includes taxonomy) as systematics. This is etymologicallysatisfying,since the ultimate aim may be conceived to be a "systematization"of relationships,that is, an accountingof the nature of heritablevariation, most expressly in a formal taxonomy, but in some alternative classificatoryor simply descriptivefashion when appropriate. PublicRelations The technical literatureand referenceworks reflect established meanings for the funandgeneraldictionaries damentalterms"systematics", "taxonomy",and "biosystematics", and encyclopediasindicate the extent to which such meaningshave achieved acceptance beyond the scientificcommunity. These key words are importantnot only for clear communicationamong specialists,but also with others in the scientificfraternityand beyond. Extractsfrom standardreferencesare presentedin Appendix 1. Taxonomyand systematics tend to be equated. Some of the arcane (and contradictory)distinctions that have been made between taxonomy and systematicsare also presentin the examples of Appendix 1, is relativelyrarely and unfortunately probablyconfusethe averagereader."Biosystematics" found, and when given is usually defined in the increasinglyarchaicsenses of cytogenetisystematics.As noted earlier,taxon is misdefinedin cally-basedor experimentally-based the acclaimed Webster'sdictionary (Mish et al., 1986). All things considered, these key systematicterms are not well presentedto the generalscientificand lay communities. Half a centuryago, there was a tendency to view "taxonomy"as effete, old-fashioned, boring,obsolescent,and marginallyrelevant,except as an identificationservice, and even in those gentler times at least part of the solution was seen in the coining of striking and "biosystematics". like "thenew systematics" Sincethen, competitionamong buzzwords the sciences for talentedrecruits,public support,and resourceshas become very keen, and this hard reality is perilously ignored. It does not seem to sufficethat systematics is the foundation of all biological and agriculturalscience, and has contributed more to the collective knowledgeand welfareof mankind than any other field of science. "Successin science depends not only on rational argumentbut on a mixture of subterfuge,rhetoric, and propaganda" (presentedby Broad (1979) as the viewpoint of the controversialphilosopher P. K. Feyerabend). to examine not just the literal, but also the aesthetic and Accordingly,it is appropriate political qualities of the titles of classificatoryscience. In English,at least, the word taxonomy has a strikinglyjarringquality to those unfamiliarwith it, and is absolutelyuninformative to the uninitiated(who are most likely to conclude that it has somethingto do with taxidermy).By contrast,"systematics"is less abrasive,and if still not informativeas to content, is suggestiveof efficientorganizationas opposed to being "unsystematic".It happens that systematic biology does not have an exclusive claim to the adjective (e.g., systematic theology, systematic philosophy), and so it is natural to specify "biological systematics",and to contrastthis to the melodious "biosystematics"(comparethe highly It is regrettable thatthe meaning successful"biocontrol","biodiversity", "biotechnology"). of this combination was so poorly conceived and has become so confused, but, as noted,

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a growing usage of it is essentially to equate it to biological systematics. Taxonomy and systematics need an attractive catchword to convey their essential vitality and relevance, and that need is fulfilled by treating biosystematics as a synonym. Indeed, the term biosystematics has recently been deliberately enlisted to communicate to the public and to other scientists the urgent need to correct the current deplorable and deteriorating support for taxonomy and systematics (Oliver, 1988a, 1988b; cf. O'Hara et al., 1988). Systematics has evolved so rapidly that most of the specialized fields that have been named now seem to be dubiously distinguishable; this dynamism of systematics should lead one to be very reluctant to coin special names for the exciting new developments that are periodically assimilated into our science. The semantic madhouse of subsciences that has been created by systematists is already embarrassingly full. Much of the nomenclatural confusion resulted from lack of focus on the essence of systematics, and therefore its unity. As I have argued, this essence needs to be captured in attractive, simple labels, such as biosystematics. However, the terms taxonomy and systematics are thoroughly established and, separately from whatever distinctions seem appropriate for academic circumstances, these words also need to be presented attractively and comprehensibly to the lay public and scientific community at large. These sciences need to be formulated (and perhaps periodically reformulated) to emphasize their quintessences while permitting sufficiently flexible boundaries to adapt to changing circumstances. Although I have objected to the inclusion of non-heritable variation, Simpson's (1961) definition "systematics is the scientific study of the kinds and diversity of organisms and of any and all relationships among them" is well and concisely formulated. I prefer to characterize systematics as the science of organization and pattern of heritable relationships among the kinds and diversity of organisms. The best contemporary conception of taxonomy is that it is a very substantial but imprecisely separated part of systematics, that is especially concerned with the production of formal classifications of living things on the basis of genetic relationships. Acknowledgments I thank L. Knutson for correspondence on the public relationsvalue of the term biosystematics, and J. Bain, P. M. Catling, A. Cronquist,W. F. Grant, P. G. Holland, J. McNeill, and several anonymousreviewersfor constructivecomments. Some of the criticism has suggestedthat the presentationin parts of the manuscriptmay be controversial.Of course, I alone am responsiblefor its faults. Literature Cited Adams, M. 1968. The foundingof populationgenetics:Contributionsof the ChetverikovSchool, 1924-1934. J. Hist. Biol. 1: 23-39.
Anderson, S. 1974. Some suggested concepts for improving taxonomic dialogue. Syst. Zool. 23:

58-70.
Arnett, R. H., Jr., G. A. Samuelson, J. B. Heppner, G. J. Nishida, J. C. Watt and R. E. Woodruff. 1986. The insect and spider collections of the world. E. J. Brill/Flora & Fauna Publications, Gainesville, Florida. Ayala, F. J. 1968. Biology as an autonomous science. Amer. Scient. 56: 207-221.

Baker,H. G. 1952. The ecospecies--Preludeto discussion.Evolution6: 61-68. Baum, B. R. 1974. Article36 and numericalclassification.Taxon 23: 652-653. in genecology. RecordScottishPlantBreeding Station 1964: Bennett,E. 1964. Historicalperspectives 49-115. Benson, L. 1943. The goal and methods of systematicbotany. Cact. SucculentJ. 15: 99-111. L. von. 1968. Generalsystem theory.GeorgeBraziller,New York. Bertalanffy,
Biosis. 1987. Serial sources for the Biosis data base. Biological Abstracts, Inc., Philadelphia, Pennsylvania. Sydney.

R. E. 1967. Taxonomy.A text and reference book.J. Wiley& Sons, New York,London, Blackwelder, and A. Boyden. 1952. The natureof systematics.Syst. Zool. 1: 26-33.
Bi6cher, T. W. 1970. The present status of biosystematics. Taxon 19: 1-5.

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Boivin, B. 1960. A classical taxonomist looks at experimental taxonomy. Rev. Can. Biol. 19: 435444. Broad, W. J. 1979. Paul Feyerabend: Science and the anarchist. Science 206: 534-537. Cain, A. J. 1959. The post-Linnean development of taxonomy. Proc. Linn. Soc. London, 170 Session (1957-58): 234-244. Camp, W. H. 1951. Biosystematy. Brittonia 7: 113-127. 196 1. The pattern of variability and evolution in plants. Pp. 44-69. In: P. J. Wanstall (ed.), A Darwin centenary. The Botanical Society of the British Isles, London. I and C. L. Gilly. 1943. The structure and origin of species. Brittonia 4: 323-385.

de la botanique,ou expositiondes principesde la Candolle, A. P. de. 1813. Theorie lWmentaire naturelleet de l'art de d&crire et d'tudier les vegetaux,Ist ed. Paris (Deterville). classification
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Appendix1. Definitions of taxonomy, systematicsand biosystematicsin standardreferences(not presentin standardreferenceif not presentedhere). Generalencyclopedias Brittanica(Encyclopedia The New Encyclopedia Brittanica,Inc., 1978) "branchof biology concernedwith the diversityof living thingsand theirgroupingor Systematics: classification" Taxonomy:"the science of biologicalclassification" The Encyclopedia Americana(Mayr, 1981) Taxonomy:"The theoryand practiceof classifyingorganisms" The CanadianEncyclopedia (Savile, 1985) "the scientificstudyof kinds and diversityof organismsand of any and all relations Biosystematics: betweenthem" "the actualarranging, groupingand namingof organisms" Systematics: Taxonomy:"covers the bases, principlesand rules of classificationand nomenclature(naming), includingthe hierarchicframework,and refersnot to a researchdiscipline but to a supporting methodology" La GrandeEncyclopedie (LibrairieLarousse,1976) Taxinomie:"definieprimitivementpar AugustinPyramede Candolle(1813) comme traitantdes cette science se donne en outre pour objectifsde circonscrire, theoriesdes classifications, ... de classer,de definirexplicitementet de denommer ... On emploie souvent la graphieoriginelle, taxonomie,qui resulted'une erreurd'interpretation 6tymologique" (Note: Mayr(1966) contendsthat taxonomyis more correctlyderived than taxinomy.) Dictionaries Webster'sThird New International Dictionary(Gove, 1981) taxonomy esp. as based on cytogenetics" biosystematy;"experimental Biosystematics:
Systematics: "1: the science of classification ... 2: a system of classification ... taxonomy ... 3:

scheme or structure an organizational ..." Taxonomy: "1: the study of the general principlesof scientific classification:systematics 2: the and namingof type groupswithina subjectfield:classification ordering, systematicdistinguishing, Random House Dictionary(Stein and Urdang, 1973) "the study of systems or of classification" Systematics: 2. the sciencedealingwith the identification, Taxonomy:"1. the scienceor techniqueof classification. naming,and classificationof organisms" The New EnglishDictionary("The Oxford EnglishDictionary";Murrayet al., 1919) ".. . 4. Nat. Hist., etc. Pertainingto, following,or arranged accordingto a system of Systematics: of or pertainingto classification,classificatory" classification; esp. in relation to its general laws or principles;that departmentof Taxonomy:"Classification, science or subject,which consists in or relatesto classification" science, or of a particular Technicaldictionaries Featherly(1954) "Taxonomicstudies involving cytologyand genetics" Biosystematy: Systematicbotany:"A study of plants in their mutualrelationshipsand taxonomicarrangement" Taxonomy:"The systematicclassificationof organisms" Jackson(1928) and taxonomicarrangement" Systematicbotany:"the study of plantsin their mutualrelationships Taxonomy:"classification" Torre-Bueno (1978) aftera distinct plan, or method" System:"an orderof arrangement accordingto a system" Systematic:"in definiteorder,or arranged Systematist:"a student of the systematic aspect of classificationin biology, who discriminates forms or groupsaccordingto their relationshipsand affinitieswith betweenspecies and arranges others" of species and groups thereof into a system which shall exhibit their Taxonomy:"the arranging relationshipto each other and their places in a naturalclassification"