small, 1989 systematics or taxonomy of taxonomy

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Systematics of Biological Systematics (Or, Taxonomy of Taxonomy)

Author(s): Ernest SmallReviewed work(s):Source: Taxon, Vol. 38, No. 3 (Aug., 1989), pp. 335-356Published by: International Association for Plant Taxonomy (IAPT)Stable URL: http://www.jstor.org/stable/1222265 .

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TAXON 38(3):335-356. AUGUST 1989

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

SummaryTaxonomyandsystematics,whichclarifyorganization mongorganismsby analysisofrelationships,classification, ndnaming, ronically hemselvesrequire larification f theseparameters.Thefollowingare concerns:circumscription f researcharea;relationshipwith otherbiologicalsciences--whetheroverlapping r not, and whetherhierarchical r not;subdisciplinarylassification; ndnomenclaturalanalysisof the disciplinary itles: taxonomy, systematics, biosystematics,experimental axonomy,genecology,andpopulationgenetics.Taxonomyandsystematics,and their nternal ubdivisions,maybe conceivedvariously n theirrelationships o each otherand to other sciences.All such systema-tizations sufferfrom the same problemas does biologicalclassification: omplex,overlapping,andindeedevolvingrelationshipsare distortedfor the purposeof simplification.Systematicdisciplinarytitles are problematical,ncludingG. G. Simpson'swidelyemployeddefinitionsof systematicsandtaxonomy.Systematicsandtaxonomyhave been interpreted s equivalent,mutuallyexclusive,con-tainingeach other(mostdesirablywithtaxonomyas a subfieldof systematicsconcernedwith formalclassification), ndoverlapping.There should be encouragementorthe trendof usingbiosystematicsas a synonymof biological systematics,rather hanin any of the numerouswaysthe wordhas beenemployed.

So oft in theologic wars,The disputants, I ween,Rail on in utter ignoranceof what each other mean,And prate about an ElephantNot one of them has seen!John Godfrey Saxe (1865),The blind men and the elephant

IntroductionThe six blind men in the above extract from the well-known Hindu fable inspecteddifferent 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 appreciatethe limitations of their own viewpoints, and disregard of their companions' observations.To avoid the distorted conclusions of the blind men, scientists must comprehend therestrictions of their perspectives and the relative perspectives of the sciences.In response to the explosion of knowledge and the search for more, biology has becomeextensively subdivided into areas of specialization. Disciplines and sub-disciplines can berelated in the same ways that formal taxonomic groups are related (that is, they can haverelative 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 providea clear systematization and clear nomenclature for their own field and its components.It has been argued, with appreciable justification, that "vague" terminology may beadvantageous in science (Rosenberg, 1975): disciplinary titles are often mental straightjack-1 Biosystematics Research Centre, Agriculture Canada, Research Branch, CentralExperimental Farm,Ottawa,Ont. KIA OC6,Canada.

335


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336 TAXON VOLUME 38

ets, limiting investigators to the theoretical foundation, if not indeed models, methodologyand 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 beenif he had remained disciplinarily restricted to taxonomic malacology. In attempting tounderstand the perspective and goals of a science - especially by defining it - there is dangerof placing that science in a straightjacket.Nevertheless, as the taxonomist is well qualified to appreciate, a comprehensible ter-minology for scientific fields is desirable simply for clear communication. As will be re-viewed, 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 inresponse to what has been considered new and fashionable; and have mutated over theyears, sometimes to the point of extreme ambiguity. In short, what is needed is a systematicexamination of biological systematics, and a nomenclatural analysis and consequent syn-onymization of terms denoting systematically-oriented disciplines. Many of the semanticdifficulties treated here could have been avoided if the well-known and easily comprehendedterm "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 nounimplies a classification" (cf. various meanings of classification in, for example, Jardine(1969)).This paper turns the methodology of biological systematics upon itself. Specialized fieldswithin systematics in its broadest sense are treated analogously to populations, and areconsidered as posing at least some similar phenomena of evolution (mutability, combin-ability, adaptive "fitness" to scientific and political needs) and problems of classification(specification of circumscriptional criteria, whether hierarchical or overlapping). I hopethat the analogy is provocatively heuristic, but it has its limitations. In particular, thereare no codified conventions for clarification of nomenclature for sciences and subsciencesas there are for groups of organisms. Also, unlike groups of organisms, concepts of fieldsof science may be inherently subjective, and are often rather vague and flexible. It will benoted that I have reversed the normal order of taxonomic examination, by beginning withestablished names of scientific fields, and then searching for discontinuities between thedesignated concepts, rather than vice-versa. It would be simpler, if of academic interestonly, to circumscribe scientific fields anew, and subsequently seek out the most applicablenames.

There is ample literature on what taxonomists and systematists do, and how and whythey do it, which of course provide the bases for circumscribing these fields. It is particularlyimportant to specify purpose (Gilmour and Turrill, 1941). McNeill (1979) concluded: "atleast above the species level, the purpose of classification in biology is to provide a simplifiedexpression of the interrelationships among organisms". The more general systematic ob-jective is simply to clarify relationships among organisms. "Relationship" may be phe-notypic or genotypic (or both), a grade or a clade (or both), combinatorial (simple hybrid-ization, 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 contextof evolutionary theory (Anderson, 1974). Another principal consideration is human psy-chology, and here there are several constraints, perhaps the most important of which isthe visual Gestalt. As Heywood (1973) noted, "because of man's visual sense, the biologicalclassifications 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 processescan begin". (Compare Stafleu's (1969) discussion of "non-visual systematics".) Anotherfactor 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, indeeddesirability of artistic expression in systematizations. Cowan (1955) remarked "the 'best'classifications are those made by artists with the keenest appreciation of what is both usefuland intellectually satisfying".


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AUGUST 1989 337

Relationshipof ClassificatoryBiologicalScienceand OtherSciencesAny field of research implies restriction to a discipline or subdiscipline, with its associatedmethodology and accumulated data. In the classical scheme of Tschulok (1910; see analysisin Major (1958)), eight quite separate "viewpoints" were applied to biology, namely com-position, structure, physiology, genetics, ecology, history, chorology and taxonomy. Theresulting sciences were alleged to be independent. However, when one considers the re-lationships between major fields, such as biology, geology, and chemistry, it becomesapparent that there can be considerable overlap. Biogenetic pathways may be as importantto 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 in-creasingly problematical (Garfield, 1986; Maddox, 1985). Clearly, there is so much overlapbetween fields of science as presently conceived that their circumscription (horizontalcategorization) is often difficult. Without belabouring the point, it is clear that sciences andsubsciences are subjective, arbitrary, human constructs, susceptible of different kinds ofseparation.A second problem is that of hierarchical recognition (vertical categorization), and in thisregardthe organization of university departments is illuminating. Gardiner (1988) remarkedthat 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 themost recent edition (Custer, 1979a): "classification may be applied not only to tangibleobjects 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 likenessesas 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 identificationof organisms) is done in key-like fashion by considering the set of alternatives at anydecimal 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 todifferentclasses by different librarians. This is of course different from identifying organismson the basis of their characteristics, where the discriminatory value of an attribute shouldnot arbitrarily vary with the subjective opinion of the observer.The Dewey system is an example of hierarchical organization of mutually exclusivegroups, and for the purpose of systematizing biological science, is both advantageous anddisadvantageous. It is good in being a stable, widely used, and at least somewhat reasonablesystem (perhaps like the Englerian system on which many herbaria are organized). It isbad in having been erected subjectively and somewhat arbitrarily, in lacking the flexibilityof permitting the incorporation of better organization as it is conceived, and indeed inproviding a ratherrigid (rathertypological) way of consideration. (Only minor modificationsof the Dewey system have been made since its inception in 1873; the Library of CongressClassification system is more flexible.)A "logical grouping or classification of the zoological sciences and in particular of thesystematics fields" was attempted by Blackwelder and Boyden (1952), whose hierarchicalresult 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 concludedthat insofar as systematics is concerned, the many contributing sciences are completelyinterdependent, producing one comprehensive class.Some ways of interpreting the relationships of systematics and some principal relatedfields are shown in Figs. 1-4. The hierarchical arrangement of Fig. I is simplistic, butemphasizes well the highly varied data base of systematics. Figure 2, in which systematicsis viewed as a central or core discipline receiving data from many other sciences, is probablythe most attractive of the four figures to systematists themselves. Figure 3 interprets sys-


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338 TAXON VOLUME8Table 1. Extractof DeweyDecimal classification orbiologicalsubjects.

560 Paleontology Paleozoology561 Paleobotany562-569 Specificanimalsandgroupsof animals570 Life sciences571 (Unassigned)573 Physicalanthropology574 Biology575 Organicevolution andgenetics576 Microbes577 Generalnatureof life578 Microscopy n biology579 Collectionandpreservationof biologicalspecimens580 Botanical ciences581 Botany581.1 Physiology of plants (581.11 = genetics; 581.19 = biophysics and biochemistry)581.2 Pathologyof plants581.3 Developmentand maturationof plants581.4 Anatomyand morphologyof plants581.5 Ecologyof plants581.6 Economicbotany581.7 (Not assigned)581.8 Tissue,cellular,molecularbotany581.9 Geographicalreatmentof plants

tematics as one of many overlapping sciences of equal status, and probably is nearer tothe viewpoint of most biologists than the preceding model. Alternatively, this could beillustrated by a completely joined network. Figure 4 illustrates three axes of a hyperspacein which each interacting discipline contributes a dimension (see Sattler (1987) for ananalogous concept from the point of view of morphogenesis).Theoretically, one could conceptualize a systematic subdiscipline for every approach tobiology: morphological (-anatomical), evolutionary (perhaps including paleobiological),genetical, chemical (-molecular), and mathematical systematics seem reasonable in termsof principal current approaches. Ecological (see Duke's (1977) discussion of "ecosys-tematics"), economic, geographical, physiological, physical, geological (i.e., stratigraphic-fossil), psychological (i.e., ethological), and pathological (see Savile's (1979) use of suscep-tibility to fungi as an indicator of relationship) systematics are other activities of biologicalsystematics involving separately recognized fields, although the hybrid disciplines are notusually conceived of as meriting subdisciplinary recognition. Although it may seem far-fetched, there are even possibilities of hybrid fields between biological systematics andareasof inquiry usually conceived of as not belonging to the realm of science. Thus theologycould be joined with biological systematics (as it indeed once was) to clarify the naturalorganization of living things by God(s); and art intermingles with biological systematics inproducing aesthetically appealing systematizations.Biological classification clearly is most related to the companion sciences evolution andecology, which are also synthetic, holistic, genetically-based, and organismic-populational.In practice, these fields are so complementary that one cannot avoid characterizing themas overlapping. Kruckeberg (1969b) noted "it is plant taxonomists, not taxa, that makedistinctions 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) heldthat "for its basic data ecology-again like taxonomy-is almost wholly dependent uponother disciplines, and the attempt by some to maintain that ecology is quasi-independent


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AUGUST 989 339Table 2. Blackwelder nd Boyden's(1952) classificationof the zoologicalsciences.A. Dealingwith one or a veryfew kinds

Dealingwith the speciesHomo sapiensMedicinePsychologyPhysicalanthropologyDealingwith domesticand a few other animalsVeterinarymedicineApicultureAnimal breedingSubjectswith a predominantlymedicalaspectPhysiological mbryologyHistologyCytology

B. Dealingwith kinds as suchDeal principallywith determininghe differencesand similaritiesbetweenkindsComparativeanatomyComparativephysiologyAnatomicalembryologyComparativehistologyCytologyZoogeographyGeologicaldistributionAssemble the data into orderand make them available foruseTaxonomyClassification

Draw conclusionsor generalizations f broadsignificancerom the classifieddataPhylogenyEvolutionInvestigate he causesof the observed facts and thus contribute o theirrecognitionMorphologyStratigraphyGeneticsC. Dealingwith kindsin relationto theirenvironmentEcology =bionomics= naturalhistory)

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 geneticorganization among organisms, and clarifying adaptive environmental relationships, mayrespectively distinguish evolution, taxonomy, and autecology. Munroe (1964) wrote "sys-tematics 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 plantswhich overlaps the viewpoints of other sciences. Considered in these perspectives, system-atics may be characterized as an approach to or viewpoint of biology stressing geneticorganization from the level of the individual upward. Similarly, evolution is an approachstressing (populational) change, and autecology is an approach stressing adaptation (cf.Fig. 4).Egocentricity of "Biological Systematics"; Systematics sensu latissimoChauvinism is common in science, and some semantic distinctions obscure the essentialnature of biological classification. The position has been taken that classificatory scienceis more fundamental or elementary than all other biological sciences, because all othersproceed from a taxonomic base (Simpson, 1961). This is of course true, but other sciences


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340 TAXON VOLUME8SYSTEMATICS \LOG\ CAL SC/

SEVOLUTION ECOLOGYGENETICS ECOLOGY MORPHOLOGYEVOLUTION BIOGEOGRAPHY BIOCHEMISTRY

GENETICSBIO GEOGRAPHY

MORPHOLOGY BIOCHEMISTRYSYSTEMATICS ECOLOGY

ECOLOGYEVOLUTION

EVOLUTION BIOGEOGRAPHYEVOLUTION

GENETICS BIOCHEMISTRY

MORPHOLOGY

SYSTEMATICS

Figs. 1-4. Models of relationshipsof some biologicalsciences.(Scienceschosen are exemplary,not exhaustiveof the possibilities.)1. Systematicsas the apexof a hierarchy; . Systematicsas a corescienceto which all otherscontribute; . Systematicsasone of many overlapping ciences;4. System-atics as one of manyscientificdimensions or 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 scienceshave their indispensable values. Another overstated argument for the uniqueness of sys-tematics is that it is the most inclusive of all biological sciences, synthesizing data fromall other disciplines while generating none of its own (Simpson, 1961). The same claimscan be made for ecology and evolution, and in any event many biological data are not ofinterest to systematics. Furthermore, systematists do collect their own data as evidencedby specialization in fields that have proven especially useful in providing data, such asmorphology and chemistry. Simpson (1961, p. 5) stated that taxonomy has "almost asuperscientific 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 involvesordering, so that accordingly systematics and science are coextensive. Hennig (1950) stressedthis idea, which seems to recur independently; Ayala (1968) stated "the goal of science isthe systematic organization of knowledge about the universe". Indeed, science may beconsidered to be a "system". As pointed out by Rowe (1961), "since at least the time ofCompte, the idea of arrangingand classifying the sciences in a hierarchy according to levelof 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 thecomprehensivenss of the units studied, and notes systematic products of each (cf. the highlyimaginative 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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AUGUST 989 341Table3. An analysisof disciplineson the basisof unitsstudied and systematizations chieved.

Unit studied Field of study Exampleof systematizationEnergy, ubatomicparticles Physics Classification f elementaryparticlesAtoms, molecules Chemistry Periodictable of elementsTissues,organs Histology, anatomy Tissue,organclassificationsSingleorganisms Morphology s. str.) GrowthformsystemsGroupsof geneticallyrelatedor- Taxonomy TaxonomicsystemsganismsGroupsof habitat-related rgan- Phytosociology Phytosociological ystemsismsGroupsof geographicallyelated Zoogeography, hy- Zoogeographic rovinces,phytogeo-organismsandorganism-related togeography graphicregionsregionsEcosystem Synecology Ecosystematic ystems

species"). As is clear from Table 3, systematization of biochemicals, biological organs, andbiotas could legitimately be called "biological systematics", although in common with mostcurrent usage, the phrase is used in this paper for the systematics of groups of wholeorganisms, associated on the criterion of genetic relationship. "Organismic systematics" ismore accurate than biological systematics, but could tIeoretically refer to classification oforganisms on grounds other than the purely genetic, and for example could include theclassification of biotas.In General system theory, von Bertalanffy (1968) presented an expansive "systematic"perspective of science. He sought natural "systems" (defined as complexes of interactingelements) 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. Bertalanffywas opposed to the reductionist approach of seeking explanation of organization simplyon the basis of the most elementary (physical) constituents. Griffiths (1974) wrote in regardto Bertalanffy's approach, "I believe that his work is important for understanding physicalsystems, but am not convinced of the validity of generalizing the system concept in thisway". It may be that von Bertalanffy has overstated the parallelism of organization ad-dressed by different sciences. Griffiths (1974) pointed out difficulties with the delimitationof populations as systems sensu Bertalanffy (and hence of so-called "biosystematics"); andsimilarly difficulties with the delimitation of ecosystems (and hence of so-called "ecosys-tematics"). It is intriguing, if premature, to speculate that there is a common systematicstructure and nature to all of reality that transcends elementary physical constitution.Frequency of Use of the Principal Titles: Systematics, Taxonomy, BiosystematicsAs remarked by Mason (1950) in his examination of these three terms, "it is usage andthe history of usage that ultimately moulds the meanings of our words and terms". Toclarify the comparative frequency of usage, several surveys were conducted, as describedbelow and tabulated in Table 4.

1. The frequency of use of the terms was compiled, manually from Biological Abstractsfrom 1943 (when "biosystematy" was coined) to 1968, and by computer from 1968 to andincluding 1986 employing the Biosis service (Biosis, 1987). In Table 3, the title taxonomyin fact is based on taxa and *taxon*, where * represents any addition; thus cytotaxonomy,chemotaxonomy, taxonomic, taxonomical and taxonomist are incorporated in the total of42,871. As a matter of interest, this included 1034 cytotaxonom* and 2189 chemotaxo-nom*. The term systematics is based on systematic* (systematic and systematics occurredin about equal frequency), and similarly biosystematics was based on biosystematic*. Of


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342 TAXON VOLUME8Table4. Frequencyof use of the majorheadingsof biologicalclassification cience.

Biosys-tematics Systematics Taxonomy1. Use in ca. 12,000,000 titles of publications,1943-1986 283 10,064 42,8712. Use in 1746 titlesof the BiosystematicsRe-searchCentre,1973-1986 16 16 763. Use in ca. 17,500names of journals (basedonBiosis, 1987) 0 22 114. Use in names of ca. 2600 institutesandde-

partmentswith systematiccollections(afterHolmgrenet al., 1981;Arnettet al., 1986) 1 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, theBiosystematics Research Centre ("Institute" until 1986) was surveyed from its inceptionin 1973 up to and including 1986. The ratio of use of biosystematics : systematics :taxonomyis about 1:1:5. It seems that being in a "biosystematics institute" does stimulate one touse 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, basedon 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 termfor journals than taxonomy, likely because of the prestige and greater generality attachedto 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 herbariaof the world), and from Arnett et al. (1986; listing locations of collections of insects andspiders). The only "biosystematics" institute was the one I represent (the Biosystematicsand Beneficial Insects Institute of Beltsville, Maryland came into being in 1986, and wasdisbanded in 1988). There were 11 taxonomic institutes and 33 systematic institutes, onceagain systematics apparently having a more desirable connotation than taxonomy, althoughthe 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 authoritativetexts. In the following I have limited the discussion to what I consider to be the pivotalconsiderations and key illustrative literature. The various ways of interpreting the rela-tionships 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 laTaxonomie (2), ou de la Theorie des classifications appliquee au regne veg6tal"; and infootnote (2): "Mot form6 de ra4Ls ordre, et votos loi, regle". Mason (1950) concluded thatin the context of de Candolle's book, classical taxonomy was concerned with relationships(primarily morphological), classification, hierarchical categorization, nomenclature, anddescription. Mason was unable to find a precise derivation for systematic, but noted thatits use in biology as referringto a hierarchical arrangement of organisms may have predatedthe 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 synony-mously, but that the latter term was often used to indicate comparative biological study,


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AUGUST 989 343TAXONOMY SYSTEMATICS

TAXONOMY= SYSTEMATICS

5 6

TAXONOMY SYSTEMATICS

SYSTEM- TAXONOMYATICS

7 8

TAXONOMY SYSTEMATICS

9Figs. 5-9. Conceptionsof relationshipbetweentaxonomyandsystematics.5. Equivalent; . Mu-tually exclusive; 7. Systematicsas 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 rel-atively narrow definition of systematic botany centred on approach: "we may define Sys-tematic Botany as the comparative study of any related (systematic) group of plants utilizingthe 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 usuallyconsidered the end of classificatory science (i.e., the system). In a similar vein, Davis andHeywood (1963, p. 452) wrote "much of the debate, in fact, centres around whether it isthe aim of taxonomy to study products or processes". Of interest, Mason's implicit defi-nition of systematics (a word not used in his 1950 article), which does not reflect commonusage, may be construed as excluding taxonomy, and vice-versa (see Fig. 6).Simpson (1961), who also noted that taxonomy and systematics were commonly usedsynonymously (as he did in his earlier writings), proposed a distinction by which systematicsis broader than taxonomy, including it (see Fig. 8; he also included classification, andnomenclature, which as noted in the following he excluded from taxonomy). Simpsonstated "systematics is the scientific study of the kinds and diversity of organisms and ofany and all relationships among them". This definition has been parroted in numeroustexts, usually without appreciation, in my opinion, of what he went on to state: "in thisdefinition 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, includingspecifically all associations of contiguity.. .". "Relationship" in the context of classificatoryscience is normally assumed exclusively to designate genetic ("phylogenetic", "evolution-ary", "patristic", etc.) affinity; Simpson made clear, however, that he included biogeo-graphical and ecological relationships. Since these are not heritable, they do not bear directlyon what taxonomists-systematists have conceived of as the core of their domain. Homo-plastic relationship (whether due to convergence or parallel evolution), chronistic relation-ship, spatial relationship, ecological relationship, and still other kinds of relationship mayserve to clarify heritable relationships, and therefore may be of "systematic" relevance,but their inclusion in a definition of systematics is challengeable. Most systematists wouldprobably 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 abranch of systematics, since the study of community relationships of organisms addressesa conceptually different kind of organization than systematics addresses.

Simpson's comprehensive definition of systematics leaves the impression that any kindof relationship is appropriately studied under the heading of systematics. Similarly, theadjective "systematic" has been so frequently applied as in "systematic anatomy" that itmight seem that any kind of comparative study is appropriately termed systematic (noteMason, 1950). However, Simpson (1961, p. 9) held that comparative autecology is essentialfor some aspects of systematics, but he nevertheless specifically excluded autecology fromsystematics. The position is taken here that the ultimate aim of systematists is the clari-fication strictly of genetic aspects of relationships, and thus it might seem that one isproposing 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 sciencesand subsciences have hindered rather than promoted communication. While it is partic-ularly appropriate that systematists inquire on what, fundamentally, distinguishes system-atics (self-analysis is sometimes useful), it would be regrettable if this resulted in constraintson either research or outlook. It is not easy to find boundaries between the sciences, anymore than circumscribing a city from its suburbs, satellites, and sister cities. Often indi-viduals working in a city, and making valuable contributions to it, are not technicallyresidents of that city, but of another. So long as the boundaries are permeable, they arenot 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 fourmain types of relationships between plants, based on: descent (phylogeny), similarity (phe-netics), space (geography), and trophic relations; he stated that systematists deal with allfour, although emphasizing phylogenetic and phenetic relationships. In my view it is mis-


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AUGUST 1989 345

leading to assign the latter two kinds of relationship to systematics, although admittedlysuch may bear indirectly on interpreting genetic affinity. Benson (1943, p. 99) expressedthe idea well: "the real goal of systematic botany isorganization--arrangement

of themyriads of living plants into a readily understandable system made up according to theirgenetic or 'blood' relationships".For taxonomy, Simpson echoed a distinction of Gregg (1954) who in effect distinguishedtheory 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 the-ory-concerned definition of taxonomy (note later comments on theory vs. practice). Mayr(1982, p. 146) made a similar distinction, dividing taxonomy into microtaxonomy, dealingwith methods and principles of recognizing and delimiting kinds (species), and macrotax-onomy, by which kinds are arranged as a classification.

Simpson's resolution of the meanings of the terms systematics and taxonomy has beenwidely 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 worriedabout this subject will too. That is, what is wrong with Simpson's classical definitions ofsystematics and taxonomy? I have always regarded them as the best statements of thedifferences between the terms, and a large number of later authors have simply modifiedthem. One of them deals with relationships--any and all relationships--and the other withclassification." Above, the viewpoint is taken that including all relationships in the defi-nition of systematics is not desirable because it includes a breadth of non-heritable variationthat dilutes the focus of classificatory science. Further, Simpson himself arbitrarilyexcludedthe study of some relationships. For example, it is instructive to note that he substantiallyexcluded 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 diversityof 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 allocateclassification under the rubric taxonomy. The problem, as pointed out in the preceding, isthat Simpson did not fully do this; he assigned only theoretical aspects of classification totaxonomy, and in segregating the practical aspects of classification to systematics, he re-moved from taxonomy what most taxonomists understand to be included.The attempt to distinguish taxonomy and systematics on the basis of theory vs. practicehas been widespread. However, as should be clear from the following, different writershave had contradictory ideas of the way the distinction should be be made, and indeed ofjust what is theory and what is practice. Camp (1951) specifically distinguished taxonomyas theory, systematics as practice: "we should keep clearly in mind that as originally definedby A. P. de Candolle (1813), the term 'taxonomy' must be retained for the study of thelaws and principles underlying a system of classification. Systematics ... refers only to theclassification 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 theclassification of organisms", and systematics as "the practice of describing, naming andclassifying 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 somewhatlike that of theoretical physics to the whole of physics" (i.e., taxonomy is a theoreticalcomponent of systematics; Fig. 8). Similarly, Heywood (1973), who accepted Simpson'sdefinition of systematics ("the scientific study of the diversity and differentiation of organ-isms 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 inrecent years to restrict it to its original meaning, namely that part of systematics whichdeals with the study of classification, its bases, principles, procedures and rules. It thereforecovers ... 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 ofpractical aspects of classification is clearly inconsistent with Simpson's narrow circum-scription of taxonomy.) Blackwelder (1967) used taxonomy to refer to "the day-to-daypractice of dealing with the kinds of organisms", and systematics to "the study of the kindsand diversity of organisms, their distinction, classification, and evolution" (Fig. 9).In most texts treating classificatory science (perhaps more evidently in North Americathan elsewhere), systematics is equated with taxonomy (e.g., Ross, 1974; Jones and Luch-singer, 1979; Radford et al., 1986; Fig. 5). In practice when a distinction is made, systematicsis usually considered to be more inclusive than taxonomy. This is illustrated by Solbrig'sstatement (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 isweak. Ornduff (1969) noted that "many of us would rather be called systematists thantaxonomists, since in some circles the latter carries with it connotation of anachronism"."Biosystematics": Ambiguous, Pretentious, Superfluous, But UsefulThe publication of a collection of papers as The New Systematics (Huxley, 1940) pre-cipitated problems concerning how biological classificatory science should be conceivedand labeled. Although the general tone of the book was highly respectful of traditionaltaxonomy, and not suggestive of a need for new fields for taxonomy, nevertheless this wasthe 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" (crit-icized by Blackwelder (1967, p. 342)). The climate of the time was marked by excitementthat the "new", "experimental" techniques of chromosomal, breeding, and populationalanalysis were rejuvenating what many conceived of as the rather unscientific, exceedinglydull 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 NewerSystematics, Experimental Taxonomy, or Biosystematics". Blackwelder (1967, p. 4) ap-propriately criticized the situation: "During the past several decades some discussions oftaxonomy have used terms that may be either derogatory in tone, ostentatious in impli-cation, or pedantic in application. Among these may be cited ... The New Systematics ...biosystematics". Stafleu (1969) observed that "the new systematics" was "a somewhatoverly optimistic term, since nothing ages as rapidly as the new".

Camp and Gilly (1943) proposed the field ofbiosystematy: "As here defined, Biosystematyseeks (1) to delimit the natural biotic units and (2) to apply to these units a system ofnomenclature adequate to the task of conveying precise information regardingtheir definedlimits, relationships, variability, and dynamic structure". At the time, they entertained thethoughts, naive in retrospect, that "natural units" based on mode of reproduction existedbelow the species level, and a nomenclature for different kinds of breeding situations wouldbe useful. None of an orgy of words they coined for their concepts was taken up exceptbiosystematy, which soon mutated to biosystematics (the earliest use of the word I haveencountered is Epling (1943)). Vickery (1984) pointed out that "biotaxonomy" would havebeen 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 thecoining of biosystematy (Moore, 1984), and the word became associated with botanicalcytogenetic studies. Constance (1964) observed that "the cytogenetic period of taxonomyhas 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 fieldsother than cytogenetics were coming to the forefront of systematic research, biosystematicsas a term mutated to become intolerably ambiguous, as noted in the following.


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Jones (1984) wrote: "Both Camp and BiScherhave protested at the perversion of theoriginal definition of "Biosystematy" (Camp, 1961; BScher, 1970) and I find myself verymuch in agreement with them"; and with regard to the derivative biosystematics, "it mightseem that there is today some universal agreement as to its meaning, but I doubt it". InBiosystematic literature, Solbrig and Gadella (1970) tellingly comment "Compilers in manycountries have worked hard at compiling this index. However, their ... definition ofBiosystematics ... varied". In Plant Taxonomy and biosystematics, Stace (1980) quicklyrevealed (p. v) the lack of conviction of some who use the term biosystematics: "despiteits title, the subject matter of this book is taxonomy". Monroe (1977), in a summary of asymposium on biosystematics, stated: "Some seemed to think of biosystematics as beingan 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 whatwas formerly called 'the new systematics,' that is, the whole array of indirect, theoretical,experimental, and modem methods as opposed to 'conventional' systematics". In theauthoritative Principles ofangiosperm taxonomy, Davis and Heywood (1963) stated "muchof what is included under the general title of biosystematics is simply part of the modernapproach to taxonomy and represents usually an extended state of knowledge of taxa. Theterm biosystematics itself is in fact largely superfluous and serves mainly to describe oneof the stages through which taxonomy passes". Johnson (1970b) 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 inso 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, infavor of 'plant population biology' and various other more specific alternatives".A growing number of authors has arbitrarilychosen to define biosystematics in a personalway, or to make biosystematics essentially coextensive with taxonomy and/or systematics.Stafleu (1969) observed the use of biosystematics as "evolutionary systematics of the livingorganisms 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 ofthe 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 syn-onymous with systematics, although with an emphasis on the study of genetical and cy-tological phenomena", offered the following personal definition: "biosystematics can nowbe defined more precisely as the application of genetics (and cytogenetics), statistics, andchemistry to the solution of systematic questions in order to provide explanations aboutthe 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 tobe that part of it which fell within the sphere of biology". Ross (1974) stated "I am usingbiosystematics here as the investigational field of systematics based on any scientific in-formation that can be brought to bear on the problems of the evolution of species, whetherthey concern speciation or phylogeny". Stace (1980, p. 6) wrote: "it is unfortunate that theterm biosystematics has been widened by some taxonomists to cover virtually any taxo-nomic 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 + EvolutionTuresson (1923) stressed ecology in coining the term genecology: "It seems appropriatefor several reasons to denote this study of species-ecology by the term genecology (fromthe Greek 'genos', race, and 'ecology') as distinct from the ecology of the individual or-ganism, 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. Hes-lop-Harrison (1964) interpreted genecology in the original Turessonian sense as a "syntheticdiscipline", merging ideas and methods from taxonomy, genetics, ecology, and physiology.Bennett (1964) stressed the evolutionary aspect of genecology in her redefinition of it: "thestudy of the genetic mechanisms which operate within organisms, and between organismsand their environment, at the level both of the individual and of the population viewedas a process, which at the population level produce those changes known collectively asmicro-evolution." She recorded her view that the terms taxonomy, experimental mor-phology, new morphology, new systematics, genonomy, biosystematics, and neotaxonomyare not equatable with genecology, and the terms experimental evolution and micro-evo-lution are more restricted in scope than genecology. (Epling's (1943) ill-defined term ge-nonomy, which was proposed to designate studies that then also fell under the terms "thenew systematics", population genetics, and biosystematics, never received acceptance.)Lawrence (1951, p. 169) stated about genecology, "this term has not been accepted inAmerica where it is confused euphonically with the medical term gynecology".A current dictionary definition of genecology (Gove, 1981) has a taxonomic aspect: "abranch of ecology concerned primarily with the species and its genetically variant subdi-visions, 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 andecological variation of natural populations was independent of taxonomy. Langlet (197 1,p. 659) implied that while genecology could help taxonomy, unless the two were maintainedas distinct fields, the term genecology ought to be abandoned. Nevertheless, as pointed outby Constance (1953), "genecology slipped away from the realm of plant ecology to that ofplant taxonomy". Heslop-Harrison (1964, p. 237) did not quite concede this transfer: "anunfortunate trend during the post-war period has been apparent in recurrent attempts toassimilate genecology into taxonomy".So-called experimental classifications began with Turesson in 1922, and were based ontransplantation, breeding, and analysis of environmental adaptations (for details see, forexample, Baker (1952); Davis and Heywood (1963); Stace (1980)). It has been clear for atleast two decades that genecological classification is a sterile exercise from the point ofview of classification (e.g., Constance, 1957; Merxmiiller, 1963), whether based on arbitrarydegrees of capacity to interbreed, adaptive differentiation, or a combination of these.

Mayr (1982, p. 277) and Heslop-Harrison (1964, p. 237) noted the common interpre-tation of biosystematics as genecology. Stace (1980, p. 6) wrote: "biosystematics may ...be considered as the taxonomic application of the discipline known as genecology-thestudy of the genotypic and phenotypic variation of species in relation to the environmentsin which they occur". However, Heslop-Harrison (1964, p. 238) insisted that "biosys-tematics (or experimental taxonomy) should ... be preserved as something distinct fromgenecology ... ecological data and observations on genetic systems are a sine qua non ofgenecology, although by no means an essential part of taxonomy. Conversely, the nomen-clatural and bibliographic studies which are an obligatory part of any taxonomic study arenot necessarily significant for a genecological investigation of a species".Kruckeberg (1969a) stated "genecology constitutes, to date, the greatest point of contactbetween ecology and plant systematics", and "we may look upon genecology as a specialbranch of microevolutionary study". Clearly genecology is a mixture of systematics, ecology,and evolution, and insofar as it addresses "racial systematics" (cf. the following discussionof 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 fre-quencies in (within, among) populations, is an ambiguous term, on the one hand reflectinga tradition of research on mathematically-based, theoretical, statistical populations, and


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on the other hand researchon organisms.In the latter sense, the origin of populationgeneticshasbeen attributed o Chetverikov n 1926 (Adams, 1968), althoughMayr(1982,p. 553) acknowledges he contributionsof others.Mayr(1982, pp. 559-560) interpretedChetverikov's ype of population geneticsas essentiallya combination of systematics (asit applied o geographic aces)andgenetics.Camp(196 1)noted that someequatepopulationgeneticswithexperimental ystematics,and he himselfequatedmuchof it to his ownterm,biosystematics.Ford (1964) usurpedthe field for ecology as ecological genetics. Langlet(1971, p. 659) observedthat such usage essentiallyequates populationgeneticsand gen-ecology.Genes arehereditarydeterminants,andby definitionthe subjectof the fieldof genetics.Changein gene frequenciesclearly belongs to the field of evolution, and so one wouldexpecta close relationshipwith systematics.However,if one confines the subjectof pop-ulationgeneticsto the genotype,then the relationshipof it and systematicssubtly hingeson causeandeffect,since even those whoadvocateclassification n the basis of genealogicalrelationshipsn fact do so almost entirelyon the basis of the phenotype(afterdiscountingenvironmentalmodifications).The visible associateof populationalgenetics,populationalphenetics, s squarelywithin the realm of Mayr'sterm(1982, p. 247) "populationsystem-atics".Inpractice,populationgeneticsas a termis applied ikegenecology, o intra-specificstudies in which morphologicaldifferentiations usuallytoo limited for formal classifi-cation.And,likegenecology,t isgenerallynotconceived aspreciselydesignatinghe residueaftertaxonomyis deductedfrom systematics."ExperimentalTaxonomy":Deceptiveand UnnecessaryA common interpretation f biosystematics s that it is "experimentalaxonomy" (e.g.,Heslop-Harrison,1954; Boivin, 1960; Stace, 1980). As noted earlier, some have alsoassignedgenecologyto synonymy.Constance(1953) traced this phraseback to 1919 (cf.Langlet 1971, p. 658) who tracedthe term to 1920, and Heslop-Harrison 1964, p. 237)who traced t to 1934).Not only is the meaningof the phraseunclear,but also it has neverbeen obvious in exactly what respectsexperimental axonomy is experimental.It seemsthat the presenceof laboratory,expensive and complicatedapparatus,and novelty, con-comitants of muchgenuineexperimentation,have often been sufficient o label an activityas experimental e.g.,Munroe,1960).Constance 1951)stated"Ifailto see howtaxonomy,itself, can ever be regardedas experimental,unlessone assumes that the successor failureof compatibilityor biochemicaltests, for example,areper se proofsthat two individualsdo or do not belongto the the same taxonomiccategory".Presumably t is this rationalethat validates the so-calledexperimentalclassificationsmentionedin the preceding.The concernwith the "experimental" spectof taxonomywasobviouslypartof the 20thcenturyattemptto transform axonomyinto a "science",since experimentations widely(and incorrectly)consideredto be a sine qua non scientificprocedure.Lewis (1957) andMayr(1982, pp. 30-32, 521, 855-856) noted that controlled aboratoryexperimentationis just not that differentfrom intelligentobservation and interpretationof uncontrollednaturalphenomena,especiallywhen done from the comparative perspectiveof the tax-onomist. Indeed, even the so-called experimentaltaxonomist relies overwhelminglyonobservationalevidence, usually only crudelycontrolledmanipulationof a very limitedportionof the genotypeand phenotype being possible.Taxonomic and systematicinter-pretationare primarilyof observationaldata, and the use of the admittedlyatt:ractivephrase "experimental axonomy"is usuallysuperfluous.Taxonomy,TaxaBycomparisonwithde Candolle's 1813)coiningof the wordtaxonomy,the 20thcenturyterm taxon is a latecomer.Accordingto 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 almostuniversally accepted among taxonomists in this one sense. (Unfortunately, the supplementto Webster's third international dictionary (Mish et al., 1986, p. 189), presents the followingerroneous definition: "Taxon abbr taxonomic; taxonomy"). I believe that most moderntaxonomists share Moss's (1983) viewpoint, "a taxonomist's basic concern is taxa, andhow to group and explain them". In contrasting taxonomy and systematics, it is etymo-logically desirable to establish some consistency between the accepted formal unit of tax-onomy, the taxon, and the science of taxonomy, and this is achievable by stressing, as inthe following, the essentially formal classificatory nature of taxonomy.Beyond Taxonomy: Unconventional Systematization

Formal, codified botanical nomenclature reflects the classical (Aristotelean) viewpointof classification that things not merely should be, but are hierarchically ordered into ab-solutely discrete groups on the basis of attributes (Cain, 1959). This has proven to applywell to organisms, but there is a recurrentdifficulty. Membership often proves problematicalwhen groups are polythetically defined (i.e., "fully polythetically": group membership notbeing determined by invariable possession of any one character, but by preponderantpossession 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 othercharacters suggest its membership in another covering group, and there is no really satis-factory way of making the choice. Formal taxonomy cleanly cuts the Gordian Knot ofincongruent natural variation with a Procrustean solution: everything is forced into auniquely named pigeon hole in a hierarchically stratifiedpigeon coop. There is a gentleman'sagreement (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 formalschemes, but there is a constant search for the Holy Grail of classical taxonomy, the onetrue system, with its place for everything, and everything in its place. When nature providesvariation patterns that are just too complex to be accommodated by conventional tax-onomy, 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 recognitionof "overlapping taxa" as contrasted with conventional, mutually exclusive taxa. DuPraw(1964, 1965a, 1965b) observed that mathematically representing characters by a reducednumber of multidimensional axes is quasi-taxonomic, and he used the oxymoronic phrase"non-Linnean taxonomy" (as noted by, for example, Johnson (1970a), while an ordinationis an "arrangement", it is not a "classification"; hence it is not a taxonomy). In effect, suchsimplifying mathematical representations are obviously "systematizations", imposing and/or revealing order. They are particularly useful for depicting non-hierarchically structuredand/or overlapping variation, as is commonly found intra-specifically and in reticulaterelationships, and in this respect can be in conflict with the nomenclatural assumptions ofhierarchy and discreteness. Baum (1974) attempted to introduce multidimensional axisdefinition into formal nomenclatural taxonomy (without disturbing the time-honored prin-ciple of discreteness of taxa), but the unenthusiastic reception of this proposal (Stafleu andVoss, 1975) suggests suspicion that any form of numericlatural systematization couldcompromise formal taxonomy.Turrill (1935, 1938) conceived a gradient of taxonomic activity and associated classifi-cations, that he suggested could be labelled sequentially by Greek letters. Alpha classifi-cations are the best that can be done with available information, and one proceeds astaxonomic knowledge increases to the ideal, omega classifications. A parallel scheme ispresented in Mayr, Linsley and Usinger (1953, p. 19) who noted a trend of designating


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characterizationndnamingof speciesasalpha axonomy,placing hespecies n ahierarchyas betataxonomy,andrelatedevolutionarystudiesas gamma taxonomy.It is an oversim-plificationto imply, as do these Greek-labelled tagesof taxonomic clarification, hat alltaxonomicproblemsare solvableby acquiringenoughinformation.As is well recognized,there is a substantialamount of weakly or confusinglystructuredvariation that simplydoes not lend itself to "classification"n the acceptedsense of the word. Indeed, Mayr(1982, p. 145) stated thatgamma taxonomyis not strictlytaxonomy.It is at the pointthat conventionalformaltaxonomyprovesinsufficiento "systematize"heritablevariation thata needhas beenrecognized o designate he additional nterestsofspecialistsstudyingthe organizationof that variation. This need is met by the practiceofdesignating he more comprehensivefield (that also includes taxonomy) as systematics.This is etymologicallysatisfying,since the ultimate aim may be conceived to be a "sys-tematization"of relationships, hat is, an accountingof the natureof heritablevariation,most expresslyin a formal taxonomy, but in some alternativeclassificatoryor simplydescriptivefashionwhen appropriate.PublicRelationsThe technicalliteratureand referenceworks reflect establishedmeaningsfor the fun-damental erms"systematics","taxonomy",and"biosystematics", ndgeneraldictionariesand encyclopedias ndicatethe extent to which such meaningshave achieved acceptancebeyondthe scientificcommunity.These key words are importantnot only for clear com-municationamongspecialists,but also with othersin the scientific raternityandbeyond.Extracts rom standard eferencesarepresented n Appendix1.Taxonomyandsystematicstend to be equated.Some of the arcane(and contradictory)distinctions that have beenmade betweentaxonomyand systematicsare also present n the examplesof Appendix1,andunfortunately robably onfusetheaveragereader."Biosystematics"s relativelyrarelyfound,and whengiven is usuallydefinedin the increasinglyarchaic senses of cytogeneti-cally-basedor experimentally-based ystematics.As noted earlier,taxon is misdefined nthe acclaimedWebster'sdictionary (Mish et al., 1986). All things considered,these keysystematictermsare not well presented o the generalscientificand lay communities.Half a centuryago, therewas a tendencyto view "taxonomy"as effete,old-fashioned,boring,obsolescent,and marginally elevant,exceptas an identification ervice,and evenin those gentlertimes at least part of the solution was seen in the coining of strikingbuzzwordsike "thenewsystematics" nd"biosystematics". incethen,competitionamongthe sciencesfor talentedrecruits,public support,and resourceshas becomeverykeen,andthis hard realityis perilouslyignored.It does not seem to suffice that systematicsis thefoundationof all biological and agricultural cience, and has contributed more to thecollectiveknowledgeand welfare of mankind than any other field of science. "Success nsciencedependsnot only on rationalargumentbut on a mixture of subterfuge,rhetoric,and propaganda" presentedby Broad(1979) as the viewpoint of the controversialphi-losopherP. K. Feyerabend).Accordingly, t is appropriateo examine not just the literal,but also the aestheticandpolitical qualitiesof the titles of classificatory cience. In English,at least, the word tax-onomy has a strikingly arringqualityto those unfamiliarwith it, and is absolutelyunin-formativeto the uninitiated(who are most likelyto conclude that it has somethingto dowithtaxidermy).By contrast,"systematics"s less abrasive,and if still not informativeasto content, is suggestiveof efficientorganizationas opposed to being "unsystematic". thappensthat systematicbiology does not have an exclusive claim to the adjective (e.g.,systematic theology, systematicphilosophy),and so it is natural to specify "biologicalsystematics",and to contrastthis to the melodious "biosystematics"compare he highlysuccessful"biocontrol","biodiversity","biotechnology").t is regrettablehat themeaningof this combination was so poorlyconceived 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 andsystematics 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 bio-systematics has recently been deliberately enlisted to communicate to the public and toother scientists the urgent need to correct the current deplorable and deteriorating supportfor 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 beennamed now seem to be dubiously distinguishable; this dynamism of systematics shouldlead one to be very reluctant to coin special names for the exciting new developments thatare periodically assimilated into our science. The semantic madhouse of subsciences thathas been created by systematists is already embarrassingly full. Much of the nomenclaturalconfusion 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 asbiosystematics. However, the terms taxonomy and systematics are thoroughly establishedand, separately from whatever distinctions seem appropriate for academic circumstances,these words also need to be presented attractively and comprehensibly to the lay publicand scientific community at large. These sciences need to be formulated (and perhapsperiodically reformulated) to emphasize their quintessences while permitting sufficientlyflexible boundaries to adapt to changing circumstances. Although I have objected to theinclusion of non-heritable variation, Simpson's (1961) definition "systematics is the sci-entific study of the kinds and diversity of organisms and of any and all relationships amongthem" is well and concisely formulated. I prefer to characterize systematics as the scienceof organization and pattern of heritable relationships among the kinds and diversity oforganisms. The best contemporary conception of taxonomy is that it is a very substantialbut imprecisely separated part of systematics, that is especially concerned with the pro-duction of formal classifications of living things on the basis of genetic relationships.AcknowledgmentsI thankL. Knutsonfor correspondence n the publicrelationsvalue of the termbiosystematics,and J. Bain, P. M. Catling,A. Cronquist,W. F. Grant, P. G. Holland, J. McNeill, and severalanonymousreviewers or constructivecomments.Some of the criticism has suggested hatthe pre-sentation n partsof the manuscriptmay be controversial.Of course,I alone am responsible or itsfaults.LiteratureCitedAdams,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--Preludeo discussion.Evolution6: 61-68.Baum,B. R. 1974. Article 36 and numericalclassification.Taxon 23: 652-653.Bennett,E. 1964. Historicalperspectivesn genecology.RecordScottishPlantBreedingStation 1964:49-115.Benson,L. 1943. The goal and methods of systematicbotany. Cact.SucculentJ. 15: 99-111.Bertalanffy,L. von. 1968. General ystem theory.GeorgeBraziller,New York.Biosis. 1987. Serial sources for the Biosis data base. Biological Abstracts, Inc., Philadelphia, Penn-

sylvania.Blackwelder,R. E. 1967. Taxonomy.A textandreference ook.J.Wiley&Sons,NewYork,London,Sydney.- 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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356 TAXON VOLUME8Appendix1. Definitions of taxonomy, systematicsand biosystematics n standardreferences notpresent n standard eferencef not presentedhere).General ncyclopediasThe New EncyclopediaBrittanica EncyclopediaBrittanica, nc., 1978)Systematics:"branchof biologyconcernedwiththe diversityof living thingsandtheirgroupingorclassification"

Taxonomy:"the science of biologicalclassification"The EncyclopediaAmericana Mayr,1981)Taxonomy:"Thetheoryandpracticeof classifyingorganisms"The CanadianEncyclopediaSavile, 1985)Biosystematics: the scientific tudyof kinds anddiversityof organismsandof anyand allrelationsbetweenthem"Systematics:"theactualarranging, roupingand namingof organisms"Taxonomy:"covers the bases, principlesand rules of classificationand nomenclature naming),

includingthe hierarchic ramework,and refersnot to a researchdisciplinebut to a supportingmethodology"La GrandeEncyclopedieLibrairieLarousse,1976)Taxinomie:"definieprimitivementparAugustinPyramede Candolle(1813) comme traitantdestheoriesdes classifications, ette sciencese donne en outrepourobjectifsde circonscrire,.. declasser,de definirexplicitementet de denommer .. On emploiesouvent la graphieoriginelle,taxonomie,qui resulted'une erreurd'interpretationtymologique"(Note:Mayr(1966) contends that taxonomy s morecorrectlyderived thantaxinomy.)DictionariesWebster'sThirdNew InternationalDictionary Gove, 1981)Biosystematics: iosystematy;"experimental axonomyesp. as based on cytogenetics"

Systematics: "1: the science of classification ... 2: a system of classification ... taxonomy ... 3:an organizationalcheme or structure.."Taxonomy:"1: the study of the generalprinciplesof scientificclassification: ystematics2: thesystematicdistinguishing,rdering, ndnamingof type groupswithin a subject ield:classificationRandom HouseDictionary SteinandUrdang, 1973)Systematics:"thestudyof systemsor of classification"Taxonomy:"1.the scienceortechniqueofclassification. . the sciencedealingwith theidentification,naming,and classificationof organisms"The New EnglishDictionary "TheOxfordEnglishDictionary";Murray t al., 1919)Systematics:".. . 4. Nat. Hist., etc. Pertaining o, following,or arranged ccording o a systemof

classification; f or pertainingo classification, lassificatory"Taxonomy:"Classification, sp. in relationto its generallaws or principles; hat departmentofscience,or of a particularcienceor subject,which consistsin or relatesto classification"TechnicaldictionariesFeatherly 1954)Biosystematy:"Taxonomicstudiesinvolving cytologyandgenetics"Systematicbotany:"A studyof plantsin their mutualrelationshipsand taxonomicarrangement"Taxonomy:"Thesystematicclassificationof organisms"Jackson 1928)Systematicbotany:"thestudyof plants n their mutualrelationships nd taxonomicarrangement"Taxonomy:"classification"Torre-Bueno1978)System:"an orderof arrangement fter a distinctplan,or method"Systematic:"in definiteorder,or arranged ccording o a system"Systematist:"a student of the systematicaspect of classification n biology, who discriminatesbetweenspeciesandarranges ormsor groupsaccording o theirrelationships ndaffinitieswithothers"

Taxonomy:"the arranging f speciesand groupsthereof into a systemwhich shall exhibit theirrelationshipo eachotherand theirplacesin a naturalclassification"

small, 1989 systematics or taxonomy of taxonomy

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